TRANSACTIONS

OP THE

WISCONSIN ACADEMY

SCIENCES, ARTS AND LETTERS

JU N 2 9 1927

Xli-

N ATU RAP' SPECIES RATIOQUE

MADISON, WISCONSIN

Volume XXI of the ^ Transactions of the Wisconsin

Academy of Sciences, Arts, and Letters is issued under the

editorial supervision of the Secretary.

GHANCEY JUDAY, f

' Secretary,

TRANSACTIONS

OP THE

WISCONSIN ACADEMY

OP

SCIENCES, ARTS AND LETTERS

VOL. XXI.

i.^ JUN 2 9 1927

iT V 7 V 3^ vv

^4I[eNAi. wHS’

MADISON, WISCONSIN

1924

V

'i ‘ ^,.~.„jiii;[V>’AV''j7l. ;■ ■; : - ■ • , ^

r • . . . / li'’ sCi^ y' : ■•

. v'( ■*»'''. "./.' ■• •'.I'.'

"■I V

■A ■■ • ■ ■”

\:a A

We

Ml

smsxjrmAAp'

Th-'A^; '> ■ '•

St'.

W&fl||£-fS

, "' . ^mS7

'• <

A

■/■\ .'.i i-

■"■ 'i

' ' ^ ¥'!?

■ ■ ^ ■ ■ ’.fe’-W.'

A

Am

M •'

mmi

:;|||ppi

KKSa

si ■ v-...v'«f:e.=«.K,

•■■■idDriStv.’ j».

^■'^#^^i;‘,'. «"' ■■■■■■' ' ’ ■• ■■■■ ■ ■' ■

„ ' - , f d

Ar"'y. ' '■'/■•''

'■ ■ *" ■

B ■ J v/;j;|'i?'ii'' ,

t- iW ' A'v ..>'

...PM. ''"■

■'-AmMsmth

CONTENTS

Page

The Unification of South Africa : A Study in British Colonial

Policy. Paul Knaplund . 1

The Removal of the Winnebago. Louise Phelps Kellogg. . . 23

Augustine of Hippo qua Patriot. Robert K. Richardson ... 31

Milton as a Writer on Education. Oliver M. Ainsworth. . . 41

Floundering in Modernity. George C. Clancy . 51

Bibliographical Evidence of the Vogue of Shaftesbury in the

Eighteenth Century. William E. Alderman . 57

Notes on New Names in Table of Formations and on Physical

Evidence of Breaks between Paleozoic Systems in Wis¬

consin. (With two Figures) E. 0. Ulrich . 71

The Fauna of the Lake Winnebago Region. (With one Fig¬

ure.) Frank Collins Baker . 109

Observations on Parasitic Worms from Wisconsin Fishes.

(With Plates I-III.) A. S. Pearse . 147

»

The Parasites of Lake Fishes. A. S. Pearse. . . . 161

The Anatomy of Troctes divinatorius Muell. (With Plates

IV-VI.) Ruth Chase Noland . 195

Arrhenuri from Washington and Alaska. (With Plates VII

and VIII.) Ruth Marshall . 213

New and Corrected Names of Certain Milk Bacteria. W. D.

Frost and Ruth Chase Noland . 219

The Characteristics of Certain Fecal Bacteria as Shown by the

Little Plate Method. (With Plate IX.) Ola E. Johns¬

ton and William D. Frost . 223

On the Nature of Disease Resistance in Plants. J. C. Walker 225

Some Ferns of Southwestern Wisconsin. Sister M. Ellen. . 249

Notes on Parasitic Fungi in Wisconsin, IX. (With four Fig¬

ures.) J. J. Davis . 251

Notes on Parasitic Fungi in Wisconsin, X. (With two Fig¬

ures.) J. J. Davis . 271

Page

Notes on Parasitic Fungi in Wisconsin, XI. (With two Fig¬

ures. ) J. J. Davis . . 287

The Cytology and Physiology of Venturia inequalis (Cooke)

Winter. (With Plates X and XI.) Charles N. Frey. . 303

Cytological Studies of Taphrina coryli Nishida on Corylus

americana. (With Plates XII and XIII.) Ella May

Martin . 345

The Structure and Behavior of the Nucleus in the Life History

of Phycomyces nitens (Agardh) Kunze and Rhizopus

nigricans Ehrbg. (With Plates XIV and XV.) E. A.

Bairdi . 357

A Quantitative Study of the Larger Aquatic Plants of Green

Lake, Wisconsin. (With seven Figures.) H. W. Rickett 381

The Rotifer Fauna of Wisconsin, II. A Revision of the No-

tommatid Rotifers, Exclusive of the Dicranophorinae.

(With Plates XVI-XLIII.) H. K. Barring and F. J.

Myers . 415

Proceedings of the Academy, 1921 to 1923 . 551

THE UNIFICATION OF SOUTH AFRICA: A STUDT IN

BRITISH COLONIAL POLICY

Paul Knaplund

By a treaty with the Netherlands of August 13, 1814, Britain ob¬

tained permanent possession of the Cape of Good Hoped At this

time the white population of the new dependency consisted almost

exclusively of descendants of the Dutch and of Huguenots brought

there during the latter half of the seventeenth century by the

Dutch East India Company. The settlements were small and cen¬

tered largely around Table Bay. Little effort had been made to

trace the northern boundary of the colony. Few realized then the

agricultural possibilities of the great veld of the interior and none

suspected the existence there of fabulous treasures in gold and

precious stones. The Cape was valued as a convenient half-way

station on the road to India. Isolated, the colony enjoyed almost

complete immunity from foreign attacks. No European power had

possessions within a wide radius from it ; and the nearest, the neg¬

lected Portuguese colony of Mozambique, belonged to a friendly

and allied state. Britain had no rival on the sub-continent.

During the last years of the Dutch occupation the Boers had

been restive. Lessons of the American and of the French revolu¬

tions were not lost on a frontier community kept under the pater¬

nalistic management of a trading company. When Britain secured

control, the political situation changed for the worse. The burgh¬

ers lost whatever share they formerly had possessed in their own

administration, and for more than a decade the English governor

ruled with all the power of an autocrat.^ Discontent with this

regime soon manifested itself, and it grew in strength with the

arrival of about 2,000 English settlers in the eastern provinces of

the colony. The newcomers clamored for their rights as English¬

men and soon the imperial government, somewhat grudgingly,

* For the text of this treaty see G. W. Eybers, Select Constitutional Documents Illus¬

trating South African History 1795—1910 (London, 1918), pp, 19—23.,

2 G. E. Cory, The Rise of South Africa (4 Vols., London, 1910- .... ), I, pp. '62, 63;

II, pp. 240; 241.

2

Wisconsin Academy of Sciences, Arts, and Letters,

conceded to the colonists a small shar6 in the management of local

affairs.®

Less salutary was, however, the cleavage which the coming of a

new racial element 'created between the eastern portion and the

older settlements in the west. Sectional strife looms large in the

subsequent history of the Cape.^ That the government seized upon

the arrival of “very numerous British born subjects’’ as a pre¬

text for attempts at suppressing the Dutch language must be con¬

sidered even more disastrous. The Boers were to be denationalized

by employing Britons, especially Scots, “in the parochial duties

of the Keformed Religion”; and a decree of 1825 provided that

after a given period “the English Language shall be exclusively

used in all judicial and official Acts”.® Although a later procla¬

mation modified this decision,® the failure to place the two lan¬

guages on an equal footing added fuel to the flame of discontent.

And the subsequent abolition of slavery created, in the opinion of

many Boers, an intolerable situation.'^

Beginning in 1836 a large number of the most enterprising

among the Dutch migrated or “trekked” in search for new homes.

Actuated by one great desire, to be freed from British control, they

pushed courageously into the wilderness. This object proved,

however, difficult to obtain. No sooner did the Boers settle in Natal

than Britain annexed the province. When they crossed the Orange

and established themselves between this river and the Vaal, the

Queen’s sovereignty was proclaimed also over this territory.® But

the more persistent among the malcontents crossed the Vaal and

continued their quest for freedom. Faced with the possibility of

having to extend their possessions to the equator in order to retain

a handful of recalcitrant subjects, the British government finally

decided to let them go.

* A proclamation of May 2, 1825, provided for the appointment of a council ‘‘to ad¬

vise and assist in the administration” of the colony. This consisted, however, wholly of

officials and not until 1852 was a representative government established. Eybers,

Constitutional Documents, pp. 24, 45— 5^.

^ In 1836 a separate district government was established in the eastern provinces;

ibid., pp. 39-41.

Ibid., pp. 23, 24. This proclamation was issued by the governor. Lord Somerset,

and dated July 5, 1822.

® By ordinance of December 13, 1826; ibid., p. 107. Not until 1882 did the Dutch

secure equal language rights in the Cape parliament; ibid., p. 66.

* A “Manifesto of the Emigrant Farmers” printed in the Qrahamstown Journal, Feb¬

ruary 2, 1837, is given by Eybers, pp. 143—145. For additional declarations see ibid.,

pp. 154, 155, 167-169.

* Several important documents dealing with these events are found in Eybers, Con¬

stitutional Documents, pp. 145—166; 260—275.

Knaplund — A Study in British Colonial Policy

3

This step was in accordance with the views on colonial policy

propounded by the leading economists of the period ; but in taking

it the colonial office seems to have been actuated more by a desire

to solve pressing practical problems than by regards for abstract

theories. With the growth of a sentiment favoring and the final

adoption of free trade and with the repeal of the navigation acts

many Englishmen considered the overseas dependencies a bur¬

den — a source of weakness and expense. Limitation rather than

expansion of the boundaries of the empire became their slogan.

Neither the governments nor the leading statesmen of the period

formally accepted this doctrine, but it influenced the attitude of

the public towards such questions as colonial defense. Since Water¬

loo a large portion of the British army had been kept in the col¬

onies.^® When, by the end of the forties, it was deemed necessary

to strengthen the home defenses Parliament appeared unwilling to

increase the army estimates; and the desired increase in the force

stationed in the United Kingdom could be obtained only by with¬

drawing the outlying garrisons.

This was the opinion of Earl Grey, secretary of state for war

and the colonies 1846-1852. He deplored the agitation of the

Little Englanders”, but he was deeply impressed with the views

of the Duke of Wellington, and other eminent authorities, who

warned England of the danger in neglecting to secure adequate

protection for the British Isles. By recalling the scattered garri¬

sons, already considered useless in case of war, a strong force could

be created at the imperial base. Steam navigation which had in¬

creased the vulnerability of Britain also facilitated the sending of

reinforcement to any dependency threatened by a foreign foe.^^

In South Africa the presence of numerous warlike natives com¬

plicated the situation. Here the military expenditures were heavy

and a further extension of British territory would necessitate an

increase of the local force at a time when the troops were needed

elsewhere. Therefore, quite apart from the problematic value of

such new possessions, an expansionistic policy might tend to aid in

® A summary of these views is given by H. Duncan Hall, The British Commonwealth

of Nations (London, 1920), pp. 39—53.

For the opinions of leading British statesmen see debates in Parliament Jan. 16,

1838; April 13, May 29, June 12 and 30, 1840; February 8, April 19, and May 6,

1850. Hansard, 3rd series, XL, cols. 34-73; LIII, cols. 1063, 1064; LIV, cols. 731

732, 1121, 1158; LV, cols. 239-245, 268, 269; CVIII, cols. 546-566, 606, 607, 1009; CX,

cols. 565, 566,, 678, 589, 1171, 1172. See also General Sir Robert Biddulph, Lord Card-

weU at the War O^ce (London, 1904), pp. 38-41.

“ Biddulph, Lord Cardwell, p. 39,

4 Wisconsin Academy of Sciences, Arts, and Letters.

jeopardizing the safety of the whole empire. That Lord Grey had.

this under consideration appears certain, and it was he who de¬

cided to limit the empire’s responsibilities on the sub-continent.^^

By the Sand River Convention of January 17, 1852, and the Bloem¬

fontein Convention, two years later, the Boer settlements beyond

the Vaal and those between the Orange and the Vaal respectively

were cut adrift.^^ These agreements laid the foundations for the

future republics. White South Africa became divided.

Hardly had this policy been adopted before protesting voices,

were heard.^^ The year of the latter of the two conventions wit¬

nessed the appearance of Sir George Grey as governor of the Cape

and high commissioner for South Africa. Although still a young

man, he had already served with distinction as governor of South

Australia and of New Zealand.^® In the latter colony he faced a

native problem bearing some similarity to that of South Africa

and had perceived serious danger in having foreign powers estab¬

lished in the neighborhood of a weak British colony.^® Independ¬

ent Boer republics might, indeed, likewise become potential rivals,

or enemies of Britain, while a union of all the white communities

besides precluding this, would also enhance their safety.

Soon the able and energetic, but also proud and imperious, pro-

consul actively championed unification. In 1856 he inquired

whether the government ‘ ‘ might not be disposed to retrace the step

which led to the abandonment of the Orange River Sovereignty ’ ’ ;

and he advocated ‘‘a united South Africa under the British flag”.^^

Although Henry Labouchere, then colonial secretary, declined ta

discuss the question Grey persisted in his efforts and received some

encouragement both in South Africa and, later, from the home

government. The Volksraad of the Orange Free State passed a

^2 Hansard, 3rd series, CXXXIII, cols. 72, 77.

For the texts of these documents see “Reports on the Cape of Good Hope” in British

Parliamentary Papers, hereafter cited P. P. 1853—54, Vol. Ill, Part I, pp. 36, 37 ; and

Eybers, Constitutional Documents, pp. 281-285, 357-359.

Inhabitants of the Orange River Sovereignty protested against the abandonment of

the territory. Whether these protests represened the -wishes of the majority is, of course,

difficult to ascertain. See P.P. 1853—54, Vol. Ill, Pt. II, pp. 8—18, 21.

No adequate biography of this remarkable man has yet been written. The two we

have, G. C. Henderson, Sir George Grey, Pioneer of Empire in Southern Lands (Lon¬

don, 1907), and W. L. Rees and L. Rees, The Life and Times of Sir George Grey (2

vols., London, 1892), leave much to be desired.

Grey urged in vain the annexation of various groups of islands in the South Pacific-

to forestall action of any other European power. Rees and Rees, Sir George Grey, I,,

pp. 128-132.

Henderson, Sir George Grey, p. 168. ^

Knaplund- — A Study in British Colonial Policy

5

resolution favoring “a union or alliance with the Cape Colony’’/®

and in September, 1858, Sir E. Bulwer Lytton invited Grey to give

liis opinion on the question of federating the South African col¬

onies. Replying in a dispatch of November 19th, the high Com¬

missioner urged consolidation of all the European communities

and suggested that the government should take the initiative by

.securing the passage of an enabling act. In doing this Grey doubt¬

less went further than Lytton had originally intended him to go.

The considerations of imperial interests which had made the pre¬

vious withdrawal desirable were opposed to the forward policy

suggested by the zealous governor. In his eagerness to promote

the cause of union Grey disobeyed orders and was recalled. While

the colonial office later cancelled this recall, the attitude towards

federation remained unchanged. Disappointed, the governor

voiced his fear that ‘‘the opportunity of establishing such a fed¬

eration as I had proposed has now been lost forever.^*^

Two events of the succeeding decade deeply influenced Britain’s

colonial relations and caused the imperial statesmen to modify

their views regarding a South African union. The first of these,

the withdrawing of the garrisons from the self-governing colonies,

took place largely because it was deemed necessary to strengthen

the home defenses without increasing the burden on the British

taxpayers. The wishes of the colonies were completely disre¬

garded.^® On the other hand, the second, the establishment of the

Dominion of Canada, was the result of a movement which orig¬

inated in British North America and received the hearty support

of the colonial office “on the ground, among others, that [federa¬

tion] was eminently calculated to render easier and more effectual

the provisions for the defence of the several Provinces.

Once having accepted the principle of colonial federation the

Rome government showed considerable zeal in attempting to secure

a wdde application for it. The Leeward Islands were federated

according to a plan prepared by Governor Sir B. C. C. Pine. In

this instance the impetus came from home. The Gladstone govern¬

ment seem to have entertained hopes that consolidation would ulti-

G. McCall Theal, The History of South Africa Since 1795 (5 Vols., London, 1908),

III, p. 175.

Despatch of July 31, 1859, P.P. 1860, Vol. XLV No. 357, p. 1. For the text of

Sir George Grey’s confederation despatch see Frank R. Cana, South Africa from the

Great Trek to the Union (London, 1909), pp. 298—309.

2** See Paul Knaplund “Intra-Imperial Aspects of Britain’s Defence Question” in

The Canadian Historical Review, III, pp. 120—125.

Cardwell to Arthur Gordon, April 12, 1865, P.P. 1867, XLVIII, cd. 3769, p. 117.

6 Wisconsin Academy of Sciences, Arts, and Letters.

mately lessen the charge upon the imperial exchequer. The island¬

ers manifested little enthusiasm although they complied with the

wishes of the colonial office and accepted the governor’s scheme.^“

A similar method of procedure was adopted in dealing with the

South African situation. Frequent complaints had been made of

the heavy military expenditures at the Cape, nor did the repre¬

sentative system of government, established in 1852, function prop¬

erly. A change was deemed necessary, especially one which would

enable or induce the colonies to assume greater responsibility for

local defense.^^ In his instructions to the newly appointed gover¬

nor and high commissioner. Sir Henry Barkly, Lord Kimberley

discussed the various problems of the colony and called attention

to the success of the Canadian federation.-^ This the former inter¬

preted as clothing a desire to have a similar change affected in

South Africa. Immediately upon assuming the duties of his offices

Barkly, therefore, took steps toward the establishment of a federal

state. Disregarding the protest of the Cape executive council he

appointed a commission to consider the advisability of dividing the

colony into three or more provinces and with these form a federa¬

tion which might ultimately embrace all of white South Africa.^^

Lord Kimberley approved. In a despatch of Nov. 16, 1871, he

authorized Barkly “to sanction the convening of Delegates from

[the Boer republics] and Natal for the purpose of considering the

conditions of a Union”. Hopes were also expressed that such a

union, if established, would assume the responsibility for defense.^®

The policy was endorsed by the House of Commons,^^ but South

Africa remained indifferent. Only the diamond diggers of Griqua-

land West supported federation.^® A strong majority at' the Cape

2sp.P. 1871, XLVIII, cd. 353.

23 Granville to Governor Sir P. E. Wodehouse, Dec. 9, 1869. P.P. 1871, XLVII, cd,

459, pp, 13—15. See also Kimberley to Sir Henry Barkly, Nov. 17, 1870; ibid., p. 66.

2^ Kimberley to Barkly, Oct. 17, 1870; ibid., p. 47.

Ibid., pp. 170, 171 ‘We . , , record our conviction that . . . the attempt to feder¬

ate upon any satisfactory basis the eastern and western provinces of this Colony — the

Basutos and adjacent native territories — including the Diamond-fields, the two Boer

republics, and Natal, or indeed, any two or more of them, will present the very greatest,

if not insuperable difficulties; and we fail to see the practical object or advantages of

any such federation in the absence of that external pressure and rivalry of a powerful

adjacent nation, which has had the chief share in effecting the consolidation of the

Dominion of Canada.” Minute of the colonial secretary, treasurer general, auditor

general, and collector of customs, April 26, 1871; ibid., p. 180. See also P.P. 1872,

XLIII, cd. 508, p. 11.

^<^Ibid., cd. 508, p. 14.

2’’ Hansard, 3rd series, CCXI, cols. 806—815. Debate May 28, 1871.

28P.P., 1873, XLIX, cd. 732, p. 121. The address is dated August 28, 1872. It

should be remembered, however, that Griqualand West was under the crown colony form

of government which would naturally prove distasteful to the miners, the majority of

whom were Englishmen.

Knaplund — A Study in British Colonial Policy

7

resented the recognition afforded the separatist tendencies in the

eastern provinces and especially a suggestion for moving the capi¬

tal.^® By annexing the diamond fields, claimed by the Orange Free

State, Britain had antagonized the Boer republics.^® Voluntarily

these would not surrender their independence in exchange for a

colonial status. Lacking the most essential elements for success,

popular support, the Barkly-Kimberley attempts at federation

accomplished nothing.

This failure might have taught British statesmen how to proceed

if they desired to establish a South African union, but it did not.

Lord Kimberley’s Conservative successor, the Earl of Carnarvon,

who had had the good fortune of steering the British North American

Act through Parliament, erred even more egregiously. Deeply

impressed with the success of the Canadian experiment, he was

determined to confer the boon of federation upon South Africa

even to the extent of imposing it upon the colonies. In dealing

with this question the noble earl, who as a statesman possessed

many admirable qualities, reminds one of the hero in Kuneberg’s

poem ^^Sven Dufva”, his heart was sound, but his head was

rather weak.

At first Carnarvon seems to have recognized the necessity of

securing public approval for his cherished plan. In 1875 he pro¬

posed to call a conference at some place in South Africa which

should discuss native policy, the enforcement of criminal law,

and federation. Probably fearful lest' the frontiersmen should not

know how to proceed, he also suggested how the delegates should

be distributed and chosen and repeated the blunder of his predeces¬

sor in recognizing the sectional interests at the Cape.^^ When the

colonists appeared lukewarm^^ Carnarvon sent James Anthony

Froude, the historian, to South Africa as his personal representa¬

tive. The choice proved singularly unfortunate. While touring

^Ibid., pp. 43, 44.

P. A. Molteno, The Life and Times of Sir John Charles Molteno (2 vols. London,

1900), I, pp. 185, 186. When the Cape Colony received responsible government President

Burgers of the South African Republic, in a letter of December 27, 1872, congratulating

the first prime minister, J. C. Molteno, on the occasion, said: “I am confident that it

will direct the spirit of the nation in that proper channel which will ultimately lead to

a closer union between the different colonies and states of South Africa.” Ibid., p. 202.

A little more generosity on the part of the imperial government would have smoothened

the way for the desired federation.

See despatch, Carnarvon to Barkly, May 4, 1875. P.P., 1875, LII, cd. 1244,

pp. 1-3.

^^Ibid., 1876, LII, cd. 1399, pp. 4, 5; Life of Molteno, I, pp. 335, 336, 346: Un¬

qualified support was received from Natal and Griqualand West, P.P., 1876, LII, cd.

1399, pp. 19, 25, 55.

8 Wisconsin Academy of Sciences, Arts, and Letters.

the Cape Colony, Fronde, in the words of a competent observer,

violated ‘‘publicly and preserveringly a constitutional obligation

which the Colonists have a right to view as one of the essential

safeguards of constitutional right. This emissary of the colonial

secretary virtually carried on a campaign against the ministry at

the Cape.^^ By appealing to the particularism in the eastern

provinces, he could truthfully report enthusiastic receptions; but

these appeals also increased the suspicion with which the majority

viewed his agitation and the cause he was supposed to further.^^

Despite, or rather partly because of, Froude’s activities. Lord

Carnarvon finally realized that the desired conference would not

meet in South Africa. He then decided to hold it in London. On

August 3, 1876, representatives of South Africa met at the colonial

ofiSce. Only those from Natal had been duly elected, the others

were officials chosen by Carnarvon. J. C. Molteno, the prime

minister at the Cape, was in London but refused to attend, deeming

himself bound by the defeat of the federation project in the colonial

legislature; and President Brand of the Orange Free State at¬

tended only after he had received definite assurances that there

would be no serious “discussion of the merits of South African

Confederation.^’^® Under such conditions the conference could

hardly be expected to achieve anything.

Carnarvon showed, however, a remarkable, although misplaced,

perseverance. Encouraged by Messrs. Blaine and Paterson from

the eastern provinces where the belief prevailed that confederation

would bring them local anatomy, the colonial secretary announced,

October 26, 1876, that in his opinion the time was now ripe for the

drafting of a permissive bill.®'^ Shortly afterwards a Draft Bill,

which was in reality a complete constitution, was transmitted to

Lord Blachford, “Native Policy in South Africa” in The Edinburgh Review, April,

1877, CXLV, pp. 232, 233. See also Letters of Lord Blachford, edited by G. E.

Marindin (London, 1896), p. 364. Lord Blachford had served as permanent under¬

secretary of state for the colonies, 1860—1871.

Life of Molteno, I, pp. 400, 401.

On December 24, 1875, Eroude declared hopefully, “The ministers of the Cape

Colony have the appearance of victory, but we have the substance.” John Skelton, The

Table-Talk of Shirley (London, 1895), p. 153. He had run hut not read.

Present at the conference were Lord Carnarvon, Sir Garnet Wolseley, Sir Theophiius

Shepstone and Messrs. Froude, Akerman, and Robinson. See P.P., 1876, LII, cd. 1631,

pp. 51, 52. In a communication to Lord Carnarvon, dated October 2, 1876, Molteno

expressed the opinion that a South African consolidation must assume the form of a

legislative union. See ibid., p. 11.

Ibid., 1877, LX, cd. 1732, pp. 13, 14. See also Herbert Paul, The Life of Froude

(New York, .1905), p. 269.

Knaplund — A Study in British Colonial Policy

9

Sir Henry Barkly to be presented before the people of South

Africa.^®

It met a fate similar to that of the proposal for a conference.

The Cape saw in it another attempt to govern the colonies from

Downing Street; Natal was dissatisfied; and even the high com¬

missioner subjected the measure to severe criticism. In the

republics the reception appeared decidedly hostile. The Orange

Free State could not ‘ ‘ accede to a union by which this State would

sacrifice its independence”; and the Volksraad of the South Afri¬

can Republic rejected the bill.^^

Suspecting Barkly of being too much under the influence of

the Cape ministry Lord Carnarvon replaced him with Sir Bartle

Frere, who received definite instructions to obtain a confederation

of the various colonies and states.^® To further this the Transvaal

was annexed/^ and the amended Draft Bill was introduced in Par¬

liament, where it met with general approval, and upon receiving

the Queen’s assent it became the South African Act, 1877

All in vain. Natal favored the convening of a conference to

discuss the question, but the Cape legislature remained obdurate.

The annexation of the Transvaal had increased the hostility

with which the Dutch, both in the English colonies and in the

Orange Free State, regarded this purely English project.^^ With

the uprising in and the retrocession of the Transvaal the cause

of confederation became entirely hopeless. In August, 1882, the

South African Act expired.

Lord Carnarvon’s persistent attempt at federating the sub¬

continent throws interesting sidelights upon Britain’s attitude

towards the dependencies and upon the colonial policy of Dis¬

raeli’s Ministry. One discerns a tendency to revive the paternal¬

istic methods of “Mr. Mother Country” of the pre-Durham-Buller

period and, in addition, a certain un-English faith in legal form¬

ulas. Federation had triumphed in Germany and proved benefi¬

cial to Canada, therefore it ought to be adopted in South Africa.

Carnarvon to Barkly, Dec. 14, 1876. P.P., 1877, LX, cd. 1732, pp. 20-29.

pp. 32-36, 41-43; 1878, LV, cd. 1980, pp. 7-9, 18; Cana, South Africa, p. 80.

41) Frere was in the opinion of Carnarvon, “The statesman . . . most capable of

carrying my scheme of confederation into effect . . Carnarvon to Frere, Oct, 13,

1876. John Martineau, The Life and Correspondence of the Right Honourable Sir

Bartle Frere (2 Vols., London, 1895), II, p. 162.

J. A. Fronde, Oceana (London, 1886), p. 53.

*2 P.P., 1877, VI, “House of Lords Bill, 271.” For the changes in the original

Draft Bill see ibid., XLX, cd. 1732, pp. 43—52.

Life of Molteno, II, pp. 364, 427, 428.

10 Wisconsin Academy of Sciences, Arts, and Letters,

Few efforts were made to ascertain whether the conditions there

were similar to those which had made the Canadian federation

inevitable and, above all, whether the people concerned really

desired consolidation in any form. The powerful social and eco¬

nomic factors which proved so potent thirty years later in creating

a sentiment in favor of union were largely non-existent in the

seventies. At this time even the most tactful of colonial secre¬

taries would probably have failed in an endeavor to create a

federation.

Several of those who had been responsible for these early

attempts finally realized that a different method of procedure

would be necessary in order to achieve success. Sir Bartle Frere,

in writing to Sir George Colley, August 26, 1880, said : ‘ ‘ One great

mistake hitherto seems to me to have been trying to hasten and

push on what can only result from natural growth, which must of

necessity be tardy if it is to be enduring. Lord Kimberley,

upon resuming the duties of the colonial office, instructed Frere ’s

successor, Sir Hercules Kobinson, to work for federation but

added, ‘‘It will be more convenient that any fresh movement for

federation or union should be initiated spontaneously by the

Colonies. ’ Froude, who had sinned grievously, also saw the light.

Commenting on the South African confederation effort in his

Oceana, he says : “ If South Africa is to rule itself under a consti¬

tutional system, we must cease to impose English views of what

is expedient on a people unwilling to act upon them.’^^® And this

became the established policy of the home government.

During the eighties and the nineties the Transvaal drifted

politically farther and farther away from British connections.

The London convention of 1884 increased the degree of inde¬

pendence enjoyed by the republic, but did not accomplish a com¬

plete reconciliation. Boers and Britons continued to regard each

other with a suspicion which increased when the latter blocked all

efforts on the part of the South African Republic to secure an

outlet to the sea or to expand into the interior.

Powerful centripetal forces came into existence, however, with

the discovery and opening up of the great Witwaters Rand gold

mines. This region became the economic center of South Africa.

It provided a market for the produce of the colonies and states,

^ Martineau, Life of Sir Bartle Frere, II. p.-387.

«P.P., 1881, LXVI, cd. 2754, p. 4.

J. A. Froude, Oceana, pp. 59, 60.

Knaplund — A Study in British Colonial Policy

11

proved enormously important for the commercial and shipping

interests of the coast towns^ and stimulated the construction of

railroads which converged on the Eand, Needs arose for adjusting

railway rates and customs duties; and the Cape Colony, Natal,

and the Orange Free State formed a customs union.^^ The Trans¬

vaal kept aloof, rather sullenly. With the construction of a rail¬

way to Delagoa Bay, in the Portugese territory, this republic

became practically independent of the Cape and the Natal ports

and showed little inclination to make sacrifices for the benefit

of its neighbors. These, on the other hand, faced serious losses

both among private concerns and especially for the state owned

railways which had been built primarily for the purpose of secur¬

ing the Rand traffic. A conflict of economic interest ensued which

had a disturbing influence in South Africa and affected adversely

the attitude of British imperialists to the Transvaal.

In the period between the collapse of Carnarvon’s effort and

the outbreak of the Boer war, several organizations kept the feder¬

ation issue before the people of South Africa. Prominent among

these was the Afrikander Bond which aimed at ^^the formation of

a South African nationality.”^® Founded by Boers, the Bond at

first gave a narrow interpretation of the term ^^nationality” and

often opposed Boer as Boer to Englishman as Englishman. Later

its vision broadened, and in 1896 it was described as ^‘an organiza¬

tion which draws together and unites for common purposes a

number of the early colonists and others holding certain views on

social and political matters.”^® Other bodies, such as the South

African Political Organization and the South African League,

openly advocated union of all the whites.®®

Cana South, Africa, pp. 138, 139. The Times expressed the hope, Nov. 9, 1891, that

practical necessity for concerted action would overcome the existing prejudices and

bring about union. British officials also voiced the need of a common policy in despatches

to the colonial office. P.P., 1890, XLVIII, cd. 5897, pp. 4, 5.

Sir Lewis Mitchell, The Life and Times of the Bight Honourable GeeU John

Rhodes (2 vols., London, 1910), I, pp. 293, 294. See also Basil. Williams, Oecil Rhodes

(London, 1921), pp. 60, 61.

Olive Schreiner and C. S. Cronwright Schreiner, The Political Situation (London

1896), p, 24; Williams, GecU Rhodes, p. 67.

The Times, Sept, 7, 1896. Nor had the imperial government given up all hopes

for such a union. The colonial secretary, the Marquess of Ripon, said, Sept. 6, 1894:

“What I look to is a sort of Federal Union of South Africa— British Territory, South

African Republic, and Orange Free States — 'in which we, of course, should have the

hegemony, hut no more .... and I should care little whether the Transvaal be¬

came a British Colony or remained the South African Republic within such a Federor

tionP Lucian Wolf, Life of the First Marquess of Bipon (2 vols., London. 1921), II,

p. 222. On this point the views of Lord Ripon seem to have almost coincided with

those of Cecil Rhodes; see Williams, OeeU Rhodes, p. 68.

12 Wisconsin Academy of Sciences, Arts, and Letters,

Tlie greatest single proponent of this idea was doubtless Cecil

John Rhodes. A true empire-builder he combined the ability to

dream dreams and see visions with a great capacity for practical

affairs and an element of ruthlessness, often so necessary for suc¬

cess. Rhodes saw clearly the benefits which could be derived from

union, presented its cause in several public addresses, and worked

persistently to arouse popular interest in its favor. His private

secretary testifies that the union of all the white communi¬

ties in South Africa was Rhodes’ “Lifelong dream. Unfortu¬

nately his zeal for this proved at times stronger than his scruples

and even than his discretion. It led him to plot the Jameson

Raid — a crime and a blunder which discredited a great cause and

precipitated a bloody and protracted war.®^

Pew wars fought by Britain in modern times have caused

fiercer partisan controversies than that in South Africa, 1899-1902.

Condemned by hostile critics as an unjust imperialistic attack upon

two weak states at the behest of a small band of adventurers and

capitalists, its defenders contended with equal vigor that it had

been forced upon the empire by Kruger ’s violation of treaty

obligations ; his unfair treatment of the foreign born, the majority

of whom were British ; and necessitated by regards for the safety

of the South African colonies. The Kruger government in the

Transvaal was doubtless narrow in it's conservatism, possibly cor¬

rupt and certainly galling to the Uitlanders. Germany’s activities

might also justify serious apprehensions.^^ But it appears prob¬

able that this war, like most confiicts of ancient and modern times,

could have been avoided if a more conciliatory policy had been

adopted by the British government. Chamberlain, Milner, Rhodes,

and their lieutenants, were the proponents of great plans for the

safeguarding and consolidation of the overseas possessions and

favored aggressive methods in securing their aims. The republics

lay close to the highway to the interior. Their annexation rounded

off the South African Colonial empire, made it safer against for-

P. Jourdan, Cecil Rhodes, His Private Life hy Hw Private Secretary (New York,

1911), pp. 162, 163. Among the advocates of union were also found Mr. P. A.

Molteno, the son of the first prime minister at the Cape. He favored a federation

similar to that of the United States. See P. A. Molteno, A Federal South Africa (Lon¬

don, 1896).

The best brief account of the Raid, revealing Cecil Rhodes’ connections with it

and the results, is found in Williams, Cecil Rhodes, pp. 242-275.

Wolf, Ripon, II, pp. 231-233. German plans for an expedition to the Transvaal,

1896, seem to have been thwarted only by the common sense shown by the Portugese

foreign minister, the Marquis de Several. See Baron von Eckardstein, Ten Years at

the Court of St. James (London, 1921), pp. 84, 85. i

Knaplund — A Study in British Colonial Policy

13

eign attacks, and benefitted important economic interests both

there and at home.®^

Upon the cessation of hostilities®^ enthusiasts for the cause of

union believed that it could be obtained by quick and decisive

action. Lord Milner shared this view. He therefore advocated the

suspension of the Cape constitution, whereby all the British pos¬

sessions would be placed more nearly on an equal footing and ‘ ‘ the

restoration of self-government in all the colonies [could be made

to] coincide with the establishment of federation.’’®® But Cham¬

berlain, who favored strongly the desired consolidation, refused to

sanction coercive measures. Mindful of past mistakes he abstained

from dictating to South Africa; Lord Carnarvon’s blunders were

not without beneficial results.®’^

Although waiting watchfully. Downing Street was not inactive.

Chamberlain himself called the attention of the South Africans

to the benefits which might be derived from federation and ap¬

pealed to them to think of their country as a whole.®® And the

numerous young men sent to assist Lord Milner in reorganizing

and rebuilding the new colonies, proved to be active and persistent

advocates of union. Milner’s ‘ ‘ Kindegarten ” combatted par¬

ticularism in all its forms and attempted throughout to direct' the

thought and aspirations of the colonists so as to foster a spirit of

unity.®®

The work of reconstruction offered, indeed, favorable oppor¬

tunities for this. A common police force, the South African Con¬

stabulary, was created and a joint loan of thirty-five million pounds

secured for the Transvaal and the Orange Eiver Colony.®® During

the war the railways of the two republics had been seized and

organized as one system. This was continued, and an Inter-Colonial

Council was established which assumed charge of the railways.

Wolf, Ripon, II, pp. 253-257; Sir William Butler, An AutoMography (London, 1913),

pp. 404-455.

The Orange Free State and the Transvaal were annexed by proclamations of May

24 and Sept. 1, 1900. Peace was not concluded until May 31, 1902, by the Treaty of

Veereeniging. For the texts of these documents see Eybers, Constitutional Documents,

pp. 344—347, 514, 515. The territories of the two republics received the crown colony

form of government.

The Times' History of the War in South Africa, edited by L. S. Amery (7 vols.,

London, 1909), VI, pp. 66, 67. See also Mitchell, Life of Rhodes, II, p. 283.

Harold Spender, General Botha (New York, 1916), p. 154.

See speech at Cape Town, February 23, 1903. Mr Chamberlain’s Speeches, ed. by

Charles W. Boyd (2 vols., London, 1914), II, pp. 109—112.

W. B. Worsfold, The Union of South Africa (Boston, 1913), p. 121. Called Mil¬

ner’s “Kindergarten” because of their youth.

J. Buchan, The African Colony Studies in Reconstruction (London, 1903), p. 245.

14 Wisconsin Academy of Sciences, Arts, and Letters.

the constabulary, and a number of minor services. The net earn¬

ings of the railways w^ere used in defraying the expenses connected

with the loan, the constabulary, and other charges.®^ Thus the two

new colonies had a common administration of important depart¬

ments and local barriers were broken down. The revived customs

union included all the colonies and the conference in 1903, dealing

with this question, expressed the hope that the time might not be

far distant when a Commonwealth of South Africa would be

created; and it also recommended the appointment of a commis¬

sion to study the native problem, so important for all the colonies.®^

Prominent among the factors which aroused the people of South

Africa to a realization of the necessity for union were questions

connected with the customs, railway administration, labor supply

for the mines on the Rand, and native policy. The last came

especially into prominence on account of the Natal rebellion of

1906 which revealed that the Kaffirs had not lost the martial

spirit of their ancestors; In the report of the Native Affairs Com¬

mission of 1905, attention was called to a situation highly dis¬

quieting. Outnumbered more than five to one the whites faced a

tremendous problem in dealing with native races noted for their

fecundity and virility. While the old order among them disap¬

peared rapidly no new restraining influence seemed ready to take

the place formerly occupied by the chiefs. And the spread of a

feeling of racial solidarity, as a result of the Ethopian movement,

gave further cause for alarm.®®

All these elements were cleverly utilized by the young English¬

men employed in the crown colony administrations of the Trans¬

vaal and the Orange River Colony. Their well-written and well-

documented arguments were finally published in book form entitled

The Government of South Africa. And in another work. The

Framework of Union, they showed the characteristics of some of

the leading federated and unitary governments of the world.®^

After the promulgation of self-governing institutions in the new

colonies the advocates of union found powerful allies among the

Lord Milner saw great possibilities in the Inter-Colonial Council “affecting much

more than the two Colonies,” Milner to Chamberlain, April 6, 1903. P.P., 1903, XLV,

cd. 1641, p. 4.

^Ibid., cd. 1640, p. 15.

Ibid., 1905, LV, cd. 2399; The South African Natives: Their Progress and Present

Condition, edited by The South African Native Races Committee (London, 1908), pp.

192, 228.

^ The Government of South Africa (Cape Town, 1908); The Framework of Union

(Cape Town, 1908).

Knaplund — A Study in British Colonial Policy

15

Boers. The Afrikander Bond urged ‘‘the development of a

feeling of national unity in South Africa and a federal union of

the British South African Colonies, keeping in view the mutual

interests of these Colonies and the supreme authority of the Brit¬

ish Crown. Mr. F. S. Malan, leader of the Bond forces in the

Cape assembly, began in 1906 a series of vigorous articles favoring

unification in 0ns Land, the leading Dutch newspaper in South

Africa.®® Of even greater significance was the aid received from

the leaders of the old republicans. Some of these had, indeed,

agitated for a united South Africa in the years preceding the

war, but by this they meant union under the republics. Now they

advocated consolidation under the Union Jack.

M. F. Steyn, ex-president of the Orange Free State, related in

language of pathetic simplicity how the view of a pool of blood

from soldiers representing the Free State, the Transvaal, and the

Cape had brought him to realize the folly of internecine strife.®’’

His weighty influence was thrown in the scale favoring unification.

General Smuts had already in 1895, as Mr. Advocate Smuts, urged

cooperation among the whites.®® With the war over he pleaded

eloquently for reconciliation and union. Speaking at Potchef-

stroom in February, 1905, he refused to “hide the fact that the

source of all our evils was disunion, disruption.’’ “Our object

of old,” he continued, “was to found a United South Africa,

stretching as far as Zambesi or farther, but because we were at

sixes and sevens we did not succeed ^ . Let us take the

hand of brotherhood.” At Klerksdorp “he advised his hearers

to let ‘ The union of Boer and Briton resemble that of England and

Scotland not that of England and Ireland. Let us cooperate in

order to attain our old object: A United South Africa.’ ”®®

Some of Smut’s appeals for union thus antedated the grant

of self-government to the Transvaal and the Orange Free State,

“South African Union” in The Edinburgh Revieiv, vol. OCX, p. 8 (July, 1909).

p. 9.

“De dag na de slag te Graspan nabij Reitz, waar generaals De Wet en De la Rey

’n vronwelager verlosten, kvam ik op ’t slagveld en wer mij de plaats getoond waar

drie van onze mannen gesneuveld warn — ’n Kolonialer ’n Vrijstater en ’n Transvaler,

Ik zag hoe hun levensbloed in een grote plas tezamen gestroomd was, Ik stond als

genageld bij die plas! Ik sprak met niemand, ’t Was voor mij ’n heilig ogenblik.

Nieuwe hoop en nieuwe moed vervulden mij. Ik Klom op mijn paard en reed weg

overtuigd in mijn hart dat ik de ware vereniging van Zuid-Afrika gezeien had, want dat

bloed kon geen mens meer scheideni” — Prom a speech of March 19, 1908. N. J. van

der Merwe, Marthinus Theunis Steyn (2 vols., Cape Town, 1921), II, p. 221.

®* For quotations from this speech see N. Levi, Jan Smuts (London, 1917), pp. 25—27.

^Ibid., p. 74.

16 Wisconsin Academy of Sciences^ Arts, and Letters.

but this timely concession increased his zeal and put the Boers in

a receptive mood. Britain had proved generous and trusted them.

Now their great commander in the weary years of a life and death

struggle came forward with appeals for union. ‘ ‘ The old Boers, ’ ’

said General Botha, “were pioneers of the Transvaal and as they

were the pioneers in that matter so they should be on the question

of Federation.”^® It held, indeed, promises of a great future

for their race. They formed the majority of the total white popu¬

lation and in 1907 three out of the four governments were con¬

trolled by them. If united, the Boers had hopes of dominating a

state which would rank with the great federations of Canada and

Australia. Not all the Boers could appreciate these possibilities,

but the majority of them were good “followers” and in Generals

Botha and Smuts they possessed leaders and statesmen cast in a

heroic mould.

When the conditions appeared favorable members of Milner’s

“Kindergarten” took prompt action. In 1906 a statement was

drawn up which showed the urgent need for an administrative

union.’^^ Through the assistance of Dr., later Sir, Starr Jameson,

then prime minister at the Cape, this was brought before the people

of South Africa. In a minute of November 28, 1906, his govern¬

ment invited the high commissioner. Lord Selborne, “to review

the general situation in South Africa in such a manner as may

enable the people of this country to appreciate the difficulties of

administration under the present system, and to consider whether

(and if so by what means) it is advisable to establish a central

national government embracing all the British Colonies and Pro¬

tectorates. ’

Lord Selborne forwarded the minute to the governments of

Natal, the Orange Kiver Colony, and the Transvaal and to the

administrator of Southern Ehodesia.'^^ When these concurred in

the request he issued, January 7, 1907, his famous Federation

Memorandum, which included the above mentioned statement.

The Memorandum presented in a clear, concise, and convincing

manner the actual and unsatisfactory condition on the sub-con¬

tinent as well as the dangers connected with a continued disunion.

Quotation from a speech at Standerton, Jan., 1907. Spender, General Botha, p. 194.

The authors of this statement were Messrs. Lionel Curtis, W. L. Hichens, Patrick

Duncan, R. H. Brand, and Feetham. See Worsfold, Union of South Africa, pp. 122,

123; and Worsfold, The Reconstruction in the New Colonies Under Lord Milner (2

vols., London, 1913), II, 398-399.

’'“P.P., 1907, LVII, cd. 3564, p. 3.

-‘^Jhid., p. 9.

Knaplund~A Study in British Colonial Policy

17

The many serious problems which confronted the various states

could not possibly be solved by piecemeal measures. Three roads

were open: ‘^The make-shift regime of the High Commissioner,

the jarring separatism of the States of South America, the noble

union of the States of North America. ’ Although recommending

union. Lord Selborne disclaimed any intention of attempting to

force it upon the colonists. In his opinion ‘‘no healthy movement

towards federation can emanate from any authority other than the

people of South Africa themselves.”^® This official and temperate

analysis of conditions which most thinl^ing men admitted were

serious had a marked effect upon public opinion and encouraged

those who worked for union. An active educational propaganda

was inaugurated and it continued till the goal had been reached.”^^

By 1907 it was generally felt that South Africa approached

a crossroad. The existing customs union and railway agreements

failed to satisfy the different colonies.'^® Since 1905 both the Cape

Colony and Natal had experienced an economic depression which

yearly grew more acute. Large deficits were accumulated and

both colonies demanded higher tariff and railway rates. The

finances of the Transvaal were, on the other hand, in a flourishing

condition and here the people clamored for a lowering of the

rates."^®

As a result the government of the Transvaal served notice.

May, 1907, to terminate the customs union and the railway agree¬

ment. A conference took place at Pretoria in the following year.

Agreement could be reached only on a series of resolutions, to be

submitted to the colonial parliaments, which declared that ‘ ‘ an early

union under the Crown of Great Britain” was desirable, and sug¬

gested that delegates should be appointed to consider and report

on the most desirable form of such a union and prepare a draft

constitution.®®

Ibid., pp. 12-61. It was published as a pamphlet in July of the same year with the

title, A Review of the Mutual Relations of the British South African Colonies (Cape

Town, 1907).

P., 1907, LVII, cd. 3564, p. 6.

^^Ibid., p. 5.

R. H. Brand, The Union of South Africa (Oxford, 1909), p. 31.

Ibid., pp. 25-27; Paul Lederer, Die EntwicTclung der sudafricanische Union auf

Verlcehrpolitischer Grundlage (Leipzig, 1910). J. Conacher, Report upon the Distribu¬

tion of Oversea Traffic Between the South African Railways (Pretoria, 1908) ; H. E.

S. Fremantle, The New Nation (London, 1909), p. 6; P.P., 1907, LVII, cd. 3564, pp.

35, 36; The Government of South Africa, I, pp. 195—229, 280—282.

The Times, May 4 and 9, 1908; Fremantle, The New Nation, p. 154.

The Times, May 6, 1908; Sir E. Walton, The Inner History of the National Con¬

vention of South Africa (New York, 1912), p. 26.

18 Wisconsin Academy of Sciences, Arts, and Letters,

The Pretoria resolutions were, on the whole, well received by

the colonies. Details were criticized, but there was general con¬

currence on the main issue. All of the party leaders supported

the call for a National Convention; and the president of the

Afrikander Bond expressed the belief that the resolutions ‘‘would

be a powerful factor in speedily establishing a united South

Africa.®^ In England the reception was equally cordial. The

Times declared that “The news was heartily welcomed here as a

sign that the States of the sub-continent were entering the path

which the best and highest interests of all of them dictate.’’®^ A

resolution was passed unanimously by the House of Commons ex¬

pressing the hope that the government would welcome the adoption

of provisions calculated to render possible the ultimate inclusion

of all of South Africa in a federal union. To this Mr. Winston

Churchill and Colonel Seely, on behalf of the government, replied

that, warned by precedent, they were determined not to attempt

to lead, but to leave the matter for the decision of South Africa.

The uncertainty of the situation prevented them from making any

declaration as to what would happen in the event of federation.

They would watch and wait,®® a policy followed consistently while

the convention was at work and the fate of South Africa hung in

the balance.®^

But the young imperialists in South Africa were furiously

active. Mr. Lionel Curtis organized numerous “closer union socie¬

ties,’’ forming a network over all of South Africa. Their object

was to familiarize the people with the problems connected with the

union. When the convention was at work these societies published

a magazine. The State, which discussed the questions then upper¬

most in the minds of the people. No particular form of union was

advocated although Mr. Curtis and the majority of his followers

had a strong leaning towards a legislative union.®®

In a solemn message issued on the eve of the convention. General

Botha appealed to the people of South Africa. “South Africa

has its opportunity now,” said he, now prime minister of the

Transvaal, “and I expect South Africa to do its duty * * * I

expect the result of the Convention to include unity, and there-

The Times, June 5, 1908.

^•Ibid., May 14, 1908.

83 Hansard, 4th series, CLXXXVIII, cols. 1215-1295. Debate, May 13, 1908.

®«See ibid., CXCIII, col. 1268; CXCIV, cols. 1609, 1610.

8“ Edinburgh Review, COX, p. 12.

Knaplnnd — A Study in British Colonial Policy

19

from to arise and develop a happy, prosperous, strong and healthy

nation. ’

These appeals were apparently necessary in order to arouse

the colonists, many of whom seemed quite apathetic. The Times^

correspondent feared lest disagreements among the unionists might

“wreck the whole scheme in presence of the unenthusiastic

many. ’

Fortunately, the difficulties were overcome and the constitution

for a united South Africa came into existence. This was affected

by a series of compromises on important issues, but without them

the attempt would surely have failed. Breadth of view and a

willingness to sacrifice unessentials characterized the attitude of

both Britons and Boers during the critical months in 1908 and

1909.®® Most remarkable was perhaps the spirit of loyalty towards

the empire manifested by such men as ex-president Steyn and

Generals Smuts and Botha. Without the aid of these great Boer

leaders the task could never have been accomplished.

The attitude of the high commissioner. Lord Selborne, remained

studiously fair and correct. Loyal to his conviction that the

problem was one for the people of South Africa to settle in their

own way, he proved helpful and courteous without attempting to

meddle or volunteer advice. The home government was likewise

friendly and sympathetic, following strictly a policy of “hands

off.”

The constitution as completed met with general approval in

England.®^ The Times described it as “a political achievement of

which the statesmen who modelled it and the people who endorsed

it have every reason to be proud of.”®® A delegation from South

The Times, Oct. 12, 1908.

Ibid., Oct. 5, 1908.

Prominent among the Boer representatives at the constitutional convention were,

ex-President Steyn and generals Botha, Smuts, De Wet, de la Eey, and Herzog. These

worked side by side with Sir Starr Jameson, Sir G. H. Farrar, and others who had

taken a leading part in the controversies which preceded and to some extent caused the

outbreak of the Boer War. A number of books have been written on the situation in

South Africa during the unification period, but no authentic source of information exists

for the actual work of the convention. The nearest approach to a contemporary account

of what happened behind the closed doors when the future constitution was debated is

Sir Edgar Walton’s The Inner History of the National Convention. The author was a

prominent member of that body, but the book itself is written partly from memory and

partly from rather incomplete notes; and it cannot be considered a full and complete

account of what took place.

For its reception in South Africa and the attitude of the separate states to it see

Worsfold, The Union of South Africa, p. 128; Walton, History of the Convention, pp.

321-328.

Leading Article, July 26, 1909.

20 Wisconsin Academy of Sciences, Arts, and Letters.

Africa, headed by Mr. Schreiner, tried in vain to persuade Parlia¬

ment to amend the clause in the constitution which barred the

colored from seats in the Union parliament. A few friends of

the natives, prominent among whom was Sir Charles W. Dilke,

supported this plea.®^ It was clear, however, that the “colour-bar’’

represented the wishes of an overwhelming majority of the whites

in South Africa, and the imperial government refused to accept

amendments to the completed document. The leaders of the two

great parties gave the constitution their whole-hearted support.

Mr., now Earl, Balfour characterized the Union as unexampled in

history; and the prime minister, Mr. Asquith, warned the House

of Commons “not to wreck this great' work * * * of freedom

and reconciliation.”®^ It was finally passed without a division.

The union of South Africa was officially announced by royal

proclamation of December 2, 1909 ; and May 31, 1910, the eighth

anniversary of the peace of Vereeniging, was fixed as the day of

union. General Louis Botha became the first prime minister and

Lord Gladstone, a son of the great Liberal leader, was appointed

as the first governor-general of the Union.

Thus was accomplished what Sir George Grey, Lord Carnarvon,

Cecil Rhodes, and others had attempted to affect during the latter

half of the nineteenth century. These early attempts were, no

doubt, premature, but lessons were taught which proved valuable

in the final effort. And it should also be remembered that without

the Boer War South Africa could not have become united. Strong

as were the economic and social forces favoring a union, an inde¬

pendent Transvaal was largely unaffected by them as long as its

natural outlet to Delagoa Bay remained accessible, and labor for

the mines could be recruited in Portuguese territory. Likewise, it

appears certain that without the aid of the young Britons trained

in the neo-imperialistic school of Chamberlain and Lord Milner

the union could not have come into existence at this early date.

The views in regard to federation presented by the members of

the “Kindergarten” were shared by their superiors. Conservatives

as well as Liberals both in South Africa and in England. They

labored for an end ardently desired by the imperial government

while employed by it in the crown colony administration of the

two new dependencies. While no evidence is available to show that

Stephen Gwynn and Gertrude M. Tuckwell, The Life of the Right Honourable Sir

Charles W. Dike (2 vols., London, 1917), II, pp. 375, 376.

The Parliamentary Debates, 5th series. House of Commons, IX, cols. 1006, 1564.

Knaplund — A Study in British Colonial Policy

21

Mr. Lionel Curtis and his friends worked under orders from

Downing Street, their eiforts must have heen noticed and approved

by the colonial office. Carnarvon’s policy of dictation, slightly

modified by crude attempts at persuasion, had failed; it was the

part of wisdom to abandon the former and evolve more finesse in

the use of the latter.^^ The presumption appears well-founded that

the imperial government was at least indirectly an important factor

in stimulating, during the period 1901-1909, that federation senti¬

ment which more direct methods had failed to arouse thirty years

earlier. But the sentiment itself could not have prevailed if great

centripetal forces had not existed and the Boer leaders had not

identified themselves with the cause of union.

Of considerable interest, in this connection, is the following observation by Sir Wil¬

frid Laurier: “At each imperial conference,’’ says Sir Wilfrid, “some colonial leader

was put forward by the imperialists to champion their cause. In 1897 it was obvious

that they looked to me to act the bell-wether, but I fear they were disappointed. In

1902 it was Seddon; in 1907, Deakin; in 1911, Ward.” Oscar Douglas Skelton, Lif*

and Letters of Sir Wilfrid Laurier (2 vols., New York, 1922), II, p. 342, note.

■■ ■ ■ .■ : "V'

: I'' *.!:;' , r ■/.'■ ■ 'y:''; : _

■■■ i . . j’ )’:>!. li;..;i;{

THE EEMOVAL OF THE WINNEBAGO

Louise Phelps Kellogg

The Black Hawk War sealed the fate of the Bock River hand of

Winnebago, and made their cession of that country inevitable.

The whites had already looked with covetous eyes upon this rich

and well watered region, especially on the western portion that

abutted on the lead mines. After the Winnebago uprising of 1827

this tribe was summoned the ensuing summer to Green Bay to

cede their portion of the lead mining region to the United States.

Too few of the chiefs were present to form a treaty, so a provi¬

sional agreement was made for the cession of the territory west and

south of the Blue Mounds and the Pecatonica River. This ar¬

rangement was confirmed by the treaty of Prairie du Chien in

1829, when the tribal lands were sold for an annuity of $18,000

annually for thirty years. The Indians were also to be provided

with three blacksmith establishments — one at Prairie du Chien,

one at Port Winnebago, and one on the waters of Rock River; they

were also to have a cart and two yoke of oxen at the portage.

Meanwhile two sub-agencies were erected for this tribe — one at

Port Winnebago, whose incumbent was the well known John H.

Kinzie. The other agent was Henry Gratiot, whose home at Gra¬

tiot’s Grove was outside the Winnebago territory; he thereupon

built an agency house on Sugar River, and there the blacksmith,

Maurice Mata, of whom Mrs. Kinzie speaks so favorably, had his

forge.^

Matters were in this condition until the Sauk uprising of 1832.

There can be no doubt that many of the Winnebago sympathized

with the hostiles, although the Rock River band was held to its

allegiance with the United States by the efforts of Gratiot and

Kinzie, ably seconded by Henry Dodge. In fact. White Crow,

Little Priest, and other Rock River chiefs did the whifes unre¬

quited service not only as guides and scouts, but as the agents in

the rescue of two captive white girls from Black Hawk’s band.

^Juliette A. Kinzie, Wau Bun (Caxton Club edition, Chicago, 1901), 327.

2

24 Wisconsin Academy of Sciences, Arts, and Letters,

Nevertheless, it was determined that the Winnebago must be

removed from a territory so close to the settlements; so in Sep¬

tember, 1832, their chiefs were summoned to Fort Armstrong,

where General Winfield Scott and Governor John Keynolds, of

Illinois, secured the cession of all their land south of the Fox-

Wisconsin waterway. In return for this cession the Winnebago

were granted the so-called neutral tract in Iowa ceded by the

treaties of 1830 with the Sauk and Foxes on one hand and the

Sioux on the other. This large tract was well watered and full

of game. Its drawback was the fact that it lay close to the habitat

of these fierce tribes still at enmity, and that its neutrality was far

from being assured. In compensation for this exchange of lands

the Winnebago were accorded an additional $10,000 annually for

twenty-seven years; they were to be served with a school and a

band of white farmers to teach them agriculture, and the Kock

Kiver band was to be supplied annually with 1500 pounds of to¬

bacco. In addition to the new cession in Iowa, the tribe still

possessed a large tract of territory north of the Wisconsin River

extending to the sources of Black River, sparsely settled by one or

two small tribal bands. The treaty further stipulated that ‘‘In

order to prevent misapprehensions that might disturb peace and

friendship between the parties to this treaty, it is expressly under¬

stood that no band or party of Winnebagoes shall reside, plant,

fish, or hunt after the first day of June next, on any portion of the

country herein ceded to the United States.

The winter following the Treaty of Rock Island was one of

great hardship for the Winnebago. The Black Hawk War had

prevented them from planting or harvesting any corn; game was

not plentiful, and numbers of the tribe literally starved and froze

to death. Mrs. Kinzie has given a graphic picture of this starv¬

ing time, and told of the distress of her husband and herself at

their inability to alleviate the condition of their poor Indian

“children.’’ “They would climb up on the outside [of the agency

house], and tier upon tier of gaunt, wretched faces would peer

in above, to watch us, and see if, indeed, we were as ill-provided

as we represented ourselves.” Corn had been purchased by the

government to supply the Winnebago ; but it had been detained at

Green Bay during the winter, on account of the frozen water¬

ways. Finally, in the spring the boats arrived and the famine

was for the time being relieved.^

® Charles J. Kapler, Indian Treaties (Washington, 1904), 345—351.

® Kinzie, Wau Bun, 363—365, 380—383.

Kellogg — Removal of the Winnebago

25

Now the time drew near when the major part of the tribe must

abandon its ancestral home, and remove all its villages either north

of the Wisconsin or to the new cession in Iowa. The government

would not specify which region should be occupied. There was,

however, rivalry among the government agents on this score.

Agent Joseph M. Street, at Prairie du Chien, was strongly in

favor of the Iowa location; he argued that it was merely a mat¬

ter of time when the tribe must relinquish all its lands east of the

Mississippi, and it would be best if it should be removed at

once to the west bank. On the other hand, Kinzie and Gratiot,

who best knew the Kock Eiver band, urged a location just north

of the Wisconsin. Street accused them of doing so in the interest

of the American Pur Company and from self-interest for their

sub-agencies. He was, however, intensely jealous of Kinzie, who

was much more popular than himself both with the whites and the

Indians. Street was a good man, with humane views, but not well

adapted to the exigencies of the frontier, and somewhat imprac¬

ticable in his plans for civilizing the red men.

Meanwhile the Fox River and Rock River bands were besieging

their agents with importunities to be allowed to remain on their

homelands until fall. If they might plant their accustomed fields

once more and gather the harvest, then they promised that they

would take their annuities and remove without making any diffi¬

culty. Both Kinzie and Gratiot favored this course, but were un¬

able to secure the coveted permission from either the governors of

Michigan or Illinois or from the commissioner of Indian affairs.

The white settlers of the Illinois frontier were terrified by the

recollections of the hostilities of 1832. John Dixon, of Rock River,

informed the army officers of the frontier posts that he believed

that the Winnebago were plotting with the Potawatomi for a

new uprising. Gratiot was hurried to his Sugar River agency

house ; there he met an officer from Fort Winnebago who had been

sent to test the temper of the tribesmen. The latter were terrified

by what they thought was a threat of war. They could not be

dissuaded from the belief that the whites were plotting to make

war upon them. They admitted to Gratiot that the Potawatomi

had sent them a message by their trader, Thibault, but insisted

that it was a peaceful message. Gratiot himself saw the wampum

and that it was tied with a green ribbon, signifying peace. The

message was an invitation to hold a great council at Turtle village

(on the site of Beloit). Here they were to have, wrote Gratiot,

26 Wisconsin Academy of Sciences, Arts, and Letters.

a great “Medicine Peast,” or a “Smoke to the Great Spirit/’

Then they hoped to secure permission to plant large fields of corn

in that locality and to postpone their removal until the autumn.

In the meanwhile, Colonel Henry Dodge, who had been on the

southwestern frontier organizing his regiment of dragoons, ar¬

rived at his home in Dodgeville. He had also been, during part

of the winter, at Washington, where he was hailed as the victor

over Black Hawk. He then proceeded to his Wisconsin home via

Dixon’s Ferry, where he heard the news of the Indian gathering.

He wrote on April 13th, to General Alexander Macomb of the

United States army, that he had seen Gratiot, who informed him

that the Winnebago were peaceful, and that they requested until

fall to make their removal. This, Dodge asserted, would by no

means to be tolerated by the frontier inhabitants. The tribesmen

must go before June 1st. He ordered two companies of his newly-

enlisted dragoons to march at once for the Illinois River and to

be ready when ordered to advance into the Rock River region.

Meanwhile Dodge met Kinzie, who assured him that the Winnebago

were much frightened and would accede to any proposals from

Dodge and their agents; that they were still in a state of semi¬

starvation and in great dread of the future.

Dodge decided to hold a conference with the principal chiefs

at the Four Lakes, and sent them word to meet him at the old

council ground on Fourth Lake, where he had talked with them

the previous May, and restrained them from joining Black Hawk.

April 28th was the day appointed. Dodge, Gratiot, and Kinzie

arrived with their escorts; all the principal chiefs of the Rock

River were there — Whirling Thunder, White Crow, Little Priest,

Little Black, Spotted Arm, White Breast, and others. The Indians

made their plea for time, which Dodge refused; they then asked

for help to remove their families, for wagons and oxen to portage

their canoes from the Rock River headwaters to the Wisconsin,

and for provisions to enable them to live. These requests the white

men granted. Dodge ordered his two companies of dragoons

under Captains Browne and Beekes to bring wagons from Illinois;

Gratiot ordered three wagons full of corn from Galena, and wrote

to Governor Porter that he would be in person at Four Lakes by

May 15th to superintend the distribution of rations.

Meanwhile the Winnebago of the lake of their name and of the

upper Fox River had begun their removal to the Baraboo Valley.

The Rock River Indians decided to settle on Sauk Prairie, where

Kellogg — Removal of the Winnebago

27

there had previously been a large Indian village. By the latter

part of May the dragoons reached Dodgeville, whence they pro¬

ceeded at once to Fourth Lake, where they built a camp on the

northwestern shore near a great spring which they named for the

spring Camp Belief ontaine. They also built another on the Wis¬

consin Kiver, naming it Camp Knox. Their instructions were to

observe a mild but firm attitude towards the removing Indians,

and to range up and down the streams to see that all the Indians

crossed the Wisconsin. On the fifteenth of May thirty lodges of

the tribesmen gathered at Four Lakes ready to cross to Sauk

Prairie. They held a grand medicine dance somewhere near the

present city of Madison, and performed ceremonies of removal.

^‘On the eve of their departure they extinguished all their old

fires, and kindled a new one procured by the friction of two sticks

of wood, which they ‘hoped would burn clear and make them

happy,’ ” wrote their agent Gratiot. He then gave them some

few presents, after which their canoes, wigwams, and effects, al¬

ready brought up the chain of the Four Lakes, were loaded on

to wagons and transported to the shore of Wisconsin Kiver. Man

Eater’s village was left behind as the chief himself was at Fort

Winnebago; Spotted Arm also was tardy in removing his band.

June 10th Dodge himself visited the camp on Fourth Lake and

went thence to Fort Winnebago, where he learned that sixty lodges

of tribesmen were still left on the eastern branches of Rock Kiver.

Kinzie had meanwhile sent out word that the annuities would

be paid July 1st at the agency house at the portage. Unfortunate¬

ly this payment was the occasion for a disgraceful orgy. Whisky

dealers brought in a vast quantity of liquor, and opened it on

private ground, where it could not be seized by the government.

Several Indians were killed in drunken rows, and most of the silver

paid them by the government passed into the hands of the liquor

dealers and traders, so that the Indians were worse rather than

better off for their annuities.

Dodge thought it necessary to keep the troops in the ceded ter¬

ritory, being certain that many of the Indians would return to

Rock River waters after the annuity payment. Whirling Thunder

and his band, encouraged by the traders John Dougherty, Oliver

Armel, and Stephen Mack, returned to their old village. Lieuten¬

ant T. B. Wheelock, of the dragoons, followed them to Sugar

River and arrested the white men and carried them and the In¬

dian band to the Fourth Lake camp. Dodge being notified by

28 Wisconsin Academy of Sciences, Arts, and Letters,

express, rode over from Dodgeville, released the traders, and sent

a troop of fifty dragoons to escort Whirling Thunder and his party

to Portage, and there set them across the river. This chief and

some others of his tribesmen then decided to settle in the neutral

ground. They went down the Wisconsin River to Prairie du

Chien, and by mid- July Whirling Thunder with two hundred and

fifty of the tribe, many of them from Lake Koshkonong and Turtle

River, had crossed into Iowa. The dragoons ’ camp on Fourth Lake

was kept up until October, when the troops had effectually cleared

the region of Winnebago stragglers.

It would be too long a story to attempt to follow the tribesmen

to their new homes. Those who went across to Iowa speedily re¬

turned because of an outbreak of hostilities between the Sauk and

the Sioux, in which they feared to be involved. After this, the

tribe felt badly crowded in the territory north of the Wisconsin.

‘^We are too many to live in so small a country,’’ complained

Chaetar to General Street at Prairie due Chien in 1834. Early in

that year Whirling Thunder sought out his old friend and agent

at Gratiot’s Grove, who has preserved for us his pathetic speech:

^‘Father — I have come to see you and get you to write a letter in

my name and in the name of my ,Rock River band of Winnebagoes.

We are tired of having no home — ^we are scattered all over the

country like wild beasts, and wish to unite in the spring, and build

a village and plant corn.

'‘Father Cass [Secretary of War] — I call on you particularly

because you know us, you have traversed our country and know

our habits, and our needs. . . .

“Father — you know better than we do that the land you gave

us west of the Mississippi, is occupied by the Sac’s and Foxes,

the Sioux’s and other tribes, and you know it [is] impossible for

us to go and live there, because all these natives are jealous of

us — it is useless for us to ramble about as we do. . . .

“Father — The Great Spirit has made the white and the read

[sic] man, the white he made more numerous than the red, and

gave them more sence they can read and they can write — it is

for that reason the Great Spirit created a distinction between

the two. The whites were made by the Great Spirit to take care

of the red people who are ignorant.” He then stated that the

Prophet was intriguing against the whites and was at his old

home on Rock River. Black Wolf, White Crow, and Little Priest

Kellogg — Removal of the Winnebago

29

were also known to have visited their old homes during the winter

of 1833-34.

So they came back — the dispossessed — ^year after year. White

pioneers tell of the long trains of Indian visitors that used to

come each summer to their old Rock River homes, to fish once

more in the beloved waters, to stand once more beside the effigy

mounds of their clans. The dispossessed make a sad picture, yet

an inevitable one. The Great Spirit had willed that this valley

of the Rock should be no longer a wilderness, haunted by bar¬

barians, but the home of a great civilization. Let us, however, in

our plenty and prosperity prove our humaneness by giving now

and then a look into the past and a sigh of sympathy for the dis¬

possessed red man, who loved the woods and streams and lakes

of his ancestral home with a deep and abiding affection. “This,’’

said one of the Winnebago tribe to the writer, “is the home of my

spirit.” From this home they were forced to go in the early

summer of 1833, because of the unreasoning terror of the frontier

settlers, and by the stern orders of the officers of the United

States government.^

* This paper is based almost wholly upon the documents in the Indian office at Wash¬

ington, copies of which are in the State Historical Library. Dodge’s letters of this

period are printed in Iowa Historical Record, v; 337—361; vi; 391—422, 445—467. See

also a trooper’s experience, in TTfs. Hist. Colls., x; 231—234. Mrs. Kinzie’s valuable

record in Wau Bun closes just before the removal and the payment of 1833. See in¬

cidents connected with the repeated return of the Winnebago, in Wis. Hist. Colls., x;

258-259.

AUGQSTINE OF HIPPO qua PATRIOT

Robert K. Richardson

Historians differ widely as to St. Augustine’s attitude toward

his country and its misfortunes at the opening of the fifth century

after Christ. Some consider him so devoted to the Heavenly City

as to be indifferent to the sufferings of the Temporal City. Others

discover in him a sorrowing patriot of the usual and accepted

pattern. An intermediate view is that represented by Dr. Angus

in his valuable work, The Sources of the First Ten Books of Au-

gustine^s Be Civitate Dei. Dr. Angus holds that: ‘‘So far as the

testimony of Augustine’s writings is concerned, his attitude to the

fall of Rome and the state of the Roman Empire of his day was

neither of intense and deeply patriotic feeling, nor of heartless

indifference, though nearer to the latter than to the former. ” “To

say the least,” continues Angus, “he appears surprisingly calm

in the face of so terrible a calamity. Augustine’s pride in Rome

was centered in her achievements of the past, not in her present.

He was more of a Christian than a Roman.

A study of Augustine’s letters and sermons, as well as of The

City of Ood, directed to the relatively small point in question, but

controlled by consideration of the broader matter of his general

outlook on God and the World, suggests that an eclectic estimate

of his position is closer to truth than any of the three views men¬

tioned. This estimate may be subsumed in four propositions: (1)

Augustine was fond, even proud, of the Empire; (2) he took a

lively interest in the fortunes of the state; (3) not considering the

situation hopeless, he bewailed this situation less than had he been

endowed with insight more prophetic; and (4) the character of

his Neo-Platonic Christianity lent moderation of grief alike to

his thought and to his rhetoric.

Augustine was fond, even proud, of the Empire.

It is not unnatural to regard Augustine’s attitude toward the

Roman Empire as basically prejudiced by a theory that the state

^ S. Angus, The Sources of the First Ten Books of Augustine’s De Civitate Dei. 1906.

275. Gf. ibid., 64—75 for quotations representative of one or the other of the types of

view mentioned in the text.

32 Wisconsin Academy of Sciences, Arts, and Letters,

originates in sin.^ In reality he assigns no such origin to the

state. The state, for Augustine, is an entity quite neutral in

character, belonging to the sphere of things temporal to be sure,

but practically adherent to the City of God or to the Earthly City

as its rulers yield their conduct’s allegiance to the one or to the

other. Only when justice is disregarded do kingdoms become

great robberies.”^ The state qua state is the handiwork of

Providence : Prorsus divina providentia regna constituuntur

humana.^ No political theory of the state, therefore, prejudices

Augustine against his country.

Nor is his affection cooled by that country’s idolatrous past.

He finds in its pagan heroes models for the citizenry of the King¬

dom of God and yearns for the eventual merging of the Common¬

wealth itself with the Civitas Dei, The great Komans of the past,

he relates, “despised their own private affairs for the sake of

the republic, and for its treasury resisted avarice, consulted for

the good of their country with a spirit of freedom, addicted neither

to what their laws pronounced to be crime nor to lust. By all

these acts, as by the true way, they pressed forward to honors,

power and glory; they were honored among almost all nations;

. . . and at this day, both in literature and history, they are

glorious among almost all nations.”® Naturally Augustine does

not forget that his country’s ancient worthies were “overcome

by love of fatherland and ardent desire for praise.”

Vincit amor patriae laudumque immensa cupido.

Haec sunt duo ilia, lihertas et cupiditas laudis humanae, quae

ad facta compulit miranda Romanos:^ and of course for the saint

of Hippo at the opening of the Middle Ages cupiditas laudis

humanae is a motive even more despicable than it was still felt

to be at the close of the medigeval period by his vacillating fol-

2 Cf. Harnack, History of Dogma (tr.), V, 153.

2 De Civitate Dei (ed. B. Dombart, 1863), IV, 4.

^ Ibid., V, 1. In general, cf. Mausbach, Die Ethik des Heiligen Augustinus, 1909, I,

332-333: “Verkorperung des Weltreiches ist der heidnische, auf Vergdtterung des

Kreatiirlichen ruhende Staat, nicht der Staat als soleher. Der letztere gehort wie die

Giiter der Menschennatur und der Familie zu einem Mittelbezirke zwischen den beiden

Gegensatzen, dessen sich beide bedienen und bemachtigen miissen.”

° De Civ. Dei, V, 15 (tr. Dods).

^ Ibid., V, 18. Cf. ibid.. Preface: “. . . iam . . . videamus, qua causa Deus, qut

potest et ilia bona dare, quae habere possunt etiam non boni ac per hoc etiam non felices,

Romanum Imperium tarn magnum tamque diuturnum esse voluerit.” The utility of the

temporal to the celestial peace is set forth in De Civ. Dei, XIX, 13, 14, 16, 17. It is

worth noting that Augustine mentions a befitting synchronism between the founding:

of Rome and the beginning of Israelitish prophecy {ibid., XVIII, 27).

Richardson — Augustine of Hippo qua Patriot,

33

lower, Petrarch. Notwithstanding, the seekers of the City of God

may find spur and exemplar in these seekers of temporal welfare

and human praise : ‘ ‘ Let us consider, ’ ’ urges the saint, ‘ ‘ how

great things they despised, how great things they endured, what

lusts they subdued for the sake of human glory, who merited that

glory, as it were, in reward for such virtues ; and let this be useful

to us even in subduing pride, so that, as that city in which it has

been promised to us to reign as far surpasses this one as heaven

is distant from the earth . . . the citizens of so great a

country may not seem to themselves to have done anything very

great if, in order to obtain it, they have done some good works or

endured some evils, when those men for this terrestrial country

already obtained, did such great things, suffered such great

things.”^

How better, then, might Augustine express his patriotic good

will toward his earthly country than by exhorting it to become

a member of his celestial country? One of the most eloquent

passages of the entire Civitas Dei is precisely such an invitation:

Nunc jam caelestem arripe, pro qua minimum lahorabis, et in ea

veraciter semper que regnahis. Illic enim tibi non Vestalis focus ^

non lapis Capitolinus, sed Deus unus et verus

nec metas rerum nec tempora ponit,

Imperium sine fine dabit,^

An identical thought is voiced in one of the sermons : Manet civitas

quae nos carnaliter genuit, Deo graiias. [No cold patriotism!]

TJtinam et spiritualiter generetur, et nobiscum tr unseat ad aeter-

nitatem.^

Augustine took a lively interest in the fortunes of the state.

The sermons and correspondence, alike, of St. Augustine, evince

an anxious interest in contemporary political conditions, and,

notably, in the Visigothic invasion. The letter to Italica, of about

’’Ibid., V, 17 (tr. Dods). At the end of the chapter Augustine finds a shadowy type

of the Eternal Country in “asylum illud Romuleum, quo multi tudinem, qua ilia civitas

conderetur, quorumlibet delictorum congregavit inpunitas.” This is quite in the spirit

of Dante’s De Monarchia. Of. also Epistula CXXXVIII, art. 17, in Patrologia Latina,

XXXIII, 533, and Goldbacher, Corpus Scriptorum Ecclesiasticorum Latinorum, XLIV,.

144-145.

» De Civ. Dei, II, 29. Of. ibid., II, 19, 21 and IV, 3, 4.

® Sermo CV, 7 in Pat. Lat., XXXVIII. The next sentence is : “Si non manet civitas

quae nos carnaliter genuit, manet quae nos spiritualiter genuit” and the chapter con¬

cludes with a discussion of the prophecy: “Exsurget gens super gentem, et regnum super

regnum.”

34 Wisconsin Academy of Sciences, Arts, and Letters.

the time of the sack by Alaric, complains that the lady has not

kept him informed of the Italian situation: “To this last letter,

just now received, I lose no time in promptly replying, because

your Excellency’s agent has written me that he can send my letter

without delay to Kome. By his letter we have been greatly dis¬

tressed, because he has [not] taken pains to acquaint us with the

things which are taking place in the city or around its walls, so

as to give us reliable information concerning that which we were

reluctant to believe on the authority of vague rumors. In the

letters which were sent to us previously by our brethren, tidings

were given to us of events, vexatious and grievous, it is true, but

much less calamitous than those of which we now hear. I am

surprised beyond expression that my brethren the holy bishops

did not write to me when so favorable an opportunity of sending

a letter by your messengers occurred, and that your own letter

conveyed to us no information concerning such painful tribula¬

tion as has befallen you, — tribulation which, by reason of the

tender sympathies of Christian charity, is ours as well as yours.

I suppose, however, that you deemed it better not to mention these

sorrows, because you considered that this could do no good, or

because you did not wish to make us sad by your letter. But in

my opinion, it does some good to acquaint us even with such

events as these: in the first place, because it is not right to be

ready to ‘rejoice with them that rejoice,’ but refuse to ‘weep with

them that weep;’ and in the second place because ‘tribulation

worketh patience. ’ . . . Far be it, therefore, from us to refuse

to hear even of the bitter and sorrowful things which befall those

who are very dear to us!”^®

If the just quoted letter imply a personal rather than a general

sympathy, the sermons reveal Augustine’s attitude toward the suf¬

ferings of the commonwealth as a whole. However redolent of

other-worldliness, the sermons numbered CV and CCXCVI in

Migne^^ are thus redolent only as are the passages therein borrowed

from the Book of Job. As the ultimate clinging to God of the

man of Uz is the expression and fruit of the deepest anguish of

soul for the loss of sons and daughters, of wealth and neighborhood

repute, so St. Augustine’s recourse to God and to divine consola¬

tion suggests the agony of heart caused by the miseries of his

country. “Be the world prosperous, be the world overturned: ‘I

XCIX, in Goldbacher, op. cit., XXXIV and Pat. Lat., XXXIII. Tr. Cunning¬

ham.

uPat. Lat. XXXVIII.

Richardson — Augustine of Hippo qua Patriot. 35

will bless the Lord, ’ who made the world. Utterly will I bless Him.

Be it well according to the flesh, be it evil according to the flesh,

‘ I will bless the Lord at all times : His praise shall continually be

in my mouth ! ’ . . . ‘ The Lord gave and the Lord hath taken

away: as the Lord pleases, so is it done: blessed be the name of

the Lord.’ And similarly with the other sermon: ‘‘The body

of Peter, men are saying, lies in Kome; in Rome lies Paul’s body,

in Rome the body of Lawrence, in Rome lie the bodies of other holy

martyrs- — and Rome is wretched, Rome is devastated, afflicted, de¬

stroyed, burned. So numerous are death’s slaughters, by famine,

by pestilence, by sword: where are the shrines [memoriae^ shrines,

memories — an apparent play on words, quite Augustinian] of the

Apostles? What sayest thou? Lo, this is what I said: Rome

suffers so many evils, where are the shrines of the Apostles? They

are there, but they are not in thee. Would they were in thee,

whoever thou be who sayest such things, who art thus void of com¬

prehension, who, called by the Spirit, art wise after the flesh!

. . . Be patient, the Lord wills it. . . . Behold the Lord,

thy God, behold thy Head, the example of thy life; harken to thy

Redeemer, thy Shepherd: ‘0 my Father, if it be possible, let this

cup pass from me. ’ Consider Row He shows a human will and

[yet] forthwith changes aversion to obedience — ‘Nevertheless not

as I will but as Thou wilt. Father.’ In such threnodies, at

least, Augustine exhibits an emotion which is more than Neo-

Platonic disinterestedness or monkish contempt!

On occasion, moreover, this quietistic and ascetic mystic exhibits

the most earnest interest in practical measures for the suppression

of danger to the integrity of the state and of society, even to the

extent of according public security definite precedence over the

claims of private renunciation. In evidence is a letter to Count

Boniface upbraiding this official for inefficiency in office. The

count is reminded that he had inclined to enter the monastic life

but had been dissuaded by the joint advice of Alypius and Au¬

gustine, who, instead, had recommended adherence to continency in

private life, but a continued connection with civic and military

affairs. Boniface is now not only compromising his spiritual wel¬

fare by a second marriage — and that, too, with a lady who has

been an Arian heretic^ — ^but is proving recusant to his military

trust. “But what shall I say,” complains Augustine, “of the

“ Sermo CV.

Sermo CCXCVI.

36

Wisconsin Academy of Sciences, Arts, and Letters.

devastation of Africa at this hour by hordes of African barbarians,

to whom no resistance is offered, while you are engrossed with

such embarrassments in your own circumstances, and are taking

no measures for averting this calamity! Who would ever have

believed, who would have feared, after Boniface had become a

Count of the Empire and of Africa, and had been placed in com¬

mand in Africa with so large an army and so great authority,

that the same man who formerly, as Tribune, kept all these bar¬

barous tribes in peace, by storming their strongholds, and menac¬

ing them with his small band of brave confederates, should now

have suffered the barbarians to be so bold, to encroach so far, to

destroy and plunder so much, and to turn into deserts such vast

regions once densely peopled! Where were any found who did

not predict that, as soon as you obtained the authority of Count,

the African hordes would be not only checked but made tribu¬

taries to the Koman Empire! ... If these benefits, though

earthly and transitory, are conferred on you by the Roman Em¬

pire, — for that Empire itself is earthly, not heavenly, and cannot

bestow what it has not in its power, — if, I say, benefits are con¬

ferred on you, return not evil for good. . . These, no

doubt, are the words of a mediaeval transcendentalist, but they

are hardly the language of a luke-warm patriot nor of an ill-bal¬

anced fanatic.

Augustine misjudged the seriousness of the contemporary situation.

As everyone knows, the rhetoric of the great church father is

frequently characterized by a refrain of weariness and woe: Con-

cutitur mundus, excutitur vetus homo; premitur caro, liquescit

spiritus.^'^ To translate is to lose the sob of it. Perit mundus,

senescit mundus . . . lahorat anhelitu senectutis.^^ Notwith-

Ep. CCXX, in Goldbacher, op. cit., LVII, and Pat. Lat., XXXIII. Tr. Ounningbam.

Of. Ep. COXXVIII, to bishop Honoratus, advising the clergy to remain by their flocks

in time of danger. The advice rests on distinctly religious grounds but illustrates,

none the less, the active and practical side of Augustine’s genius (Goldbacher and Pat.

Lat., ut sup.) Of., also, Joseph McCabe, St. Augustine and His Age, 1903, 487 et seq.

Pew first class biographies are less sympathetic in their treatment of their subjects.

^ Sermo CCXCVI, ut sup.

isSermo LXXXI, in Pat. Lat., XXXVIII. Of. Sermo CCXCVI; “Audistis fratres,

simul audivimus: Erunt bella, erunt tumultus, erunt pressurae, erunt fames. Quare nobis

ipsis contrarii sumus, ut quando leguntur credamus, quando implentur, murmuremus?

Sermo CV : Virgil is made to say that he placed the famous lines : His ego nec metas

rerum, nec temper a pono; imperium sine fine dedi, into the mouth of Jove as not be¬

lieving in their truth, and to affirm that his true views are to be found in the Georgies:

non res Romanae perituraque regna. Ep. XCIX (to Italica) : omnes autem nos dominus

consolatur, qui et haec temporalia mala praedixit et post haec bona aeterna promisit.

Ep. CXXVII, in Goldbacher, XLIV and Pat. Lat., XXXIII: Labores et pericula et

.exitia huius transitoriae vitae. Sermo XI.

Richardson — Augustine of Hippo qua Patriot.

37

standing, the attentive auditor is again and again conscious of an

iterated non desperandum. As will be explained directly, this

is in part due to the Augustinian philosophy; in part, however,

it is due to an inadequate estimate of the seriousness of contempor-

rary conditions. As often before, so now, the state is being bet¬

tered, not destroyed, by its afflictions. . . the Koman Em¬

pire is afflicted rather than changed, — a thing which has befallen

it in other times also, before the name of Christ was heard, and

it has been restored after such affliction, — a thing which even in

these times is not to be despaired of. For who knows the will of

God concerning this matter The sack of A. D. 410 is perhaps

the worst of Romeos misfortunes — Augustine is by no means sure

of it^®— -but the chastisement is after all a merciful one.^^ ‘‘Be¬

hold,” he exclaims in one of his discourses, it is said “Rome is

perishing in the times of the Christians: perchance Rome is not

perishing; perchance it is being scourged, not cut off: chastized,

not destroyed. Perchance Rome perishes not provided Romans do

not perish. For they will not perish if they shall praise God:

they will perish if they shall blaspheme. For what is Rome, if

not Romans f ’ The most remarkable example of this minimizing

of the catastrophy of 410 is the Sermo de Urhis Excidio^^ which

credulously cites the escape of Constantinople from a marvelous

manifestation of divine wrath during the reign of Arcadius and

maintains that the situation of 410 is analogous; “By the hand,

therefore, of an amending God is that city receiving correction

rather than destruction. Like a servant who, knowing the mas¬

ter’s will, does deeds worthy of stripes, it shall be beaten with

many blows.”

Both in his apologetic, or quasi-apologetic, writing, as in the

above sermon, and in his familiar correspondence, Augustine treats

the disasters of the day as casual and not as final. Nor is it proper

to ascribe this to apologetic requirements.^^ So thorough-going a

De Civ. Dei, IV, 7 {tr. Dods). Cf. nn. 8, 9 sup. The passage cited is written 415:

A. F. West, in Angus, op cit., 60.

Sermo CCXCVI, Pat. Lat. XXXVIII : Sed plus inquiunt, plus vastatur modo genus

humanum. Interim considerata praeterita historia, salva quaestione, nescio utrum plus.

Sed ecee sit plus: credo quia plus.

De Civ. Dei, V, 23.

Sermo LXXXI, Pat. Lat. XXXVIII. Cf. Sermo CCXCVI: after all, Rome has been

burned twice before; Modo te quid delectat contra Deum stridere, pro ea quae consuevit

ardere ?

21 Pat. Lat., XL.

22 Cf. Dill, Roman Society in the Last Century of the Roman Empire, 2nd ed., 1905,

313-314, 70, n. 4. (On Orosius).

38

Wisconsin Academy of Sciences, Arts, and Letters.

pagan as Rntilius Namatianus took even a rosier view: and, in¬

deed, the sack by Alaric, in itself considered, appears to have

been, in reality, a rather tame aJffiair.^^ Surrounded by angry

fugitives from the capital,^^ Augustine kept his head, appraised

the situation as best he might,^^ avoided undue minimizing of the

evil, and equally shunned its exaggeration.^® That his estimate of

conditions was too conservative is due, not to unpatriotic aloof¬

ness, but to the lack of perspective inherent in a contemporary

position.

Neo-Flat onic Christianity lent moderation to Augustine’s grief and

to its expression.

The reaction of the bishop of Hippo to the history of his time

was doubtless largely, perhaps chiefly, the manifestation of his

religious philosophy, a philosophy based, in its turn, as much on

the harrowing experimentations of his soul as on a detached and

placid Neo-Platonic metaphysic. His poise and self-possession are

less apathy and insensibility than the behaviour of a prophet who

sees God in the affairs of men and the affairs of men subsidiary

to the eternal and loving purposes of God. Augustine is at once

a pessimist and an optimist, and the firmness of his glance upon

“the portentous events around him,’’ to employ the phrase of

Ozonam,^^ is due less to a balancing of the two tendencies than

to a system which preserves each entire and manages harmoniously

to blend the two apparent antitheses.

It is quite correct to describe St. Augustine as a thorough pessi¬

mist. The human will is, for him, since the Fall, congenitally dis¬

eased; the phenomenal world possesses but secondary existence;

nature may better be discerned in the mind of God than by direct

observation, and investigation of nature, in and for itself, is but

soul-damning curiosity; our lives are dreary and arduous pil¬

grimages to a better land ; imperfection is the necessary concomitant

of all existence short of that of Deity itself. To be gripped in this

view of the world is in advance to possess an antidote to the worst

Dill, op. cit., 309—311. H. F. Stewart, in Cambridge Medieval History, I (1911),

575-576.

2* Dill, op cit., 62-63.

Gf. Orosius’s blind partizanship. Dill, op. cit.

2“ For emphasis on the darker aspects of contemporary life, vid. : Sermo CCXCVI in

Pat. Lat., XXXVIII; Sermo XI, ibid.’, Ep. XCIX and CXXVII, in Goldbacher, op. cit.,

XXXIV; and Sermo LXXXI, in Pat. Lat., XXXVIII.

^ A. Frederic Ozanam, History of Civilization in the Fifth Century (tr. A. C. Glyn),

I, 23.

Richardson — Augustine of Hippo qua Patriot, 39

of temporal misfortune. If (Gregory the Great, strengthened by

what he could grasp of this philosophy, might not be carried off

his feet by the flood of evils of the latter portion of the sixth cen¬

tury, how much less might Augustine be moved by the lesser in¬

undation of the early fifth !

On the other hand, from his very pessimism Augustine squeezes

a cheerful and optimistic view of life. Positive though it be as

an experience,^® evil is, in its essence, negative. The only truly

evil thing is evil will, and evil will, itself, as will, is good, for the

good God could form it only like Himself. The creative Provi¬

dence, forseeing in its own eternity the necessary imperfections

of temporal creatures made from nothing, plans the proper al¬

lowances and provides the appropriate compensations. The darker

the background, the brighter the foreground : the sharper the con¬

diment, the more piquant the sauce. The great and final Judg¬

ment will explicate and justify those lesser and daily judgments

which perturb and perplex the human heart. The ills of life are

incidents in the war of the two Cities. In His own time God will

segregate the realm of Satan, and the City of God, centered ever

in its Pounder, be seised of repose unchanging and eternal peace.^®

Augustine, therefore, disciple of Plotinus and Neo-Platoniser of

Christianity, is not unpatriotic when he comforts his contemporaries

by an attempted valuation of temporal trials in terms of eternity.

In summary, then, it may be affirmed that one^s appreciation

and estimate of St. Augustine ^s attitude must necessarily be af¬

fected by one own^ inmost and deeply private reaction to that

‘ ^ other- worldliness ” and mysticism natural to historical Chris¬

tianity. For the writer, the truth has best been expressed in two

mutually supplementary statements of De Pressense: ‘^We feel

that Christian as he is, he remains still a citizen, ’ ’ and ‘ ‘ He mourns

. . . but his tears do not conceal from him the destinies of the

City of God.”««

Gf. Augustine’s Confessions, Books VII, VIII. Paul Elmer More, Shelburne Es¬

says, 6th, Series, Augustine, 88-89: “It [the Tolle, lege; tolle, lege scene] summoned

him from the intellectual consideration of evil as a negation of good to the conviction

of sin as something for which he was morally and terribly responsible. . . . Evil was

the deliberate setting apart of the human will from the divine will, the voluntary separa¬

tion of the soul from the source of life.”

For an excellent study of the Neo-Platonic character of Augustine’s theology, vid.

L. Grandgeorge, Saint Augustin et le Neo-PIatonIsme, 1896,

Quoted by Angus, op. cit., 64-65.

MILTON AS A WRITER ON EDUCATION

Oliver M. Ainsworth

Milton's contribution to educational theory, although neatly

classified and apparently disposed of by several writers, still has

possibilities for the investigator. Oscar Browning tells of a rude

shock that his ideas on the subject once received, when he first

thought of reprinting Milton's Tractate Of Education. ‘‘One

of the senior masters at my school," says Browning, “set Milton

as a subject for a Latin theme to his division, and told his boys

that they were to prove that Milton, like Burke, went mad in his

old age. I had never heard of this idea before, and I asked the

master on what grounds it rested. He replied, ‘Did he not write

a crack-brained book about education, in his old age ? ' I concluded

that my scheme was useless, and gave it up." Fortunately, how¬

ever, Browning's discouragement was not final. Thanks to him

and to others, Milton's ideas are to-day somewhat better known.

And yet, if one may judge from the readiness with which some

educators dispose of Milton, their interest does not seem to have

penetrated much further than that of Browning's senior master,

more than seventy years ago.

Milton's direct experience in teaching lasted only a few years;

but the subject of education occupied his thoughts at frequent

intervals throughout his whole life, as one may see from the many

allusions to it in his writings. Even as an undergraduate at

Cambridge, Milton was interested in educational theory. In one

of his academic exercises,^ an address delivered before his fellow-

students and his instructors, he attacks the problem, vigorously

condemning the barren subtleties of scholastic philosophy, which

was then a prominent part of the curriculum, and contrasting it

with the pleasures of history, literature, and natural science, in

which he would have preferred to spend his time. His little treatise

on education appeared some twelve or fifteen years later (1644)

in the midst of his earlier writings on public reform. Another

notable reference to education is to be found in the tract. The

^ Masson, Life of Milton, 1 :281.

42 Wisconsin Academy of Sciences, Arts, and Letters.

Likeliest Means to Remove Hirelings out of the Church (1659).

Here Milton desires better schools in different parts of England

for the purpose of educating the clergy, without the necessity of

their attending the University.- Again, in The Ready and Easy

Way to Establish a Free Commonwealth, Milton’s last direct effort

in what he believed to be the cause of liberty, he has these remark¬

able words on the relation of education to representative govern¬

ment : ' ‘ To make the people fittest to choose, and the chosen fittest

to govern, will be to mend our corrupt and faulty education, to

teach the people faith, not without virtue, temperance, modesty,

sobriety, parsimony, justice; not to admire wealth or honor; to

hate turbulence and ambition; to place every one his private wel¬

fare and happiness in the public peace, liberty, and safety.”^

Sound education, therefore, is the very basis of Milton’s political

structure. Even in Paradise Lost, as a friend of mine at a neigh¬

boring university has recently pointed out, Milton’s thought was

occupied with education as well as with theology and ethics.^

These examples show that the idea of a better education was

by no means a Utopian scheme of Milton’s. He connected it with

practical matters which he believed to be of the highest importance.

Moreover, the Tractate Of Education, Milton’s principal expres¬

sion of his views on the subject, is not a casual or isolated essay.

In The Second Defense of the People of England there is a passage

where Milton, in defending himself from the personal attacks of

a foreign writer, gives rather an extended account of his own life

and works. After mentioning his writings on the reform of church

government and on marriage, he says: “I then discussed the

principles of education in a summary manner, but sufficiently

copious for those who attend seriously to the subject; than which

nothing can be more necessary to principle the minds of men in

virtue, the only genuine source of political and individual liberty,

the only true safeguard of states, the bulwark of their prosperity

and renown.”^ Thus Milton represents this work as merely in¬

cidental to his earlier writings in the cause of reform.

That Milton’s opinions were valued, by at least one of his con¬

temporaries, is evident, for the Tractate was written at the earnest

solicitation of a friend — a man of some note in the London of

^ Prose Works, ed. by St. John, 3:27.

s lUd., 2:126.

^ Murray W. Bundy, “Milton’s View of Education in Paradise Lost,” Journal of

English and Germ. Philology, 21:127.

Prose Works, 1:259.

Ainsworth — Milton as a Writer on Education. 43

that day. This was Samuel Hartlib, who was well known and

highly esteemed as a philanthropist and reformer. Of Prussian

birth and Polish-English descent, Hartlib had important friends

and correspondents on the Continent as well as in England. His

interests ranged all the way from the promotion of mechanical

inventions to a scheme of uniting the Protestant churches through¬

out Europe. Among them was the reform of schools. At the

time of Milton’s return to England in 1639, Hartlib was espe¬

cially active in spreading abroad the ideas of Comenius, the great

Moravian teacher. Milton expresses the highest regard for Hart¬

lib ’s character and service; and it is significant of the esteem in

which Milton was beginning to be held, that such a man as Hart¬

lib should so earnestly desire his opinion on education.

Nevertheless, there are two points in the Tractate Of Education

that may scarcely have been pleasing to Hartlib. In the first

place, Milton expresses but little regard for Comenius. Milton’s

practical sense would naturally lead him to take the more skeptical

view of the schemes of Pansophia or Universal Wisdom which at

this time were bound up in Comenius’ mind with the plan of edu¬

cational reform. And it is likely that, after his years of appli¬

cation to study, and his experience as a teacher, Milton not un¬

justly believed himself quite as competent to theorize on the sub¬

ject of education as the most of his contemporaries. At all events

he expresses entire indifference to Comenius’ writings.

Furthermore, one of the most prominent features of Milton’s

plan would be looked upon by Hartlib, and also by most modern

educators, as a serious limitation. Milton’s scheme, unlike that

of Comenius, is not intended for the children of all classes; and

nothing whatever is said about the education of girls. And yet,

from Milton’s words in The Commonwealth (quoted above) and

from Sonnets 10 and 11 and the eulogy on Queen Christina of

Sweden,® we know that Milton was by no means indifferent to these

wider interests. In the Tractate he was particularly concerned

with the training of leaders in Church and State, and he naturally

looked for them among the young men of the best families.

Now, another of Hartlib ’s interests was the founding of an

agricultural school; consequently, a glance at the reading-list in

Milton’s curriculum may have pleased him better. It includes

many of the authoritative works of the day on natural science and

its applications. That these books, as natural-scientific treatises.

^Ibid., 1:249-250.

44 Wisconsin Academy of Sciences, Arts, and Letters.

are now out of date, is no fault of Milton’s. Natural science in

the modern sense had then scarcely begun. The including of

writers on agriculture, geography, medicine, and natural history

must have appealed to Hartlib’s ‘‘practical” inclinations. At any

rate, it has given rise to a widely spread notion that Milton’s

academy was informational, rather than disciplinary, in method,

and vocational, rather than cultural, in purpose.

It is significant in this connection that the oldest edition of

Milton’s treatise to be found in the library of Beloit College is

bound in the same volume with Locke’s Thoughts Concerning Edu¬

cation, a work which is a main source of utilitarian doctrine. The

association of the two is not accidental. The editors give as the

reason for it, that the subject of Milton’s treatise “seemed more

in harmony with the topics discussed by Mr. Locke than with the

contents of any other volume in the intended series.”^

Charles Lamb in his essay. The Old and the New Schoolmaster ,

clearly strikes the note of criticism. ‘ ‘ The modern schoolmaster, ’ ’

says he, “is expected to know a little of everything, because his

pupil is required not to be entirely ignorant of anything. He

must be superficially, if I may say so, omniscient. . . . You

may get a notion of some part of his expected duties by consult¬

ing the famous Tractate on Education addressed to Mr. Hartlib.”®

Dr. Johnson, however, puts the case with greater emphasis, and

I may be pardoned for quoting his remarks at more length. In

discussing Milton’s school he says:

The purpose of Milton, as it seems, was to teach something more solid than

the common literature of the Schools, by reading those authors that treat of

physical subjects; such as the Georgick, and astronomical treatises of the

ancients. . . .

But the truth is, that the knowledge of external nature, and the sciences

which that knowledge requires, or includes, are not the great or the frequent

business of the human mind. Whether we provide for action or conversation,

whether we wish to be useful or pleasing, the first requisite is the religious

and moral knowledge of right and wrong; the next is an acquaintance with

the history of mankind, and with those examples which may be said to em¬

body truth, and prove by events the reasonableness of opinions. Prudence

and Justice are virtues and excellences, of all times and of all places; we are

perpetually moralists, but we are geometricians only by chance. Our inter¬

course with intellectual nature is necessary; our speculations upon matter are

voluntary, and at leisure. Physiological learning is of such rare emergence,

that one man may know another half his life, without being able to estimate

''Library of Education, Gray and Bowen, Boston, 1830.

* The Work of Charles and Mary Lamb, ed. by Thomas Hutchinson, 2 vols., Oxford,

1908, 1:536.

Ainsworth — Milton as a Writer on Education.

45

his skill in hydrostaticks or astronomy; but his moral and prudential character

imm.ediately appears.

Those authors, therefore, are to be read at schools that supply most axioms

of prudence, most principles of moral truth, and most materials for conver¬

sation; and these purposes are best served by poets, orators, and historians.^

This is rather an uncompromising statement of the case for

humanism ; and yet for this very reason it is wide of the mark as

a criticism of Milton. For Milton himself, above all things else,

was a humanist. His life was dominated by intellectual and spirit¬

ual ideals. He devoted himself, for the good of humanity, to one

of the noblest of human endeavors — the creation of beauty in art.

And the art to which he dedicated his great powers of intellect

and imagination was the art of poetry; poetry, too, not conceived

of as amusement or pastime, but inspired by the most vital prin¬

ciple in human nature — the religious instinct.

One who has read Milton’s works with attention and without

prejudice can hardly doubt that he was in fundamental agree¬

ment with Johnson on the question of what studies are really the

most ^‘practical.” The general substance of Johnson’s remarks

may be found in different passages of Paradise Lost,^^ and per¬

haps Milton himself, in deliberate prose, could not have stated his

principles more faithfully than Johnson has done.

How, then, has the conception risen that Milton’s scheme of edu¬

cation is materialistic? Simply through judging by the lists of

reading alone, without sufficient attention to Milton’s own explana¬

tion of his purpose and method.

Two definitions of education are an outstanding feature of the

treatise. Near the beginning Milton says: ‘‘The end then of

learning is to repair the ruins of our first parents by regaining

to know God aright, and out of that knowledge to love him, to

imitate him, to be like him, as we may the nearest by possessing

our souls of true virtue, which, being united to the heavenly grace

of faith, makes up the highest perfection. After discussing

the mistakes of the schools and universities, he says : “ I call, there¬

fore, a complete and generous education that which fits a man to

perform justly, skilfully, and magnanimously, all the offices, both

private and public, of peace and war.”^^

It is possible to see in these two definitions two distinct pur¬

poses, not wholly in accord with each other. The first is a religious

^ Lives of the Poets, ed. by Waugh, 1:73-74.

Paradise Lost, 7:111-130; 8:172-197; 12:553-587.

^ Prose Works, 3 :464.

^Ibid., 3:467.

46 Wisconsin Academy of Sciences^ Arts^ and Letters.

purpose, and may be humanistic; the second, considered by itself,

is almost certainly utilitarian. And yet it seems strange, in so

brief a work, that Milton should have had two purposes in mind

at all; and stranger still, that they should have been in conflict,

or even unrelated. Such is probably not the case. The true har¬

mony of Milton’s definitions may perhaps be most clearly shown

by the analogy of faith and works. ‘‘These two divisions,” says

Milton in his Christian Doctrine, “though they are distinct in their

own nature, and put asunder for the convenience of teaching, can¬

not be separated in practice.”^® Now, the basis of Milton’s edu¬

cational doctrine is character, both as an end in itself, and as the

foundation of true service. These two values of character may be

separately studied, but in practical life they are inseparable.

Milton had no interest in a “fugitive and cloistered virtue” that

never emerges in action. Furthermore, an education which does

not produce character cannot, in Milton’s estimation, prepare a

man for service. For instance, one might suppose that the most

important part of a statesman’s education would be political

science. But what does Milton say of the young men who leave

college to enter politics? “Others,” he says, “betake them to

state affairs, with souls so unprincipled in virtue and true generous

breeding, that flattery and court-shifts and tyrannous aphorisms

appear to them the highest points of wisdom. To be sure^ Mil-

ton includes political science in his curriculum; but he sets no

great value on science of any kind without right principles to

guide and control it.

We may once for all set aside the notion that the school was

to be vocational, for Milton expressly states that it is a place of

“general studies”^® in contrast to such special schools as those of

law and medicine. The fact of its military organization is more

important. Milton labored under no illusions on the subject of

war; bul he realized the ethical value of military training, as

well as its practical necessity.

Let us now glance at one or two interesting features of Milton’s

curriculum.

The chief medium of instruction in that day was the Latin

language; therefore the pupil’s first task was to acquire it. The

customary method of learning Latin in Milton’s time was, first,

the memorizing of many cumbrous rules of grammar, (often ar-

4:13.

3:466.

^°Ibid., 3:467.

Ainsworth — Milton as a Writer on Education, 47

ranged, for greater convenience, in rude verse) ; second, the prac¬

tice of Latin composition iu generous amounts, both in verse and

in prose. Some Latin authors, of course, were read ; but composi¬

tion predominated. The pupil began the process about the age

of ten, or even younger. If he survived it, he entered the uni¬

versity at the age, perhaps, of sixteen, with a fairly good com¬

mand of Latin vocabulary, and the ability to understand lectures

in the language, and to converse with his fellow-students in it.

The process, however, was not always successful, and never very

agreeable. Eoger Ascham^s treatise. The Scholemaster, nearly sev¬

enty-five years earlier than Milton ^s, was the outcome of a con¬

versation over the case of some boys who had run away from Eton

School for fear of a beating.^® One of Comenius^ greatest achieve¬

ments was in simplifying the method of teaching Latin. The prob¬

lem was a close parallel to our modern one of how to teach the

Freshman to write English. We are attacking it very much as

Milton’s contemporaries did — ^with unlimited quantities of theme¬

writing.

However, Milton’s views on this subject were revolutionary. He

insists on the principle that composition ought to be based on in¬

formation of some kind, and not painfully spun out of the student’s

inner consciousness. He calls it a ‘‘preposterous exaction” to

demand much writing of students until they have read widely in

good authors, and acquired a moderate supply of information as

well as a sense of style. Milton believed it possible to teach lan¬

guage chiefly through extensive reading — a belief shared by some

good teachers to-day. Moreover, he regarded language, not as an

end in itself, but merely as “the instrument conveying to us things

useful to be known.” Therefore, after teaching his students the

elements of Latin grammar, he led them rapidly through a con¬

siderable amount of reading, and deferred the composition until

a later part of the course. He followed the same method with

Greek, which, of course, as a humanist, Milton felt to be indis¬

pensable.

Next to be considered is the subject-matter of the reading.

It is chiefly to this that Dr. Johnson applies his criticism. The

first authors to be studied are the Latin writers on agriculture;

and between the ages, probably, of thirteen and sixteen, the pupils

are to read treatises in Greek and Latin on such matters as ar¬

chitecture, astronomy, geography, medicine, and natural history.

“Roger Ascham, English Works, ed. by Wright, Cambridge, 1904, p. 175.

48 Wisconsin Academy of Sciences, Arts, and Letters,

not forgetting to note the practical applications of these various

arts and sciences, or omitting to call in the aid of practitioners for

purposes of demonstration. Arithmetic and geometry had been

studied in the earliest year of the course.^^

All this looks somewhat as if Dr. Johnson were in the right.

And yet, while Milton’s curriculum incidentally imparts a great

deal of information, both curious and useful, his ultimate purpose,

even in the study of external nature, was to develop the mind

through contemplation. This purpose he makes very clear at the

outset, when he says that we cannot in any other way arrive so

clearly at the knowledge of things invisible as we can by studying

the visible creation.^® In one passage of Paradise Lost Milton com¬

pares Nature, or the entire order of the Universe, to a ‘‘scale,”

or ladder,

‘ ‘ Whereon,

In contemplation of created things

By steps we may ascend to God.”^^

And in his essay on The Reason of Church Government Milton

expressly states that the knowledge of God and of his true worship,

and what is infallibly good and happy in the state of man’s life,

what in itself evil and miserable, is the only high valuable wis¬

dom indeed.^®

Furthermore, the very multitude of subjects covered in so short

a time would make it impossible for the student to become very

proficient in any one art or science. The course is distinctly a

survey; but the reading of one or more unified treatises on each

subject, though not expected to make the student even an amateur,

would give him just what Milton desired to impart — an insight into

the spirit and method of natural science. A final circumstance,

of a kind to show that Milton looked upon natural science de¬

cidedly from the cultural point of view, is this: the study of ex¬

ternal nature is to be rounded off by reading those poets who treat

of it, and add to it what Wordsworth calls “the breath and finer

spirit of all knowledge, the impassioned expression which is in

the countenance of all science.

Prose Works, 3 :469.

3:464.

Paradise Lost, 5:507-512.

-uprose Works, 2:473.

Preface to the Lyrical Ballads, 1800; Poetical Works, ed. by Hutchinson, Oxford.

1913, p. 938.

Ainsworth — Milton as a Writer on Education, 49

From the study of external nature the pupils proceed, in the

next stage of the course, to the study of human ideals and institu¬

tions. Between the ages of sixteen and nineteen, they read the

standard treatises on ethics, economies, politics, law, theology, and

history both sacred and secular. These subjects are more in accord

with what are commonly known as the humanities. They are in¬

tended, through the study of man and his works, to give the pupils

a further insight into the purposes of the Creator, as well as to

acquaint them with the nature of practical affairs. Toward the

end of this stage, the greatest masterpieces of literature are read,

and an attempt is made to impart something of their true spirit

to the students.

In the last stage of all, the pupils are to learn the laws of con¬

struction that underlie every great type of writing; and now,

but not before, are they to be formally trained in the art of com¬

position, ''when they shall be thus fraught with an universal in¬

sight into things.”-^

From Milton’s use of natural science as a means of training

the mind and building character, it may be inferred that he had

no superstitious fear of its destroying religious faith. Indeed, in

the matter of scientific truth as opposed to civil or ecclesiastical

authority, Milton was on the side of scientific truth; for in visit¬

ing the aged Galileo, he defied the Inquisition. Milton, however,

believed that true science and true theology both are ultimately

rational; and he took no interest in the ingenious opposition be¬

tween them, which has puzzled some minds since the time, prob¬

ably, of Heraclitus.

At the same time, he realized that the functions of science and

of religion are distinct, and that neither one can do the work of

both. Science may lead us to a rational conviction of divine truth,

but moral power is supplied by faith alone. Accordingly, since

his chief object is the development of character, Milton takes care

from the very outset of his course to instill religious faith into the

minds of his pupils. "After evening repast, till bedtime,” says

he, "their thoughts would be best taken up in the easy grounds

of religion, and the story of scripture.”^® He probably chose the

evening as the time when the imagination is most active, and most

open to feelings of reverence and affection. The severity of the

master would be laid aside; and in after years the pupils would

Prose Works, 3 :474.

^Ibid., 3:469.

50

Wisconsin Academy of Sciences, Arts, and Letters,

cherish the impressions of these hours with a fondness and tenacity

that formal precepts never could evoke.

The Tractate Of Education is so rich in suggestion that one is

confined in a brief space to a few such outstanding features as

I have discussed. Perhaps Milton’s own words at the close of the

treatise, since they show that he felt the work to be suggestive

rather than final, may bring this paper to a fitting conclusion.

Thus, Mr. Bfartlib, you have a general view in writing, as your desire was,

of that which at several times I had discoursed with you concerning the best

and noblest way of education; . . . many other circumstances also I

could have mentioned, but this, to such as have the worth in them to make

trial, for light and direction may be enough. Only I believe that this is not

a bow for every man to shoot in, that counts himself a teacher; but will

require sinews almost equal to those which Homer gave Ulysses; yet I am

withal persuaded that it may prove much more easy in the assay, than it now

seems at distance, and much more illustrious; howbeit, not more difficult than

I imagine, and that imagination presents me with nothing but very happy

and very possible according to best wishes; if God have so decreed, and this

age have spirit and capacity enough to apprehend. 24

3:478.

FLOUNDERING IN MODERNITY

George C. Clancy

It is surely unnecessary to remind our contemporaries that they

are living in a modern age. Our times are reeking with “modern¬

ity^” whatever that means. To be modern apparently implies

escape from the dreary dullness of mid-Victorianism, the ennui of

the fin de siecle, and the shackles of faith in anything whatsoever.

Intellectually, it is a gay time to be on earth. For the generation

latest born, life has particular zest. “Bliss is it in this dawn to

be alive, but to be young is very heaven.” Youth plays saucily

with the gray-beards, and the older generation, for all its hoary

wisdom, is hard put to it to preserve its dignity — alas, it has

to defend its innermost shrines from sacrilege by its own children.

If we were inclined to sentimental mooning we could shed a few

tears in the grave-yard where lie buried sacred traditions of litera¬

ture and art, ardent hopes for democracy, and some of the holy

faiths of our fathers. Even our efforts at virtue and culture, into

which we Americans have thrown what we think is a pardonable

enthusiasm, are the object of the indulgent smiles of the intelli¬

gentsia. A recent article by a brilliant foreign critic refers to

our orgies of prohibition and suppression as unbelievable to the

civilized European, calls our reforms debauches of virtue, finds a

malady of intellectual anaemia in the varied departments of Ameri¬

can life, and scores the books of popular idealists as the namby-

pamby product of moral soothsayers. Even the time-honored

institution of marriage is under a hot cross-fire; the family, once

a centre of intense loyalties, gives evidence of losing its unity;

“Home,” as Robert Frost says, “home is the place where, when

you have to go there, they have to take you in.” Our democracy

as expressed in the House of Representatives at Washington, we

are told is a mess of incompetence and imbecility. H. L. Mencken,

writing on “Politics,” says that our Congressmen are, in the

overwhelming main, shallow fellows, ignorant of the grave matters

they deal with and too stupid to learn, with the intelligence of

the country newspaper editor, or the evangelical divine. As a

52 Wisconsin Academy of Sciences, Arts, and Letters.

civilization we appear to be in a parlous state, at least if one’s

view is colored for him by the modern prophet of pessimism.

Yet these clever and incisive writers of our decade are more

than masters of the telling phrase. There is ample evidence that

things are not as they were with our fathers. Everywhere cables

are slipping, moorings are lost; the thought of our age flounders

amidst the tide-rips, the cross and shifting currents of an un¬

charted sea. Denial, rebellion, daring experimentation, are at the

wheel, and the compass has evidently been thrown overboard.

Thomas Hardy ventures the opinion that it would not be at all amiss

if man should undertake the education of God as an effort toward

the general improvement of things.

Modernity has entered the classic halls of learning — there is no

doubt of that. Indeed, the once sacred precincts are proving

particularly fertile soil for the growth of the new idea. In the

better institutions, there is apparently little hostility on the part

of administration and faculty towards the intellectual insurgency,

wherever it is thoughtful and genuine. Unfortunately, college

youth often assume an easy cynicism which in its sublime ignor¬

ance would be impertinent if it were not so absurd. It flaunts

itself in the face of the instructor with such sophomoric assurance

and condescending tolerance that the teacher may well be congrat¬

ulated on his self-control in not wringing the necks of his pro¬

teges. But if the instructor has the grace of God in his heart to

stand this arrant cockiness, he has reason to welcome the condition

of which it is a symptom. It is the evidence nearly always of

intellectual alertness, a thing vastly to be desired in the student,

and all too frequently sought in vain. Cynicism and rebellion are

not diseases of youth to be treated pathologically. They are grow¬

ing pains, signs of healthy expansion in the body politic. “The

most hopeful thing of intellectual promise in America today, ’ ’ says

a daring modern writer, “is the contempt of the younger people

for their elders ; they are restless, uneasy, disaffected. ’ ’ However

much our civilization may resent this contempt, it can seek within

itself at least partial cause for its repudiation by the rising gen¬

eration. Too often has it wrapped its faith about with the trap¬

pings of sanctity, and sacrificed logic to the promptings of mere

sentiment. Striking as were the changes in the thinking of the

nineteenth century, we and our fathers have been curiously help¬

less in our efforts to throw off the inhibitions of tradition. Again

and again the cake of custom has not been broken up. The intrepid

Clancy-— Floundering in Modernity,

53

spirit of the pioneer in things of the mind has found its freest

expression in the realm of physical science. Professor James

Harvey Eobinson in his much-discussed book, ‘^The Mind in the

Making,” says* While we have permitted our free thought in

the natural sciences to transform man^s old world, we allow our

schools and even our universities to continue to incalculate beliefs

and ideals which may or may not have been appropriate to the

past, but which are clearly anachronisms now. For, Hhe social

science’ taught in our schools is, it would appear, an orderly pre¬

sentation of the conventional proprieties, rather than a summons

to grapple with the novel and disconcerting facts that surround

us on every side.”

Scholarship has apparently been at its best only where the

processes of thought do not concern themselves with human motives

or become intertwined with the heart-strings; if they are so in¬

volved, argument must make assurance doubly sure and take a bond

of fate. Not that our scholarship is ever in the position of abject

fear as it faces possible conclusions, but too often the wish is father

of the thought, or the bias of heredity and temperament is an

unconsciously determining influence. Who is there of us who

would not surrender in a moment and without a pang Newton’s

law of gravitation or the solar system of Copernicus ? In the lab¬

oratory of the true scientist there is never a fear for what lies

at the bottom of the crucible when the test is over. Were the

evidence to be found there to mean the denial of the hope of

immortality, there would be no moment of hesitation, nor a quiver

of the eye-lid. Is not truth in its purity, without compromise,

goal sufficient for man or God? Yet we pursue with trepidation

the paths that explore the fields of humanistic scholarship and

cling with hazy-eyed and fond devotion to the formulae of the past.

More than that, most of us^ — unconsciously, to be sure— are special

pleaders. We have a cause to promote, which is not truth but a

particular brand of truth in which we have come to believe. The

churchman today, for example, is far too likely to start with his

conclusions and work back to his premises. Thus he is frequently

driven to unconvincing and even specious argument to support

positions rapidly becoming untenable. His cosmic philosophy will

not tolerate the conception of a Godless universe or a denial of the

immortality of the soul. And the radical thinker, on his part,

is likely to be carried away by things in general — ^in art, literature,

politics, and religion ; he loses his mental balance, becomes a

54

Wisconsin Academy of Sciences, Arts, and Letters.

shouter of empty slogans. Emma Goldman — of all people ! — turns

state’s evidence on this point. ‘‘The average radical,” she says,

“ is as hide-bound by mere terms as the man devoid of all ideas.

‘Bloated plutocrats,’ ‘economic determinism,’ ‘class consciousness,’

and similar expressions sum up for him the symbols of revolt. His

hearing is dulled by the din of stereotyped phrases.” The world

is indeed rife with special pleadings by doctrinaires. We cannot

with confidence seek our truth in the printed organ of the socialist

party, the religious journal, or in the publications of the “wets.”

Such literature serves its purpose, which is essentially propagan¬

dist, but practically none of it is free from the taint of prejudice.

To make the cause prevail — this is the primary purpose, and argu¬

ment and data are selected and displayed to that end. It would

be foolish to seek for an unbiased presentation of truth in any

one of them.

In this intellectual chaos, it would not perhaps be unfair to

say that scholarship, in its pure and detached form, is about the

only agency engaged in a wholly disinterested search for truth —

the scholarship of teacher and writer as we find them in their

highest realization, the scholarship of the man working in the

field of the pure sciences, the scholarship of the discriminating

teacher of history, of the keen analyzer of our social unrest. In

this statement no reference is made to things of purely spiritual

content, except as they rest on scientific fact and so depend for

their convincing power on the truth of a cosmic philosophy. If

scholarship does occupy and can continue to maintain such a

unique position in a world of flux, its power as a guiding and con¬

trolling force is inestimable. If it can draw to itself the best

minds of the nation, men of intellect, poise, and devotion, men

capable of mental perspective and detached judgment, it' has a

high destiny before it. It would be the supreme court in the

world of the mind, judicial, wise, incorruptible. Such a court

would command respect and win confidence wherever its qualities

were recognized. The insurgency of college students has some¬

times troubled good faculty members. They look askance at the

welcome accorded at the universities to the preacher of heresies

in religion and political economies, at the applause which a stirring

apostle of radicalism wins in the college chapel. But if a college

faculty is worthy of the great traditions and ideals of scholarship,

and by its devotion to truth and untainted judgment has won

the confidence of those it teaches, there will be little danger that

Clancy — Floundering in Modernity,

55

student opinion will be swept off its feet by the facile and intoxi¬

cating appeal of any superficial speaker. Judgments will be held

in abeyance for leisurely review. But if there were to lurk in the

mind of the undergraduate a suspicion of the integrity of the

scholarship and free mental processes of his instructor, the student

will take supreme — and, indeed, well- justified^ — ^joy in exploding

bomb after bomb within the unprofaned precincts of his college.

The student of today is quick as a flash to catch the note of insin¬

cerity and weak compromise; though often he himself is given to

specious argument, he is scathing in his criticism when he flnds

such a defect in his instructor’s reasoning. Fundamentally, the

college student is the most stalwart of moralists, he is insistent on

fair play, and his chronic radicalism is his attempt to set the world

right. But unfortunately, he is also the most lax of thinking

beings, his contempt for things in high places is not a disciplined

contempt, he seldom seeks his data far, or subjects himself to the

rigors of logical and painstaking analysis. He has the failings

of the generation he represents, he is curious about the new, dis¬

cards the old without compunction or regret, is hurried in his

judgments and superficial in his culture; his ardor for things in¬

tellectual is unkindled. But his instincts are right and his capacity

for devotion to what is fine is unlimited.

In the midst of the college and of the broader world of which

the college is a part, the scholar may hold a position of eminence.

In his make-up there can be no suggestion of the mere prettiness of

culture; his mentality rather connotes wrought-iron ; he teaches

the Greek virtues of discipline, measure, and proportion; the very

essence of his nature is a protest against the facile thinking of

the man who would reform the world in thirty days. In the midst

of an acquisitive society curiously obsessed by material values,

he looks for things that are enduring, things tried by the fierce

fires of human experience and suffering. He knows that civilization

has been won by painful struggle, that its finest achievements are

the product of disinterested searchers for truth. The continuation

of the sublime traditions of his profession is not an easy task

in these days when a thousand discordant voices are crying in the

wilderness, ‘‘Prepare ye the way of the Lord.” Vague and windy

theorizing, glittering generalities, and sweeping denunciations must

all be referred to the discriminating and wise scholarship which

is the saving hope of our civilization.

BIBLIOGRAPHICAL EVIDENCE OF THE VOGUE OF

SHAFTESBURY IN THE EIGHTEENTH CENTURY

William E. Alderman

The importance of the “contributions of Anthony Ashley Cooper,

Third Earl of Shaftesbury, to the science of ethics has long been

recognized and frequently elaborated, both in English and in Ger¬

man. Strangely enough, however, he has been notoriously neg¬

lected as a writer whose teachings entered largely into the con¬

sciousness of his century, and who, consequently, was a potent

force in suggesting the content of a large body of English liter¬

ature. So definitive a work as the Cambridge History of English

Literature profeses to be allows slightly less than two pages in its

main entry ^ to an enumeration of his works and a partial cata¬

loguing of his ideas. The utter disregard of certain writers is

even more noticeable.^ Only a very few authors of histories and

.handbooks give him the place of eminence which is indubitably

his.®

It has long been the custom to trace a certain bent of the eight¬

eenth century temper either immediately and wholly, in its be¬

ginnings, to certain poems of the preceding period, or largely, in its

later developments, to certain foreign influences which, as time

went on, became operative. To deny the partial truth of either

of these points of view would be quite contrary to the spirit and

method of this present study; but to suggest that certain poems

of the seventeenth century have been credited with greater genera¬

tive power than they, of themselves, possessed, and that certain

foreign ideas found their counterparts already firmly intrenched in

English thought and literature, is quite a different matter.

The present study, however, does not aim to suggest an elaborate

philosophical background to the moods that mark the beginnings

of romanticism, or to establish definitively the thesis that ideas

identical with those to which Rousseau gave such brilliant state-

iVol. IX, p. 334 ff.

^ The index to English Literature in the Eighteenth Century, by T. S. Perry, contains

not so much as a single reference to Shaftesbury.

“ Gosse, Eighteenth Century Literature is perhaps the most complimentary.

58 Wisconsin Academy of Sciences, Arts, and Letters.

ment, and which are variously branded as romantic, ultra romantic,

sentimental, and naturalistic, were part and parcel of English spec¬

ulations before the intellectual invasion from abroad; for this the

reader must look elsewhere.^ Prom what follows it is evident,

however, that the ideas of Shaftesbury, whatever they were, were

both popular and potent in the century to which they were given.

I.

Forty years after the first collected publications of the essays

of Shaftesbury under the title of Characteristicks of Men, Manners,

Opinions, Times (1711), “Estimate” Brown, in reviewing their

reception and influence, remarked: “It has been the fate of

Shaftesbury’s Characteristics, beyond that of most books, to be

idolized by one party, and detested by another. While the first

regard it as a work of perfect excellence, as containing everything

that can render mankind wise and happy; the latter are disposed

to rank it amongst the most pernicious of writings, and brand

it as. one continued heap of fustian, scurrility, and falsehood. ’

So accurately does this represent the varying currents of ap¬

probation and disapprobation, that it deserves the place of promi¬

nence that it has been given in this study. For it must be kept

in mind continually that the intellectual initiator is certain to

have antagonists as well as vindicators, and that the volume of

discussion that he provokes is likely to be in direct proportion to

his significance. Mediocrity rarely begets either friends or enemies

in great numbers. To greatness is reserved the honor of being

both understood and misinterpreted.

With half a century more of retrospect than had Brown, Thomas

Park, continuing and enlarging the Royal and Noble Authors

of Walpole, gave attest to the same erratic popularity of Shaftes¬

bury. “Few writings have attracted more attention, or excited

more discussion, than the works of this noble author ; who has been

applauded and condemned with equal extravagance . . . .

For a considerable time he stood in high repute as a polite writer,

and was regarded by many as the standard of elegant composition ;

his imitators, as well as his admirers, were numerous, and he was

* See The Significance of Shaftesbury in English Speculation, by the present writer,

P. M. L. A., Vol. XXXVIII, 175-195; The Style of Shaftesbury, also by the present

writer, M. L. N., Vol. XXXVIII, 209-215; Shaftesbury and the Ethical Poets, by C. A.

Moore, P. M. L. A., Vol. XXXI, 264-325.

^Essays on the Characteristics, p. 1.

Alderman — Bibliographical Evidence of Shaftesbury. 59

esteemed the head of the school of sentimental philosophy. Of late

years he has been as much depreciated as he was before extolled. ’

Although it is not at present the primary purpose to examine

definitely that host of friendly and unfriendly essays of which

the Characteristics was provocative, it must be borne in mind

continually that they do attest eloquently to the fact that Shaftes¬

bury had entered into the philosophical and theological conscious¬

ness of his own century, and that he was generally regarded as

the most typical representative of a certain body of ideas, variously

thought of as being wholesome or pernicious. It matters not, there¬

fore, whether a writer champions, extenuates, or assails our author,

his manner, or his doctrines; all attitudes substantiate the fact

that Shaftesbury was to the people of his age a figure of no incon¬

siderable prominence.

Furthermore, it should be noted that, although many of the

tenets of Shaftesbury are actually traceable to former sources,

he is, for all practical purposes, generally regarded as their

originator. It was he who gave them their most brilliant state¬

ment, and it was he, therefore, more than any one else, who was

responsible for their vogue. It was with Shaftesbury himself,

and not with his intellectual progenitors, that such prominent lit¬

erary, philosophical, and theological writers as Mandeville, Butler,

Berkeley, and Hutcheson were concerned. The frequency of his

name in titles, and the direct and indirect references to him in

poems, essays, novels, and controversial literature show how inti¬

mately and inseparably he was connected in the general conscious¬

ness with the inception and propagation of the ideas that stand

to his credit or discredit.

This close association of the name of Shaftesbury with a large

body of sentimental and optimistic ideas, and his indubitable

prominence in several of the important intellectual wars of the

century, lead one inevitably to question the validity of that form

of criticism that has been all too religious in its attempt to trace

to foreign sources all that is bad and much that is good in English

thought and literature. Such a critic as Professor Babbitt sees

immediately that ^‘Considered purely as an initiator Shaftesbury

is probably more important than Rousseau.’’^ But even a super¬

ficial examination of the writings of the century enforces the fact

that he was more than initiator; he was a sincere and vigorous

® Walpole’s Royal and Noble Authors, 5 vols., London, 1806, IV, 55.

Rousseau and Romanticism, p, 44.

60 Wisconsin Academy of Sciences ^ Arts, and Letters,

propagandist, whose writings, by reason of their manner and

matter, became immediately popular and affected deeply the specu¬

lations of the time.

Elsewhere instances of a nature different from those cited below

have been multiplied to show how Shaftesbury insinuated himself

upon the thought of the age.® In many cases the borrowings have

been immediate; in others, they are apparently secondhand. But

this by no means lessens the importance of our figure. Whether

the idea of a subsequent writer came from Shaftesbury, or Pope,

or Akenside, or Hutcheson, or any one of a dozen others, matters

little. The fact remains that Shaftesbury was a brilliant initiator,

whose disciples helped to spread his doctrines broadcast. Socrates

taught Plato, Plato taught Aristotle, and Aristotle taught Alex¬

ander the Great ; but this does not lessen the greatness of the first

of this illustrious quartette, — quite the contrary. Shaftesbury

taught Hutcheson, Pope, and Thomson, and they, in turn, taught

an increasingly large number of writers; but Shaftesbury remains

the source whence the streams of optimism and sentimentalism of

a certain hue flow.

II.

Anthony Ashley Cooper, the son of Dryden’s ‘‘shapeless lump”

and the grandson of his “ Achitophel, ” had the good fortune to be

ushered into this world with Locke as the attending physician.

By the grandfather, to this already famous philosopher was en¬

trusted the health and mental development of the child. Applying

his own theories of education, Locke had young Ashley instructed

in Latin and Greek by the conversational method. The success of

the method and the precocity of the youngster both find support

in the fact that at the age of seven the coming Earl could read

both languages readily — a fact that needs to be kept in mind

when dealing with his indebtedness to the ancients. His travels

on the continent after leaving school in 1686, his year in Holland

after his enforced retirement from Parliament in 1698, his year

in Holland in 1703-1704 when his health was “mightily impaired

by fatigue in public affairs,” and his withdrawal to Italy in 1711,

all helped to make him the cosmopolitan that he was. His politi¬

cal positions gave him prestige and wide associations at home;

and his affability and mental vigor made him the acceptable com-

® See note 4 supra; The Influence of Shaftesbury on English Literature in the

Eighteenth Century, by the present writer, in MS in the library of the University of

Wisconsin.

Alderman — Bibliographical Evidence of Shaftesbury. 61

panion of such luminaries as Bayle and Le Clere abroad. Were

not the purpose of this study other than biographical, much of

interest could be said relative to his ardent Whiggism, his constant

devotion to principle, the simplicity and sincerity of his private

life, and his benevolence toward literary men, struggling students,

and the poor of his own neighborhood. For these the reader must

look elsewhere.® Suffice it to say here that such was his general

culture that he has without exaggeration been called ‘ ‘ the Matthew

Arnold of Queen Anne’s reign. Gosse has characterized him as

“the most accomplished Englishman of his day, the man with the

widest taste and the most complete culture, while the purity of his

personal character matched well with the charm of his intellect.

Generous both in theory and in practice; prodigal with advice,

yet consistent and temperate in habits; devoted to a certain set

of principles, but catholic in his point of view; this statesman,

philosopher, and litterateur died before he had yet completed his

forty-second year.^^ He was never vigorous physically, but lived

actively and well ; and his works stand as his best memorial.

Shaftesbury’s first venture into print was in the year 1698 when

a preface by him was prefixed to a volume of Selected Sermons by

Dr. Benjamin Whichcote, the distinguished Cambridge Platonist or

Latitudinarian.^® A glance at their contents reveals the cause

of Shaftesbury’s warm feeling for them, for already at this early

age he was committed to the ‘ ‘ Benevolent Theory of Morals. ’ ’ The

sermons became very rare, but were reprinted in Edinburgh in

1742, this time with a preface by Dr. Wishart.^^ Shaftesbury’s

Preface was reprinted in connection with a volume of his letters,

which counted as the fourth volume of his Characteristics, in 1758.^®

In 1698 Shaftesbury was compelled by poor health to leave

Parliament and to seek strength in Holland. It was during this

sojourn abroad that Toland surreptitiously published the Inquiry

® See Stephen’s article in D. N. B.; Rand, Life, Letters and Philosophical Regimen;

Fourth Earl of Shaftesbury, Life of the Third Earl; Bayle, Life of Shaftesbury, in

General Dictionary.

Leslie Stephen, FreethinMng and PlainspeaTcing, p. 242.

'^Eighteenth Century Literature, p. 171.

Stephen, English Thought in the Eighteenth Century, II, 20, note, says that the

Characteristics first appeared in 1711, the year of his death. This is an error. The

Characteristics did first appear in 1711, hut Shaftesbury did not die until 1713.

For a full statement of the relationship that existed between Shaftesbury and

Whichcote see P. M. L. A., XXXVIII, 183-189.

See also The 'Wor'ks of Benjamin Whichcote, Aberdeen, 1751; Ernest P. Cam-

pagnac. The Cambridge Platonists, 1901.

15 Fowler, Shaftesbury and Hutcheson, New York, 1883, p. 48.

62 Wisconsin Academy of Sciences, Arts, and Letters.

concerning Virtue (1699)/® Robertson’s remark ‘‘that Shaftes¬

bury should at the age of eighteen have produced from his own

meditations a finished and formal treatise, of which the theses were

capable of influencing European thought for a century, would be

an extravagant assumption, is misleading, if not absolutely

wrong as to date. Evidently he is thinking of the years 1698 and

1699, for he goes on to say that Shaftesbury could have learned

his Spinoza in Holland at this time. This would make Shaftesbury

twenty-eight rather than eighteen — a considerable difference, in¬

deed! He can hardly be thinking of what Fowler^® styles a

‘‘rough draft” sketched when the Third Earl was twenty. Nor

did Shaftesbury regard this premature publication as “finished

and formal,” for in the First Edition of the Char act eristicks (1711)

the statement is made that the essay was “formerly printed from

an imperfect copy ; now corrected and publish ’d intire. ’ ’ However

complete the revision may have been,^® it is certain that long before

the publication of the Preface to Whichcote’s Sermons, Shaftes¬

bury was thinking deeply on questions that were to lead him to

his system of ethics. The contradiction between zeal and lack of

humanity in religion gave him occasion to inquire “what honesty

or virtue is, considered by itself, and in what manner it is in¬

fluenced by religion; how far religion necessarily implies virtue;

and whether it be a true saying that it is impossible for an atheist

to be virtuous, or share any real degree of honesty or merit. ’

The so-called French Prophets, or poor Cevenol Protestants,

by their extravagant enthusiasm and insane practices drew forth

the opinion of Shaftesbury in A Letter Concerning Enthusiasm

(1708) addressed to Lord Somers. Various methods of dealing

with them had been proposed, but Shaftesbury contended that

“raillery” and “good humor” were most potent against false

enthusiasm, that “Ridicule and not Punishment, is the most

effective weapon of Fanaticism.” The Letter stirred up three re¬

plies in 1708 and 1709, and Shaftesbury returned to his general

Stephen, English Thought in the Eighteenth Century, II, 20, note, remarks that the

“Essay on ‘Virtue’ had been published in an imperfect state by Toland in 1698.’’

Shaftesbury did go to Holland in 1698, hut did not return to England until November

10, 1699. Evidently the publication took place in this latter year, for the essay is

described in the first edition of the Characteristics, 1711, as “first published in 1699.’’

Characteristics, Vol. I, p. XXXII.

«P. 15.

Shaftesbury bought up the whole impression of Toland when but a few copies had

been sold; consequently it is almost impossible to get hold of the earlier version for

purposes of comparison.

^ Inquiry concerning Virtue, Bk. I, Pt. I, section 1.

Alderman — Bibliographical Evidence of Shaftesbury. 63

thesis in Sensus Communis: An Essay on the Freedom of Wit

and Humor, (May, 1709).

The Moralists, a Philosophical Rhapsody (1709) is in imitation

of the Dialogues of Plato. Bishop Hiird speaks of it as one of the

three dialogues in the English language fit to be mentioned and

this is no small praise, for some had preceded it and many others

were to follow before Hurd passed his judgment. As Theocles,

Shaftesbury unfolds his optimistic theories and theology, and sings

his Nature hymns.

His next treatise, Soliloquy: or Advice to an Author (1710), is

a mine of comment and reflection on various topics.^- Of it he

says in his Miscellaneous Reflections that ^^His pretense has been

to advise Authors and Polish Styles ; but his aim has been to cor¬

rect Manners, and regulate Lives/

Miscellaneous Reflections (1711) was first published in the. First

Edition of the Characteristics. Later it was described as first

printed in 1714^’, an error due, perhaps, to Printed in the year

M.DCC.XIV” which appears on the title page to the treatise in

the collected works of that date. These Reflections, varied and

illuminating, were intended to bolster and supplement the other

treatises. ^

In Naples, whither he had gone for his health in 1711, he wrote

A Notion of the Historical Draught or Tablature of the Judgment

of Hercules and the Letter Concerning Design. The former was

first printed in Prance, in the Journal des Sgavans for November,

1712, and appeared in English, separately, in 1713, and in the

Characteristics of 1714; the latter, although it occurs in a large

paper copy of the Second Edition in the British Museum, seems

not to have been included generally until 1732.

That Shaftesbury was an assiduous writer of letters can be seen

both from the total number that he wrote and the painstaking with

which each was written. Several Letters written by a Noble Lord

to a Young Man at the University, addressed to Michael Ainsworth,

appeared first in 1716 and again in 1732. In 1721, Toland, for¬

getful of his first surreptitious venture of 1699 that had been so

generously followed by an annual stipend from Lord Ashley, again

Worlss, London, 1811, VoL III, pp. 24—25. He says, “The Dialogues I mean are.

The Moralists by Lord Shaftesbury; Mr. Addison’s Treatise on Medals; and the Minute

Philosopher of Bishop Berkeley.” Also quoted by Warton, Essay on Pope, London, 1806,

Vol. II, p. 198.

^ For an excellent summary of it see Fowler, p, 54.

^ Cited by Fowler, p, 54.

64

Wisconsin Academy of Sciences, Arts, and Letters.

broke faith, this time with Shaftesbury’s relatives and friends, by

bringing out an unauthorized edition of certain private letters to

Robert Molesworth. The epistles, according to Toland’s ‘‘large

introduction” are “upon two of the nicest Subjects possible, and

the most important to Mankind ; the one private, the other publick ;

the first being the Choice of a Wife, and the second the Service

to one^s Country.’’-^ Subsequent volumes of letters appeared in

1746, 1750 and 1758.

All of the works intended by Shaftesbury for publication, with

the exception of the Preface to Dr. Whichcote’s Sermons, are to

be found in editions of the Characteristics beginning with 1732.

His letters, the so-called Philosophical Begimen,^^ and treatises III

and IV of Second Characters'^^ did not appear until after his death.

Little more than a mere catalogue of his works has been at¬

tempted. The chronological tabulation of them given below^^ is

illuminating, in that it shows that they were in great demand, and

that his popularity did not die out at any point in the century.

Letters to Molesworth, p. 1.

2® Published 1900, Benjamin Rand, editor, Macmillan Co., N. Y.

Rand, Cambridge University Press, 1914.

1698 Preface by Shaftesbury to Selected Sermons of Benjamin Whichcote.

1699 Inquiry concerning Virtue, surreptitiously published by Toland.

1708 Letter concerning Enthusiasm.

1709 The Moralists, a Philosophical Rhapsody.

1709 Sensus Communis : An Essay on the Freedom of Wit and Humor.

1710 Soliloquy: or Advice to an Author.

1711 Miscellaneous Reflections (published in the Characteristics).

1711 Char act eristicks of Men, Manners, Opinions, Times, First Edition, 3 vols.

1712 The Judgment of Hercules (in French).

1713 The Judgment of Hercules (in English).

1714 Letter concerning Design (appears for the first time in a large paper copy of

the Second Edition of the Characteristics in the British Museum.)

1714 Characteristics, Second Edition, 3 vols.

1716 Several Letters written by a Noble Lord to a Young Man at the University,

addressed to Michael Ainsworth.

1721 Letters from the Late Earl of Shaftesbury to Robert Molesworth, Esq. Intro¬

duction by Toland.

1723 Characteristics, Third Edition, 3 vols., 8vo., London.

1727 Characteristics, Fourth Edition, 3 vols., 8vo., London.

1732 Characteristics, Fifth Edition, 3 vols., 8vo., London

1732 Several Letters written by a Noble Lord to a Young Man at the University.

1733 Characteristics, Another Edition, 3 vols., 12mo., London.

1737 Characteristics, Sixth Edition, 3 vols., 8vo., London.

1744 Characteristics, Another Edition, 3 vols., 12 mo.

1746 Letters of the Earl of Shaftesbury, collected in one volume.

1749 Characteristics, Another Edition, 3 vols., 12mo., London.

1750 Letters of the Earl of Shaftesbury, collected in one volume.

1758 Letters of the Earl of Shaftesbury, including the Preface to Dr. Whichcote’s

Sermons.

1773 Characteristics, Fifth Edition, printed by Baskerville, Birmingham.

1790 Characteristics, with a collection of letters, 8vo., Basil.

Alderman — Bibliographical Evidence of Shaftesbury. 65

Long before Shaftesbury had established himself as a writer,

the appearances of his unsigned treatises excited wide comment

and speculation. Swift, writing from London to Robert Hunter

in Paris, January 12th, 1708-09, remarks, ‘‘I cannot forbare telling

you of your mechancete to impute the Letter on Enthusiasm to me ;

when I have some good reasons to think the author is now in

Paris. And again in writing to Ambrose Philips, September

14, 1708, he says, ‘‘There has been an essay on Enthusiasm lately

published, that has run mightily, and is very well writ. All my

friends will have me to be the author, sed ego non credulus illis.

By the free Whiggish thinking I should rather take it to be yours,

but mine it is not. . .

The appearance, during the period 1698-1790, of twenty-five

publications and republications of the works of this “noble au¬

thor” was not without its cause. The books were popular and

sold rapidly. The editor of the 1733 pocket edition triumphantly

asserts that “all the best Judges are agreed that we have never

had any work in the English language, so beautiful, so delightful,

and so instructive as these Char act eristichs.^^^^ Nor can this be

considered a commercial exaggeration, for he goes on to state what

obviously was a fact : “And five large Editions being sold off, give

a very sensible proof of their being generally liked.” Eleven edi¬

tions of the Characteristics, each of three volumes, appeared be¬

tween 1711 and 1790. “The reception these writings have met

with from all persons of good taste and judgment, has been such

as might have given great satisfaction to that truly noble and in¬

genious Author, if he had lived longer to enjoy it.”^^

III.

Nor are these numerous appearances of his works the only proofs

of his vitality. Scarcely had his unsigned Letter concerning En¬

thusiasm (1708) appeared, when speculation began as to its author¬

ship. Swift generally was thought to be its originator, but he in

turn passed the implication on to Philips and Hunter. There

were forthcoming almost immediately three replies — Remarks on

a Letter by a Lord concerning Enthusiasm, not written in raillery

but in good humor, published anonymously; Bart’lemy Fair, or

^^Correspondence of Jonathan Swift, edited by F. E. Ball, London, 1910, p. 136.

^Correspondence, London, 1910, p. 111.

Preface to 1733 edition.

Gosse, Eighteenth Century Literature, p. 387.

66 Wisconsin Academy of Sciences, Arts, and Letters.

an Enquiry after Wit, by Dr. Wotton; and Reflections upon a

Letter concerning Enthusiasm, now generally credited to Dr. Ed¬

ward Fowler, Bishop of Gloucester. All were alert to the possi¬

bility of the author ^s implied reflection upon the English clergy

as well as his open attack on the Cevenol peasants. Shaftesbury

was swift to come to the defense of his cherished doctrine with

Sensus Communis: An Essay on the Freedom of Wit and Rumor

(1709). What, apparently, is another attack is that attributed to

M. Astell entitled An Enquiry after Wit: wherein the trifling

argument and impious raillery of the late Earl of Shaftesbury in

his Letter concerning Enthusiasm . . . are fully answered.^^

Doubtless the most formidable opponent of Shaftesbury was the

arch cynic Bernard de Mandeville (1670-1733) whose A Search

into the Nature of Society (1723), added to the second edition of

the Fable of the Bees, launched his attack. As the eighteenth cen¬

tury representative of Hobbes he sees in man a ‘^Compound of

Evil Passions’’, makes a benefit of luxury, a vice of natural pas¬

sions, and a necessity of crime. “Two systems”, he says, “cannot

be more opposite than his Lordship’s and mine.” By continuing

his attacks in the Dialogues, published as a second part of the

Fable of the Bees in 1728, Mandeville, as Cleomenes, represents

Shaftesbury as an enemy of revealed religion.

But Mandeville ’s attack was too coarse to be tolerated by self-

respecting and nation-loving subjects, and again Shaftesbury had

his defenders in rapid succession. John Dennis’s Vice and Luxury

Public Mischiefs (1724), William Law’s Remarks on the Fable of

the Bees (1724), Richard Piddes’s A General Treatise on Morality

(1724), Francis Hutcheson’s An Inquiry into the Original of our

Ideas of Beauty and Virtue (1725) and his Observations upon the

Fable of the Bees (1725-27), Archibald Campbell’s Aretelogia

(1728), and parts of Berkeley’s Alciphron (1732), and John

Brown’s Essays on the Characteristics (1751) and An Estimate

(1757) are at times as much pro Shaftesbury as they are anti

Mandeville.

One of the most effective replies of the whole controversy was

that of John Bulgay, a follower of Clarke, in A Letter to a Deist

concerning the Beauty and Excellency of Moral Virtue, and the

support and improvement which it receives from the Christian

Revelation (1726). The burden of this rational work is well sum¬

marized in its own words: “In short, the question is not, which

See British Museum Catalogue under Shaftesbury.

Alderman — Bibliographical Evidence of Shaftesbury, 67

motives are the purest and most sublime ; but which are most use¬

ful, and most effectual, to prevail with degenerate man and accom¬

plish his regeneration. ^ ’ The answer is clearly implied in the title.

Joseph Butler (1692-1762) was both a friend and foe of Shaftes¬

bury. In his Preface to the 1729 edition of .the Fifteen Sermons

he gives not a little space to the discussion of Shaftesbury’s teach¬

ings. In fact the Third Earl is the only one to whom he does give

elaborate or explicit attention. Although he readily admits him

to be authoritative in the main, and says that ^^he has shown be¬

yond all contradiction that virtue is naturally the interest or hap¬

piness, and vice the misery, of such a creature as man, placed in

the circumstances which we are in this world,” he cannot accept

as adequate his moral sense” as a guide to correct action.

Bishop Berkeley (1685-1753) in the third Dialogue of Alciphron,

or the Minute Philosopher (1732), adopts a kind of Theological

Utilitarianism not unlike that of Locke. ^^Alciphron, adapting

Shaftesbury, reduces conscience to a taste, enlarges upon the

beauty of virtue, and disparages faith in a future life as a selfish

and cowardly appeal to hope and fear. Against this Euphranor

maintains that a sense of the beauty of goodness is inadequate for

making us good, as man needs for this a stronger and more awe¬

inspiring motive than taste: the springs of action must be sus¬

tained by faith in the destiny of man under God. ’ A strange

answer to Berkeley appeared two years after the Minute Philos¬

opher under the title A Vindication of the Beverend D -B— — Y

from the scandalous imputation of being the author of the late

book entitled Alciphron, or the Minute Philosopher. Affecting

to believe Berkeley’s attack a forgery, the author refutes it with

passages from Shaftesbury, and with statements none too friendly

toward orthodoxy.

A country clergyman by the name of Elisha Smith came forward

in 1736 with The Cure of Deism. In the title he charges Tindal

and Shaftesbury with having given a ‘Wery imperfect account of

the religion of Nature, and of Christianity,” and proposes ^Uhe

Mediatorial scheme of Jesus Christ” as *Ghe only true Eeligion.”

The work proved to be of great interest and was reprinted in octavo

form in 1737, 1739, and 1740. This is especially interesting in

that it shows the constant tendency of the time to regard Shaftes¬

bury as an open enemy to all those things sacred to orthodoxy.

Dr. War burton, who in his Dedication of the Divine Legation to

Fraser in Preface to Aleiphron in Worlcs, Oxford, 1901. Vol. II, p. 10.

68 Wisconsin Academy of Sciences, Arts, and Letters.

the Free-Thinkers (1738) has a scattered attack on Shaftesbury,

wrote to Dr. Hurd, January 30, 1749-50, '‘Mr. Pope told me, that,

to his knowledge, the Characteristics had done more harm to Re¬

vealed Religion in England than all the works of Infidelity put to¬

gether. Skelton,. in Deism Bevealed^^ asserts that Shaftesbury

“labours to strike out and establish a new system of morality, un¬

happily founded on a notorious falsehood. ’ His teachings as to

rewards and punishments he brands as “nothing else in effect but

downright practical Atheism.

But the attacks and defenses were not yet at an end. John

Brown, realizing the seriousness of Shaftesbury ^s vogue, brought

out his popular Essays on the Characteristics in 1751.^^ Taste is

again thought to be too vague a criterion for the morality of the

generality of mankind; only the orthodox belief in rewards and

punishments can deter man from vice. Despite the general fair¬

ness of Brown, his book only served to fan the fire of admiration

that had never died in the hearts of the friends of Shaftesbury.

Almost immediately Charles Bulkley, a dissenting minister, came

to the fore with A Vindication of Lord Shaftesbury on the Sub¬

jects of Morality and Beligion; and an anonymous writer pro¬

duced the well-written Animadversions on Mr. Brownes Three

Essays on the Characteristics.

That curious respect that even the enemies of Shaftesbury had

for his person and his works is typically stated by Leland in View

of the Principal Deistical Writers (1754). “It gives me real con¬

cern, that among the writers who have appeared against revealed

religion, I am obliged to take notice of the noble author of the

Characteristics.^ And yet these books “are so generally read,

and by many so much admired, that it is necessary to take notice

of those things in them which seem to have a bad aspect on religion,

and to be of dangerous influence and tendency. ’ Having spoken

of his style, his opposition to Hobbes, and his refined sentiments

on virtue, Leland admits with regret that these “have very much

prejudiced many persons in his favor, and prepared them for re-

See Nichols' Literary Anecdotes, II, 212; or Chalmers’ Biographical Dictionary, arti¬

cle on John Brown.

Published in 1749 and 1751.

3«Vol. I, p. 134,

IMd.

»8 Republished in 1752, 1764.

88 1757 edition, I, p. 48,

*»I, p. 49.

Alderman — Bibliographical Evidence of Shaftesbury, 69

ceiving, almost implicitly, whatever he hath advanced. De¬

spite the dangers that Leland thought himself called upon to battle

against, Shaftesbury remained popular, and the most sumptuous

edition of his works was soon to appear — the famous Baskerville

edition of 1773. Again the complaints and warnings of the ortho¬

dox were voiced, this time by John Ogilvie in An Inqury into the

Causes of the Infidelity and Skepticism of the Times (1783), in

which Shaftesbury came in for his share of the blame. But the

most significant reply was another edition of his treatises and let¬

ters seven years later.

Any writer, philosophical or other, who has it within him to

create such a controversial furor as Shaftesbury precipitated, and

who can, despite opposition, maintain his popularity and retain

his defenders, is a man of no mean vigor. Thomas Gray in a letter

to Richard Stonehewer, (August 18, 1758), accounting for his

vogue as a philosopher, said : ‘ ‘ First, he was a Lord ; 2dly, he was

as vain as any of his readers ; 3dly, men are very prone to believe

what they do not understand; 4thly, they will believe anything at

all, provided they are under no obligation to believe it ; 5thly, they

love to take a new road, even when that road leads nowhere ; 6thly,

he was a fine writer, and seemed always to mean more than he

said. ’ But these reasons are puerile when we trace the constant

republication of his works, and recall the challenge that he gave to

the best intellects of the century. Men are not likely to continue

to read the subtle speculations of a dead Earl simply because they

cannot understand them, or for the sheer pleasure of being led

they know not whither. A leader, no matter how suave or vain,

will hardly make intellectual friends or enemies if there is no

“obligation to believe’^ that which he says in all seriousness.

But Shaftesbury weighed heavily upon the conscience of the age.

As John Armstrong remarked,^®

“Ashley has turn’d more solid heads than one.”

No, Gray was all too nonchalant and facetious in accounting for

the vogue of Shaftesbury.

Montesquieu in his Pensees Diverses ranks him with Plato, Male-

branche, and Montaigne as one of the four great poets.^^ Voltaire

« I, p. 49.

Gray, Gosse edition, p. 375.

Taste, 1755, found in Chalmers XVI, p. 538.

** (Euv. Comp., Paris, 1838, p. 626.

70 Wisconsin Academy of Sciences^ Arts^ and Letters.

mentions him repeatedly in his Lettres sur les Anglais or Lettres

Philosophiques. Diderot reproduced his Inquiry concerning Vir¬

tue in Essai sur le Merite et la Virtue (1745 and 1751) ; and in

1769 a French translation of the whole of Shaftesbury’s works, the

letters included, together with a French introduction of twenty-six

pages, appeared in Geneva. Herder in 1794 said that this “vir¬

tuoso of humanity” had signally influenced the best heads of the

eighteenth century.^® Leibnitz, Mendelssohn, and Wieland drank

of his waters. He began to be turned into German in 1738, and in

1768 and 1776-1779 translations of his Characteristics appeared.

Here then we have an international as well as a national char¬

acter. In order to determine wherein lay his peculiar power and

originality, it would be necessary first to look into the ethical and

theological dogmas of his predecessors. When their thoughts had

been compared with his thoughts, and their ways with his ways, we

could move with some certainty into the literature of the century

in an attempt to see to what extent it took color from his style and

his ideas. But this, which has been done elsewhere,^® gains force

in the light of the bibliographical facts just rehearsed. Shaftes¬

bury becomes at once an originator and a propagandist, whose

ideas, regarded both as toxic and as therapeutic, were widely dis¬

seminated and increasingly accepted in the century of “prose and

reason”.

Beloit College.

See Brief e zur Beforderung der Eumdnitat, Briefe 32, 33.

See notes 4 and 8 supra.

NOTES ON NEW NAMES IN TABLE OF FORMATIONS AND

ON PHYSICAL EVIDENCE OF BREAKS BETWEEN

PALEOZOIC SYSTEMS IN WISCONSIN

E. 0. Ulrich

Introductory note hy TV. 0. Hotchkiss. The Geological^ and

Natural History Survey of Wisconsin is deeply indebted to the

writer of this paper, and to the United States Geological Survey

for the cooperation which has made possible the important results

set forth. The field work on which Dr. Ulrich’s work is based be¬

gan in 1913 and has been carried on during short periods of a few

days or weeks each summer since, as the pressure of his other duties

permitted. The work has added greatly to our knowledge of the

Paleozoic formations, and -while this is more than ample justifi¬

cation, it is satisfying to record that the results have been imme¬

diately applicable to economic uses as w^ell. The successful search

for local supplies of shale for road surfacing in western Wiscon¬

sin would have been impossible without Dr. Ulrich’s work as a

basis. The guidance of drillers of deep wells and the close identifi¬

cation of the strata in these wells which has made possible the find¬

ing of desirable water supplies also rests largely on the results of

his work.

Dr. Ulrich’s work has resulted in very much more detailed

knowledge of the strata and has shown the need for giving names

to the new units recognized. It has also been deemed advisable

in his paper to discuss somewhat fully the physical evidence on

which the limits of the various formations are determined.

The Burroughs Dolomite

Near the top of Bur rough’s Bluff, at the northern end of Savan¬

nah, Ill., and also in and above Charles Miles’ quarry near the

southeastern edge of the same city the easily recognized Brainard

shale at the top of the Maquoketa facies of the Richmond group is

succeeded unconformably by a variable succession of bluish to

*Published by permission of the Director of the U. S. Geol. Survey.

72 Wisconsin Academy of Sciences, Arts, and Letters.

grayish yellow irregularly bedded magnesian mudstones and brown¬

ish dolomite aggregating some 50 to 60 feet in thickness. On weath¬

ering the upper half or more of these beds shows more or less

of earthy chert in nodules and uneven plates. The bluish gray

lower third contains few fossils, those found being of a diplograptid

(? Mesograptus sp.) Dictyonema sp. and broadly branching car¬

bonaceous films supposed to belong to some marine plant. The lat¬

ter two seem indistinguishable from specimens found in the lower

part of the Cataract formation in Ontario. The cherty upper beds

contain a more varied though not abundant fauna. Among them

I have provisionally identified the following: Streptelasma aft.

divaricans and radicans, Streptelasma aft. rnstica, Lindstromia sp.,

Halysites sp., Favosites sp., Anaphragma aff. mirahile, Phaenopora

ensiformis, Phaenopora cf. fimhriata, N ematopora aff. delicatula,

Helopora cf. fragilis, Bhinopora sp.. Lingula sp., Pholidops sp.,

Leptaena cf. rugosa, Plectorthis cf. whitfieldi, Dalmanella cf. edge-

woodensis, Strophonella cf. patent a, Atrypa praemargmalis, Zy-

gospira putilla, Tentaculites cf. oswegoensis, Orthoceras cf. sociale,

Proetus sp., Calymene sp. Obviously this fauna is decidedly post-

Richmond ; and it is as clearly older than Clinton. Evidently then

it falls into some part of the intermediate Upper Medina stage in

which the Edgewood formation of Missouri is probably a nearer

contemporary than the Cataract of Ontario.

Lithologically the formation is so different from the Edgewood

that it is thought unwise to employ that term for the beds under

consideration, especially as there is some doubt as to their strict

equivalence. According I propose the name Burroughs dolomite.

The same beds are indicated, though mainly by debris, in the

mounds in southwestern Wisconsin. Hitherto they have been

classed with the Niagaran dolomite that caps these and similar

mounds in northwestern Illinois and northeastern Iowa.

The Names Trempealeau Formation, St. Lawrence Limestone

OR Formation, and Jordan Sandstone

Three facts have been mainly responsible for the proposal to

substitute the new name Trempealeau formation for the series of

beds to which I have previously applied the term St. Lawrence.

First, the outcrops in St. Lawrence township, Minnesota, for which

N. H. Winchell originally proposed the name St. Lawrence lime¬

stone is so limited and its relations to overlying and underlying

Ulrich — Paleozoic Systems in Wisconsin.

73

Cambrian beds so imperfectly indicated that the locality consti¬

tutes a most unsatisfactory type for the composite formation I had

in mind when I adopted Winchell’s term for it in 1914. Second,

the name was originally applied only to the magnesian limestone

that lies in the lower part of the formation. The subsequent ex¬

pansion of the application of the name to everything between the

top of the Dresbach sandstone and the base of the Jordan sand¬

stone was unwise and unwarranted by rules governing in such

cases. Third, the name St. Lawrence, as applied to a stratigraphic

unit, ever since its proposal in 1874 has had an uncertain status

and meaning. At times it was referred to the Lower Magnesian

series and then to the St. Croixan ; and only very recently Keyes^

committed the now almost unpardonable error of correlating it

with the Oneota dolomite of Iowa. Besides, prior to 1914, all who

had occasion to refer to the St. Lawrence limestone made it the

same as the really much younger Mendota limestone of Wisconsin ;

and even today a few of the more conservative geologists adhere

to this old opinion. As an additional and final reason I may say

that employed strictly in its original sense there is still a desirable

and valid use for the term St. Lawrence limestone or dolomite. It

is almost needless to say that the St. Lawrence in the type locality

is not, as recently thought by Keyes, the Oneota dolomite but a

much older and thinner bed that is widely distributed, especially

in southern Wisconsin, where it lies either at or a few feet above

the base of the Trempealeau formation. At St. Lawrence, as very

generally too in Wisconsin, the bed is marked by a definitely Cam¬

brian fauna that is entirely different from the Upper Ozarkian

fauna which is found in the Oneota dolomite.

Judging from the varied and usually critical comments made

by geologists who have kindly read the original draft of this

paper it seems desirable and perhaps necessary to give a fairly full

account of the nomenclatural history of the term St. Lawrence

limestone and of the very different meanings in which the name has

been used by authors since it was first proposed by Winchell. This

account will at the same time serve in determining the relations

of the real Jordan sandstone to other sandstones with which it has

been confused and of the Franconia formation to the St. Lawrence,

the “Sparta Shale,’' and other formations that have been named

in the past ten years.

^ Keyes, Charles, Terranal differentiation of Iowa Cambric succession, The Pan-

American Geologist, vol. 38, No. 4, p. 323, 1922.

74

Wisconsin Academy of Sciences, Arts, and Letters,

The term St. Lawrence limestone was first proposed by N. H.

Winchell in his 2nd Annual Report, Minnesota Geological Sur¬

vey, page 152, 1874, the name being derived from exposures at St.

Lawrence, Scott County, Minnesota. Here the bed so named con¬

sists of hard magnesian limestone layers speckled with green (glau¬

conite) with a total exposed thickness of 14.5 feet. According to

well records given by Upham in his report on Scott County (Minn.

Geol. Sur. Final Rept. 2, p. 120-121, 1888) this bed may reach 25

or possibly 30 feet. In 1874 Winchell regarded this outcrop as

representing the lower division of the Lower Magnesian limestone

of Owen (subsequently named Oneota, limestone by McGee) and

as underlying the Jordan sandstone which he viewed as separating

the two divisions of the ‘‘Lower Magnesian.’’ The name Shakopee

limestone was given at the same time to the dolomitic formation

which overlies the J ordan in the valley of Minnesota River. So far

as the sequence of these three formations is concerned Winchell’s

original view has been proved correct. But he was in error in

correlating the Jordan and St. Lawrence with respectively middle

and lower divisions of the Lower Magnesian. Both of these forma¬

tions are pre-Oneota deposits. The Oneota belongs between the

Jordan and the Shakopee, but in this part of the Minnesota Val¬

ley the lower and more important part of the “Lower Magnesian”

limestone is thin and in places seems to be absent entirely.

In the 2nd, 4th, and 5th Annual Reports of the Minnesota Sur¬

vey the Shakopee is confused at times with the Oneota, and in

other places it is the St. Lawrence that is regarded as the equiva¬

lent of what we now distinguish as the Oneota. In the 2nd Annual

Report, as said, the Jordan is regarded as a middle member of the

Lower Magnesian series, the Shakopee as the upper, and the St.

Lawrence as the lower division. In the 3rd Annual Report the

name St. Lawrence is in places applied to the whole of the Lower

Magnesian, the locally developed sandstone (that was later named

New Richmond sandstone and which Winchell first suggested and

later claimed to be the same as his Jordan sandstone) being absent

in such places. “It (the St. Lawrence limestone) constitutes the

principal portion of the Lower Magnesian” (4th Annual Rept., p.

33, 1876). That the “St. Lawrence” as used in the 4th Annual

Report includes Shakopee is evident not only from the thicknesses

given (“not far from 200 feet”) but is indicated also by the men¬

tioned presence in the upper part of the massive algal remains

Ulrich — Paleozoic Systems in Wisconsin,

75

wMch lie later described as Cryptozoon minnesotense and which

occur only in the Shakopee.

On page 34 of the 4th Annual Report the railroad cut section at

Clear G-rit is described as showing 16 feet of sandstone at the top

which is called Jordan^ and beneath this 30 feet of dolomitic lime¬

stone called St. Lawrence. In the first place the sandstone is not

the Jordan but may be the New Richmond. It is a red sandstone

which^ as proved by experience, is of itself sufficient evidence to

establish its age as post- Jordan, the latter being everywhere a

grayish white though often iron-stained, case-hardened, interiorly

friable sandstone, the weathered surface of which commonly is

studded with highly characteristic rounded, often botryoidal con¬

cretions. Otherwise the true Jordan is closely similar to the St.

Peter sandstone. The dolomite which underlies the red sandstone

at Clear Grit belongs to the Lower Magnesian and most probably

is the upper part of the Oneota dolomite.

At other places noted in the 3rd Annual Report the Jordan is

properly recognized, but the overlying rock is not the Shakopee

as then supposed by Winchell but the Oneota.

Winchell’s discussion of the St. Lawrence, Jordan and Shakopee

formations in the 5th Annual Report, 1877, is essentially as in his

4th Annual Report. However, in this year he introduces his sub¬

sequently more definitely stated belief that Irving ^s Mendota dolo¬

mite and Madison sandstone of south central Wisconsin corre¬

spond to the St. Lawrence and Jordan formations of Minnesota.

That Winchell still correlated the St. Lawrence with what we now

know to be the whole of the Lower Magnesian,’’ that is in places

where the locally developed reddish weathering sandstone (the

New Richmond) is absent, is clearly indicated by his statement on

page 29, regarding the thickness (250 feet) of the formation at

La Crosse.

In 1888 (Final Report, Vol. 2, pp. XXI and XXII) Winchell

had changed his mind very greatly regarding the relations of the

early Paleozoic beds in the Minnesota Valley to those exposed to

the southeast in the bluffs along Root River and to the east along

the Mississippi. He now recognized that both the Jordan sand¬

stone and the St. Lawrence limestone are older than the lower

main mass of the “Lower Magnesian limestone,” that is than the

Oneota dolomite as now known, and that the sandstone for which

76 Wisconsin Academy of Sciences, Arts, and Letters.

he now adopts Woosters^ designation New Kichmond beds or sand¬

stone and which occurs locally at the contact of the Shakopee and

Oneota dolomites is a higher bed than the Jordan sandstone with

which he had previously confused it. Moreover, he now placed the

Jordan at the top of the St. Croix and referred the underlying St.

Lawrence to a correspondingly lower position in the same forma¬

tion or series. In this work also Winchell definitely correlated the

Jordan and the St. Lawrence respectively with the Madison sand¬

stone and the Mendota dolomite of the section at Madison, Wis.

In reading Winchell ’s comments on the several parts of the

section one notes the suggested desire to expand the limits of the

St. Lawrence so as to include “shaly beds with which it is asso¬

ciated and into which it seems to graduate.” These shaly beds

lie both above and beneath the typical St. Lawrence limestone

but mainly beneath, and if these were included the expanded forma¬

tion “will include beds to the amount of nearly 200 feet.” How¬

ever, in his designation and description of the cut and table show¬

ing the sequence of the lower Paleozoic formations in Minnesota

this suggestion is not carried out, the St. Lawrence being described

as 0-30 feet in thickness and the beds between the St. Lawrence

limestone and the underlying Dresbach sandstone are separately

given as “ 7. Sands and sandy shales ... at least 200 feet ’ ’ in the

first table and as “Shales” in the second.

From 1888 on to 1895 the term St. Lawrence was used in the

restricted original sense by all who had any occasion to refer to

the formation. On two occasions during this time HalP, and Hall

and Sardeson^ so used it. Also Keyes® and Calvin.® At the close

of this period Hall and Sardeson^ published an excellent paper in

which they discuss the “St. Lawrence dolomites and shales” at

considerable length and give the thickness of these beds as 213

feet. With such a thickness it would be impossible to exclude the

Franconia part of the section. That the “shales” must refer

mainly to the Franconia is clearly indicated by the fact that in the

table of formations the item “St. Lawrence dolomite and sandy

shale, 213 feet” immediately follows the “Faunal break” at the

2 Geology of Wisconsin, vol. 4, pp. 106 and 127, 1882.

® Minn. Acad. Nat. Sciences Bull., vol. 3, No. 1, p. 134, 1889.

* Geol. Soc. America, vol. 3, p. 341, 1892.

® Iowa Geol. Survey, vol. 1, p. 23, 1893.

® Ibidem, vol. 4, p. 62, 1895.

’’ The Magnesian Series of the Northwestern States. Geol, Soc. America Bull., vol. 6,

pp. 167-198, 1895.

Ulrich — Paleozoic Systems in Wisconsin.

77

top of the Dresbach sandstone. Besides^ the succeeding Jordan

sandstone is given a thickness of 200 feet and this of course drops

the base of the Jordan far down toward the top of the Franconia.

In fact it leaves between the Jordan and the Franconia only what

would be required by the 30 feet or so of limy beds that constitute

the typical St. Lawrence limestone. Moreover^ knowing most of

the localities mentioned by them I am certain that not only the

greater part of the shales but also some of the magnesian limestones

which they correlate with the typical St. Lawrence are really in the

Franconia. But all this is difficult to reconcile with their definite

statement (p. 172) that the beds exposed at the type locality

represent ‘ ^ the lower half of the formation. ^ ’ Evidently limy beds

of the Franconia were confused with those at St. Lawrence.

However erroneous some of the correlations of beds in the “St.

Lawrence” of Hall and Sardeson’s 1895 paper may be the fact of

greatest importance in this connection is the implied and partly

carried out intention to expand the stratigraphic meaning of the

term St. Lawrence.

The first definite and unqualified change was made about 1910

when the U. S. Geological Survey Committee on Geologic Names

decided how the Cambrian and Lower Ordovician rocks in the

Upper Mississippi Valley were to be divided and what names they

should bear. These decisions appear in two Water Supply Papers,

No, 256, by Hall, Meinzer, and Fuller, and No. 293 by Norton and

others, published in 1911 and 1912 respectively. Now for the

first time since 1888— -if we disregard the work of Hall and Sarde-

son mentioned in the preceding paragraph— the St. Lawrence lime¬

stone becomes a formation and is increased in thickness from the

preceding maximum of 30 feet to over 200 feet. This expansion

was brought about mainly by incorporating the underlying Fran¬

conia sandstone which had been named in the meantime. No rea¬

son is given anywhere in these two papers for this unwarranted

and really quite inexcusable proceeding. Even though it was not

yet known that still another formation- — since described under the

name Mazomanie sandstone — wedges in from the east between the

base of the St. Lawrence limestone and the top of the Franconia,

anyone well acquainted with the field relations of the concerned

stratigraphic, faunal, and lithologic units could hardly have failed

to recognize the thorough distinctness of the Franconia on the one

hand and the beds above it on the other. The top also of this

expanded St. Lawrence is an unnatural and often indefinite bound-

78 Wisconsin Academy of Sciences, Arts, and Letters.

ary. Evidently it was drawn somewhere about the transition

from the limy Lodi shale member to the Norwalk sandstone mem¬

ber of the Trempealeau formation, this position being indicated by

the thickness (160 feet) assigned to the overlying ‘^Jordan” sand¬

stone and the fact that in the columnar sections the St. Lawrence

is extended only to the top of the limy beds.

Even if the St. Lawrence formation as defined in these Water

Supply Papers were a naturally bounded and permanently desir¬

able stratigraphic unit I see no warrant for this expansion of the

St. Lawrence except the suggestion by Winchell in 1888 and the

paper by Hall and Sardeson, both of which are referred to above.

But Winchell himself failed to carry out his suggestion, and Hall

and Sardeson did so under obvious misconceptions as to the corre¬

lation of the concerned beds. Besides, the lower part of the ex¬

panded St. Lawrence had in the meantime been named by Berkey ;

and after the term Franconia sandstone had been proposed all

possible excuse for extending the application of the name of the

superposed smaller and in every respect less important strati¬

graphic unit so as to cover and completely eliminate Berkey ^s

Franconia had been forfeited. Moreover, experience in the field

has proved conclusively that the lower two-thirds or more of Hall

and Norton ^s St. Lawrence formation — in other words, the Fran¬

conia — is as useful and as clearly defined a formation as any of

the subdivisions of the Upper Cambria series in the Upper Missis¬

sippi Valley now recognized.

In 1914, two years after the appearance of Norton’s paper, Wal¬

cott® published the sequence and preliminary classification of the

Upper Cambrian formations in Wisconsin and adjoining States to

the west that had been worked out the preceding field season by

E, 0. Ulrich, In this classification the Franconia is recognized and

the St. Lawrence formation of Hall and Norton was not only re¬

stricted to beds above the Franconia, but its definition was again

modified by extending its upper boundary so as to include a sand¬

stone — ^here distinguished as the Norwalk sandstone member— that

had been improperly referred to the lower part of the Jordan

sandstone by the Minnesota and Iowa geologists. The Norwalk

sandstone member is not present in the section at Jordan, Minn.,

but where both are found, as is generally the case in western Wis¬

consin, the two are usually separated by a well-defined boundary.

® Smithsonian Mis. ColL, vol. 57, p. 354, 1914.

Ulrich — Paleozoic Systems in Wisconsin.

79

Although this classification was regarded from the beginning

as a merely preliminary effort to bring order out of the confusion

that had previously prevailed in the published accounts of the

Cambrian sequence in the Upper Mississippi Valley it has in the

main proved a serviceable and reliable guide in the work during

the past nine years. With unimportant modifications, except in

the matter of much added detail that had accumulated in the mean¬

time, the same classification was used in 1919 by Twenhofel and

Thwaites.^ Also in 1920 and 1922^*^ by the writer, who on these

occasions introduced a new formation — the Mazomanie sandstone

— that had previously been regarded as the eastern representative

of the Franconia but is now known to be a distinct formation that

wedges in from the east in Wisconsin between the top of the Fran¬

conia and the base of the St. Lawrence limestone.

Knowledge of the lower Paleozoic section in Wisconsin and

neighboring states having reached the stage where I feel war¬

ranted in presenting my final opinions concerning the formations

and their classification, it seems desirable and important to subject

the names of the formations to critical scrutiny as well as the strata

themselves. The only nomenclatural change of importance that has

been suggested by these inquiries is the proposed substitution of

the new name Trempealeau formation for the beds that I had pre¬

viously called St. Lawrence. The need for this change was first in¬

dicated when I sought an appropriate geographic name for the

member to which Twenhofel and Thwaites had applied the desig¬

nation ‘‘Calcareous beds” and found that the concerned member

is actually the bed to which Winchell had originally applied the

name St. Lawrence limestone. Under the circumstance I felt un¬

willing to propose a new name for this member ; and this unwilling¬

ness persisted even when it became obvious that my refusal neces¬

sitated the proposal of a new name for the formation of which

the St. Lawrence limestone is a member. On the other hand, how¬

ever, the proposal of a new name for the broader unit clears up

most of the confusion that at present attends the term St. Law¬

rence. That this confusion is very real is most clearly indicated

in tabular form.

® Twenhofel, W. H., and Thwaites, F. T., The Paleozoic section of the Tpmah and

Sparta quadrangles. Jour. Geol,, vol. 27, p. 616, 1919.

Ulrich, E. O., Major causes of land and sea oscillations. Washington Acad. Sci.

Jour., vol. 10, pp. 74-76, 1920, and reprint of same in Ann. Rept. Smith. Instit., pp.

333-335, 1922.

TABLE SHOWING VARYING USE OF THE TERM ST. LAWRENCE FROM 1874 TO 1922.

80

Wisconsin Academy of Sciences, Arts, and Letters.

- J,

S3 A

c^Ph

0OU8JMB'-J -qg

TO

xioao -^S

I ^

uiaoj xioJQ •:}g

•g -g

< o c-a

& 8 ^ 1

^ o s

[*< Q W

’Si

^•1

og

o> 8

UBIStJBZQ

HBUqUIBQ

♦In the area covered by Twenhofel and Thwaites the Mendota is lacking^.

Ulrich — Paleozoic Systems in Wisconsin,

81

The foregoing table shows that in 1874-7 Winchell mistakenly

applied the terms St. Lawrence and Jordan to three distinct pairs

of formations belonging to three systems. Only the Cambrian pair

— originally described in 1874 — is correctly named, the other

usages of the terms by him being based on misapprehended cor¬

relations. The table indicates further that Winchell, and also Cal¬

vin, never included any other Cambrian beds in the St. Lawrence

limestone than the calcareous 15-30 foot lower member of the

Trempealeau formation (“St. Lawrence formation” of Ulrich,

Walcott, Twenhofel an^ Thwaites, not Hall, Norton, or Winchell)

to which I now propose to confine the term. The third fact shown

by the table is that the St. Lawrence formation of Hall and Norton

differs widely from the St. Lawrence of Walcott, Ulrich, Twen¬

hofel and Thwaites (1914-1919) in that it is extended downwards

to the top of the Dresbach sandstone and upward only to the base

of the Norwalk sandstone member of the Trempealeau formation

(St. Lawrence of Ulrich et ah).

In view of the consistent use of the term St. Lawrence by Win¬

chell in 1874 and 1888 and by Calvin in 1895 I maintain that it

was both unwise and contrary to the rules of stratigraphic nomen¬

clature to introduce the great expansions of its meaning that were

given it by Hall in 1911, Norton in 1912, and Ulrich in 1914. It

would have been better to disregard WinchelUs term entirely as

a formation name and give a new name to the broader unit of

which the original St. Lawrence is a part. The main reasons why

I did not do so in 1914 were (1) the fact that the United States

Geological Survey had previously adopted the term in the wide

significance given it in the two publications by Hall, 1911, and

Norton, 1912, and (2) the obvious need of further field studies

before a final classification and nomenclature might be warranted.

But the Franconia, proposed by Berkey in 1897, had proven too

good a formation to be ignored, so I defined its boundaries as well

as I could in that early stage of my investigations of the Cambrian

in Wisconsin and Minnesota and restricted the St. Lawrence to

beds above the Franconia.

In 1916 or 1917 another complication of the St. Lawrence ques¬

tion was introduced by Shipton,^^ who proposed the new, though

preoccupied term Sparta shale for the same beds to which Hall,

Norton, and others had applied the name St. Lawrence. And the

Shipton, W. D., Proc. Iowa Acad. Sci., vol, 23, p. 142.

&£OLO&/C^COLi/MhL

pyzso^S'/A/

iv/TH CO0KELATIOA/ rEfSLC £S2S. ZliS. PALCQMfC £S^

G£-A/£/?Al riM£ ^C^l£

t-V'jESr tH^^/SCOA/S/f</ SASr HP/SCOA/S/Af

r/AfiT

H«rr $*>'^.sco^s/Ar

t^^AfOOA/^^Z/S/

Maysi^/yie (Oh/o 4^S0ff. /s. &

shj

(^baen/)

CAbsen/ ?)

Eden (Oh/o %JOOf/: sb. S: /s.)

(Abse/ib)

(Absent ?)

Trenton (A(Y &Po- 400- GOO ffJs) <3a/ena ch/. (£50 ft i/ic. Prosser Js.)

Wmmst^z/ch (Mo. /OO ft. is.)

Decor ob (/o$*'a/ &Off sb& fs.)

(Absenf)

Ga/enodof. (/00-200ff.)

(Absent)

Decorah (SO"^ ft. sh & ts.)

HAytertotvntAtyd Tenn. 8-200 ft. ts)

P/attei^/tte ts.(7S ft)

Lot*^r///e(AtyaAp/o. 600 ft h.)

(^/oerfftue dot /5 ft

Upper Bi/ ff dot 02 ft

totter Bti/e otot 25f/.

£. oi*'er Buff dot 28 b.

''bazyan (At V & App. 7000 ft )

(Absent)

Jocb/m (Afo. too ft. ts.)

Ot Peter (Af /bn tOOft)

^ Peter ss. & sb. CO-JJ2ft.)

dt Peter ss. 3sb (O-JOO

e^^erton (ArA. t20ft ts. &ss.)

H'/tb Hmgs P/Ver ss. anct

Bneeots- ts. men/bers.

(Absent)

Upper Canad/an (App. 2000ft. ts. )

SbaAopee dot (70 ft)

SboAopee dot fO-tOOM

M/ddte Cpnaettan (App SeOAta. 2SOO/t)

(Absent)

toH/erConoctton fPcr. 700ft ts.)

(Absent)

Upper OsarA/en (App tSOO ft. dot)

Oneota dot (0-200 ft.)

Oneota dot. (0-/00 ft)

M/ddte O/sarAtan (App. 2000 ft dot)

(Absent)

Loiter QeonA/bnfPb: BA/a. 2JOOftdot)

Mad/son ss. (AO ft)

(Absent)

Mad/son ss. (dOft)

Atendota dot. (20 ft)

Deu'/ts toAe ss (tOOPtb

Jordan (Af/bn BOftss)

Jordan ss. (7d ft.)

Tre/npeateoa (Af/s. t2Sft. sb.Sk/s.)

(=St. Lat$rrence of U/r/cb, tPtA)

Ahm^atAss. member (SO ft)

lodt Sbo/e (soft)

St. laterence ts. (2Sft)

Sbe/e (tocot 2 Oft)

Jordan ss.

(Oenero//y Absent)

(Absent)

Lodt Sbate (0-2Sft.)

StLokt^rence (0-20 ft.)

Sbate (tocot)

Maxom'an/e OV/'s. /SO ft dot ss.)

(Absent)

Afazoman/e ss (tOO -t6S}

trancon/'a (At/bn tZS t ft

prss. sb. & ts.)

Upper Breensond (Sf-TO)'^

Ye//o(^ ss. (AO -SO)

Lot^er (reensond (AO)

M/coceous sh (IS)

fronton ss. nremberUS)

f20

to

f70

Usuo/ty Absent

Dreebacb (M/nn )

Dresboch ss. (AO-2SOft.)

Dresbocb ss. (AO -tdOfti

ff EaaCtatre (id/s.)

Bou Cto/re sb. (BOO-JOOff.)

£paCtoJresb (SSOft)

Mt. Simon (Id/'s )

Af/dd/e & Lon/'er Combr/on (PocAy

Mts. (2.000 ft ss. sh. & ts.)

Mt S/mon ss. ( (00 - 200 ft)

Mt S/mon ss. (700*- ff. )

(Absent)

Cbepuanseaan ss. tOOO ft

Deut/s /stand ss. JOG ft

Or/enta ss. SOOO ft

Amn/con Sbate and ArAase SOOO ft

£//een Sandstone 2000 ft

Preata/ Sandstone (2^ OOO ft

A/onesacb Sbate /20-J50 ft

Outer Cong/omerate 300 -/200ft

Bosat/s; <//obose, rbyot/te, gabbro, te/s/tes^ cong/omerates. etc

(tZOOO to SO, OOO ff)

Congtomera/tes and Qaortz/tes (22S-SOO ft)

Ty/er format ton, ^m/'ca and c/oy o/a/es^ and gray/tracAes (7,000 - //OOO ft)

IronH/0od forn?at/on, Prrug/noas cherts and cberty /ran carbonates (800-/0(00/7)

Pg/ms form ot/on (c/gy state and gaar/s/te AOQ-BOOf/)

Bad Ptirer do/om/te (200 SOO ff)

Laan&nf/an - gran/tes, syen/feSj pabbros, and gne/ssa/d egat/ra/enfs.

Peai>*raf//7 — pu-eensfones (scbts/cse baso'/ts), anct ^reen scb/sts.

84 Wisconsin Academy of Sciences^ Arts, and Letters.

case became even worse confounded when Keyes^^ a few months

ago made the St. Lawrence in its typical exposure in Minnesota the

same as the Oneota and the Jordan sandstone the same as the

New Richmond sandstone, renamed the Franconia (Albin shale),

also the typical St. Lawrence (Allamakee dol.), and proposed the

name Waukon sandstone for what Hall, Sardeson and Norton had

called Jordan.

Considering all this confusion — nomenclatural and stratigraphic

— and wishing at the same time to preserve the credit really due

to N. H. Winchell for naming two Cambrian horizons that are

well marked both lithologically and faunally not only in Minne¬

sota but also in the adjoining states of Wisconsin and Iowa, some

radical changes in names and in the definition and correlation of

the concerned stratigraphic units seem necessary. After thorough

investigation of the facts and much thought the best and only just

solution of the various difficulties and issues seems to be the one

here followed and in part newly proposed: namely

(1) The restriction of the name Jordan sandstone to the wholly

unfossiliferous, probably continental deposit of usually light gray¬

ish sandstone which occurs at Jordan, Minn., and is elsewhere in

the Mississippi Valley easily recognized by its lithologic character

and its position in the section between the top of the fossiliferous

marine Norwalk sandstone member of the underlying formation

and the more or less unconformable base of the often similarly

sandy initial deposit of the succeeding Lower or Upper Ozarkian

formation.

(2) The restriction of the term St. Lawrence, in the original

form of St. Lawrence limestone and not St. Lawrence formation,

to the well-characterized and widely recognizable calcareous zone

to which this term was originally applied by Winchell and to

which it was again confined by him in 1888.

(3) The proposal of a new name, the one chosen being Trem¬

pealeau formation, for the formation to which Ulrich in 1914 ap¬

plied the term St. Lawrence formation and of which the typical

St. Lawrence limestone usually constitutes the basal member. The

Trempealeau thus embraces the beds between the top of the Fran¬

conia formation and the base of the true Jordan sandstone.

^ Keyes, Charles, Terranal differentiation of Iowa Cambric succession. Pan-Ameri¬

can Geologist, vol. 38, 1922, pp. 313—326.

Ulrich — Paleozoic Systems in Wisconsin.

85

The Trempealeau Formation and Its Subdivisions.

The Trempealeau formation is well displayed and in fairly typi¬

cal composition in Trempealeau Bluff on Mississippi River. Other

good sections may be seen (1) at Norwalk, (2) in and below Beans

quarry, 2 miles northwest of Tunnel City, (3) in the bluff south¬

east of Mazomanie, (4) at Hillside just south of Wisconsin River

from Spring Green, (5) in the bluffs on both sides of St. Croix

River at Osceola, and many other places in Wisconsin, Most of

the beds are exposed also at Lansing, in Iowa, and at Winona,

Redwing, and Stillwater, in Minnesota.

There is considerable variation in the character and sequence of

the component beds of the formation from place to place. Thus

in eastern Wisconsin the upper (Norwalk) member is commonly

and perhaps always absent. But the yellow calcareous shaly Lodi

member is generally present and may also be called the most char¬

acteristic part of the formation. The same might be said of the

next underlying St. Lawrence limestone or dolomite member, but

in the middle and northern parts of the State this member is en¬

tirely wanting in many places, or it is so much altered by addition

of relatively coarse quartz sand that the recognition of its zone

is rendered difficult and uncertain.

In calling the Lodi member a shale it is to be understood that

as defined by me the word shale applies generically to a rock of

very fine grain, thinly laminated construction and containing some

clay but not necessarily in preponderant quantity. All of these

Cambrian shales consist mainly of finely divided siliceous matter.

Subdivisions of the Trempealeau Formation.

In its fullest development the Trempealeau formation is divisible

into four lithologically and faunally distinct members. Locally

one or two of these subdivisions might be mapped separately, but

as a rule the topographic conditions are such that only the lower

magnesian limestone member lends itself readily to such separate

treatment. Names for at least three of these members are desir¬

able if only for purposes of discussion. They differ notably in

geographic distribution, and such facts are neither easily nor

clearly to be brought out in descriptions of local stratigraphy

without definite names for each of the several members.

86 Wisconsin Academy of Sciences, Arts, and Letters.

Norwalk sandstone member. — The thickest and therefore per¬

haps the most important of these members is the one at the top for

which the term Norwalk sandstone member is proposed. As a rule

it consists of fine-grained grayish sandstone, sometimes nearly

white and often with a yellowish or brownish tinge, commonly

rather massive in its upper two-thirds and more or less thin bedded

and in even plates 1 to 6 inches thick in its lower third. In thick¬

ness this member varies from 0 to over 40 feet. At Norwalk,

where as usual it is in contact above with the Jordan sandstone,

it is about 35 feet. In Beans quarry, near Tunnel City, it is 43

feet thick, at Osceola, Alma, and in the lower Beef Valley about 50

feet. From these maximum developments the member diminishes

southeastwardly to 25 feet at Ironton and less than 7 feet in the

vicinity of Mazomanie. It has not been observed to the east of

Cross Plains, in which region the Trempealeau formation is ter¬

minated above by the Lodi shale member.

A large and varied fauna has been collected from the Norwalk

sandstone member; and most of the species are strictly confined

to its zone. Locally the sandstone is quite bare of fossil remains,

but recognizable specimens of some of its characteristic species

have been found in many places. At most of these localities the

fossils occur mainly or solely in the thin-bedded lower part, but

the reverse is the case at Norwalk where many were found in the

upper third and very few in the beds beneath. Osceolia osceola,

Saukia pyrene, S. leucosia, Illaenurus qnadratus, Eurekia eos, and

Sinnopea sweeti are the species most commonly found.

Lodi shale member. — This term is proposed for the usually yel¬

low calcareous shale-like sandstone that lies between the Norwalk

sandstone and the St. Lawrence limestone. This shale member is

widely distributed, the outcrops being everywhere recognizable

from Stillwater, Minn., and Osceola, Wis., on the north to Spring

Green on the south and the vicinity of Madison on the east.

Locally the characteristically yellow shale is interbedded with

purple shale, as in the vicinity of Mazomanie, or with layers of

sandstone and in other places with dolomitic limestone, but even

without considering the fossils there is seldom any difficulty in

recognizing the Lodi shale. The thickness of the member rarely

falls under 15 feet, with approximately 25 feet as the average and

50 feet as the maximum.

In view of the frequent miscorrelation of the underlying St.

Lawrence limestone with the Mendota dolomite it is important to

Ulrich-Paleozoic Systems in Wisconsin.

87

note that in the 4 or 5 miles distance between Farwells Point on

the north side of Lake Mendota and Pheasant Branch at the west

extremity of the lake certain beds enter the section that are not

present at Farwells Point or near-by in Maple Bluff. Namely, at

Farwells Point only two feet of St. Lawrence limestone separate

the Mendota dolomite from the underlying top of the Mazomanie

sandstone. In the quarry at Pheasant Branch, on the contrary,

the Mazomanie which outcrops on the lake shore is succeeded, first,

by an undetermined but small thickness (less than 10 feet) of soft

shaly and glauconitic sandstone beds, second, by 12 feet of St.

Lawrence limestone, and, third, by at least 5 feet of yellow and

purple calcareous sandy shale that not only has the lithological

characters of the Lodi shale but also some of its characteristic fos¬

sils. Above this shale the section is not exposed for about 35 feet,

when a three-foot ledge of coarse white and brownish sandstone

that resembles the Jordan but more probably belongs at the base

of the Oneota dolomite comes to the surface. The covered inter¬

val may be occupied by either the Madison sandstone or the Men¬

dota dolomite or by thin representatives of both, or, as seems the

more probable, by Jordan sandstone only. In the near-by Leith

well only one limestone (evidently the St. Lawrence) is found be¬

neath the Oneota.

The Lodi shale member usually is fossiliferous, and its fauna, so

far as known, is almost entirely confined to this member. As the

commonest and most characteristic of its species we may cite

Dikelocephalus minnesotensis, Saukia crassimarginata, S. lodensis,

and Aglaspis harrandei. With these, and usually more abundantly

than anything else, occur species of brachiopods provisionally

identified with Westonia stoneana and W. aurora.

St. Lawrence limestone or dolomite (as originally defined). — ^As

stated on a preceding page, some distinctive name is desirable for

the bed of highly magnesian limestone that commonly underlies

the Lodi shale member in Wisconsin, Minnesota, and probably also

in northeastern Iowa. Further, as the name St. Lawrence lime¬

stone proved on visiting the type locality to have been based by

Winchell on the very bed for which a name would now be a useful

aid in describing the several divisions of the Trempealeau forma¬

tion it has seemed eminently proper to adopt Winchell ’s term in

the restricted sense in which it was originally used. There is no

doubt whatever in my mind that the ‘ ‘ St. Lawrence limestone ’ ’ de¬

scribed by N. H. Winchell in 1874^® is the bed here referred to as

88‘ Wisconsin Academy of Sciences, Arts, and Letters.

underlying the Lodi shale member of the Trempealeau formation.

As described by Winchell the St. Lawrence limestone member at

the type locality consists of grayish magnesian limestones, mottled

with red and yellowish brown blotches and more or less profusely

speckled with the green grains of glauconite, in layers 2 to 18

inches thick and aggregating a total thickness of between 14 and

15 feet. Under it is a shaly bed, 18 feet of which was observed

by W. 0. Hotchkiss and the writer in 1915 in a ravine north of

the quarry. The latter exposure did not extend down to the top

of the Franconia which probably underlies the Trempealeau in

the valley of Minnesota River as elsewhere. Whether the Fran¬

conia is present here or not the observed shale bed is lithologically

at least like the bed that is found at many places in Wisconsin

at the base of the Trempealeau and usually beneath the calcareo-

magnesian zone that is correlated with the typical St. Lawrence

limestone or dolomite.

Regarding the propriety of the latter correlation it would be

amply warranted by the similarity of lithologic characters alone —

especially the likeness in the sequence and thickness of the litho¬

logic units concerned. But we have corroborating fossil evidence

that with the lithologic evidence of the rocks themselves settles

the question beyond all reasonable doubt. Fossils are seldom plen¬

tiful in the St. Lawrence limestone and only rarely are they in a

good state of preservation. Despite these disadvantages a consid¬

erable fauna from this zone has finally accumulated. More im¬

portant is the fact that this fauna includes a number of character¬

istic species that are sufficiently distinctive to be recognized even

in poor condition. Nearly always this zone affords some brachio-

pods, particularly a form of Billingsella that we have not yet suc¬

ceeded in distinguishing from B. coloradoensis — a. common lower

Franconia shell. Also a variety of Dicellomus politus, a species

otherwise not found above the basal layers of the Franconia. With

these usually is the Finkelnhurgia osceola corrugata which so far

seems confined to this zone. Perhaps a surer but rarer guide fossil

is a large trilobite that may be the species described by Walcott

under the name Dikelocephalus vanhornei. Specimens of all of

these four species rewarded a half hour’s search in the quarry at

St. Lawrence, Minn.

In Dane and Iowa counties, Wisconsin, the St. Lawrence dolo¬

mite is notable for certain interesting additions to its more usual

Second Ann. Rept. Geol. and Nat. Hist. Sur. Minnesota, p. 152.

Ulrich — Paleozoic Systems in Wisconsin.

89

faunaj referred to and discussed by Ulrich in 1916d^ These addi¬

tions— in all 10 species, most of them obviously primitive gastro¬

pods — make up what may be called a distinguishable but unde¬

niable preoccurrence of species typically developed in the much

younger Lower Ozarkian Mendota dolomite. Many even more

striking instances of recurring faunas are now known, so that no

particular importance is to be attached to this at first perplexing

case.

In southern and western Wisconsin the zone of the St. Lawrence

limestone is seldom absent. Where present it is always more limy

than the superjacent and interjacent beds; and as a rule it is un¬

questionably indicated by thick, often quarried layers of character¬

istically colored crystalline magnesian limestone. No other bed

that resembles it at all closely occurs in the prevailing sandy Upper

Cambrian series in the Upper Mississippi Valley. However, it

varies greatly in thickness from place to place. In the vicinity of

Sparta it usually is only about 2 feet thick, between Mazomanie and

Black Earth 3 or 4 feet, at Spring Green about 6 or 7 feet, at

Pheasant Branch (west end of Lake Mendota) 12 feet, at Farwells

Point, on north shore of Lake Mendota, only 2 feet. Apparently

it is entirely absent to the north and east of the Baraboo Range.

In the bluffs along the Mississippi its development is variable. It

was not observed in the section at La Crosse, but to the north of

that place, as far at least as Stillwater, Minn., its zone is generally

recognizable.

Basal shale. — ^The base of the Trempealeau is often a shale or

shaly sandstone with considerable greensand and occasionally thin

layers of dolomitized sandstone. At the bottom usually there is a

thin layer of sandstone conglomerate. In thickness it varies from 0

to 15 or even 20 feet, the latter maximum being found on the east

side of Blue River Valley about 4 miles southwest of Muscoda.

This place also is the only one at which satisfactory fossils were

procured from this shaly bed. The fauna as indicated by this

collection of seven or eight species comprises three or four seen

nowhere else and two small species of Saukia that can be compared

only with S. pyrene and S. leucosia, two species known elsewhere

only in the Norwalk member of the formation.

It is not at all certain that the shaly bed at the base of the

Trempealeau is always of the same age as the one found in this

Correlation by displacements of the strandline. Geol. Soc. America, Bull., vol. 27,

p. 477, 1916.

90 Wisconsin Academy of Sciences, Arts, and Letters.

position in the valley of Blue Eiver. Indeed, this correlation is

decidedly doubtful in the case of the basal Trempealeau shale in

sections where, as at Norwalk and below Beans quarry near Tun¬

nel City, the St. Lawrence dolomite member is either absent or

unrecognizable. In such places the basal shale may well be of the

age of the St. Lawrence dolomite. Until this question can be fur¬

ther investigated and determined it is thought advisable to defer

proposing a special name for the basal shale.

The Feanconia Formation and its Subdivisions

The Franconia formation is divisible everywhere in the western

half of the State into four, five, or six members, but these differ

regionally so much in character and sequence that it is difficult

and as yet impossible to correlate exactly those prevailing in the

northwestern quarter with those found in the southwestern quarter

or in either of these with those seen in the middle part of the

State.

In the area lying to the south of Trempealeau we can usually

distinguish two main greensand zones, one — 50 feet to 70 feet

thick — at the top of the formation; the other — 30 feet to 45 feet

thick — in the lower half. The upper of these greensand zones is

characterized faunally by several species of Ptychaspis, among

which the most notable is P. miniscaensis. The lower greensand

contains few fossils.

Between the greensand members is another usually well defined

zone — 10 feet to 40 feet thick — that differs from the adjoining

beds in consisting mainly of yellowish platy sandstones of which

occasional layers often are fiUed with highly characteristic re¬

mains of tribolites. These include Ellipsocephalus curtus, Chari-

ocephalus wJiitfieldi, various species of Ptychaspis, and species of

the new genus Wilburnia, Walcott ms., including the typical form of

W. diademata (Hall).

Under the lower greensand is a fourth member that is dis¬

tinguished from the others by its shaly and thin-bedded micaceous

sandstone and sandy limestone. The latter occurs in the lower

third of this member and usually is marked by an abundance of

valves of articulate brachiopods. These belong to two or three

species of Eoorthis, of which E. remnicha probably is the most

characteristic. Many small trilobites occur with these and in the

shaly sandstones above.

Ulrich — Paleozoic Systems in Wisconsin.

91

Finally at the base of the formation is a 2 to 15~foot bed of

reworked Dresbach sand. Usually this forms a single massive ledge

and bench at the top of the Dresbach bluffs^ commonly holding this

position because its top is more or less silicified and therefore more

resistant than the relatively incoherent mass of sandstone beneath

it. It contains some highly characteristic fossils which distinguish

this initial deposit of the Franconia from all preceding and suc¬

ceeding beds in this region. Some of the more useful of these fossils

are mentioned on a following page.

Of these five members only the lowest and possibly the shaly

micaceous bed above it are clearly recognizable by their respective

lithologic and faunal characters in the Franconia as developed in

the tops of the mounds in Adams County. In these mounds the

soft greensand members are represented, as is clearly established

by their fossil contents, by much harder red and gray sandstones ;

but the intervening yellowish sandstone member is not easily dis¬

tinguished from the beds on either side of it. However, the sand¬

stones that are believed to correspond to the upper part of the

intermediate member form unusually massive, coarsely grained and

but sparingly fossiliferous ledges.

In the northwestern quarter of the State, speaking particularly

of the area between Mondovi and Alma on the south and St. Croix

Falls on the north, no two of the many Franconia sections are

strictly alike. And yet a general similarity in both the character

and the sequence of the beds is manifestly maintained. Compared

with the Franconia in the southwestern quarter only one of the

members usually determinable there is clearly identifiable in the

northwest. The zone referred to is the yellow sandstone member.

In places, particularly in the area between Hudson and Menomonie,

this member maintains in fair approximation the color and char¬

acter of bedding that distinguish it in the region between Sparta

and Lavalle. However, followed from Hudson both to the north,

as at Franconia, Minn., and to the southeast, as in Beef Eiver

valley between Alma and Mondovi and also at Durand, nearly the

whole of this member becomes so strongly charged with glauconite

that were it not for its abundant and highly characteristic fossil

contents it would hardly be distinguished from the associated green¬

sand beds.

In the northwestern quarter of the State there also, is a fairly per¬

sistent “Lower Greensand zone. At its top is a bed of variable

92 Wisconsin Academy of Sciences, Arts, and Letters.

thickness (2 to 15 feet) that contains some magnesian limestone

layers with dismembered plates of crinoids or cystids and other

fossils that is recognized as far south as La Crosse. At this latter

place, however, it rests on the initial Franconia deposit of re¬

worked Dresbach sand containing Camaraspis convexus (Whitf.).

At Hudson, on the contrary, it is underlain by about 27 feet of soft

sandstone more or less profusely charged with greensand ; and this

ought to correspond to the “Lower Green sand” of the Sparta

region. Granting this decidedly questionable conclusion it would

follow that the “Micaceous shale” of the Sparta region is unrepre¬

sented in the sections at Hudson and elsewhere in the area to the

north of La Crosse. It would follow also that the lower half of the

Franconia contains two distinct limy zones, the one in the north

containing crinoidal fragments and lying above the ‘ ‘ Lower Green¬

sand,” the one in the south without crinoidal remains and occur¬

ring beneath or at the base of the “Micaceous shale.” But these

doubtful correlations can as yet be viewed only as provisional sug¬

gestions to be entertained until proved or disproved by the results

of detailed comparisons of sections and fossils now being carried on.

At present it seems more probable that the limy zones are the same

and that the “Lower Greensand” of the south, rather than the

^ ‘ Micaceous bed, ’ ’ thins out or becomes unrecognizable to the north.

As in southwestern Wisconsin, so also in the sections at Fran¬

conia, Minn., Hudson, Durand, west of Mondovi and as far south

as Trempealeau in Wisconsin, the base of the Franconia is made

by a fossiliferous bed that consists mainly of reworked Dresbach

sand. But in the mentioned latter places the initial Franconia

deposit contains more glauconite. Its fossils also are quite differ¬

ent from those found in the basal Ironton sandstone member of

the formation in the southwestern quarter of the State. In the

latter region the fossils in the basal sandstone consist almost en¬

tirely of trilobites, most of which have been found only in this

bed. On the other hand, the fauna of the basal sandstone in the

mentioned northern localities consists entirely of a few species of

inarticulate brachiopods. Of these brachiopods a species of Dicel-

lomus, usually referred to D. politus — a common fossil in the Eau

Claire shale — is most abundant. A small elongate and very narrow

species of Lingulella is more characteristic of the zone.

These regional variations in the composition of the lower third

or half of the Franconia, when all is considered, can be interpreted

only as indicating differential oscillation of surface and conse-

Ulrich — Paleozoic Systems in Wisconsin,

93

quent varying local migrations of the strandline during the early

stages of the Franconia age. Abundant data indicating the char¬

acter and order of occurrence of these oscillations and the areas

affected by them are available, but their citation and discussion

would require more time and space than the present opportunity

warrants.

The Ironton sandstone member, — Of the various members of the

Franconia formation above briefly discussed only one is named on

this occasion. The member so distinguished is the basal sand¬

stone of the formation in the southwestern quarter of the State.

The Ironton sandstone member, as it is proposed to call it, has been

recognized and studied at many places in Sauk, Richland, Vernon,

La Crosse, Monroe, Jackson, Adams and Juneau counties. So far

as observed in these counties the bed varies in thickness from

about 2 feet to 12 or possibly 15 feet. At Ironton, the type local¬

ity, it varies from 5 to 10 feet. The top of the bed is even, the

rapid inequalities in thickness being due to unevenness of its

base. As a rule it forms the slightly hardened top of Dresbach

sandstone bluffs and for this reason has usually been regarded as

the terminal bed of that formation.^^ However, it is now quite

clear and commonly accepted that it is of later date and, in fact,

the initial deposit of the Franconia.

The Ironton member is composed mainly of reworked washed

and relatively coarse residual grains of Dresbach sandstone, the

surface of which had previously been subjected to subaerial leach¬

ing and wear. The line of the break between the two formations

— Dresbach and Franconia — lies at the undulating plane where

washing and sorting of the loose quartz grains of the underlying

formation is first indicated. In other words, the Ironton sand¬

stone member extends downward to the lowest plane indicating

reworking and redeposition of the weather-loosened top sands of

the underlying Dresbach formation. Commonly the new deposit

includes a few grains of glauconite and other material that is not

present in the undisturbed underlying beds of Dresbach sandstone.

But to make sure of the identification of the Ironton member

it is advisable to search for its characteristic fossils. In the

Dresbach proper no organic remains — except perhaps worm bur-

15 Twenhofel and Thwaites in their paper on the Paleozoic section in the Tomah

and Sparta quadrangles, Jour. Geo., vol. 27, p. 616, 1919, apply the term “Wormstones”

to relatively firm sandstone at the top of the Dresbach that may be a part of the Ironton

sandstone member.

94 Wisconsin Academy of Sciences, Arts, and Letters.

rows — have so far been observed. The overlying basal sandstone

of the Franconia, however, only rarely fails to reward a few min¬

utes’ use of the hammer with indubitable evidence of the presence

of such remains. The most abundant and characteristic of these

are the nearly hemispheric cephalic shields of several species of a

new genus of trilobites, one of which was long ago described by

Whitfield under the name Arionellus convexiis. As this trilobite

does not belong to either Arionellus or any other previously estab¬

lished genus it is proposed to change its designation to Camaraspis

convexus (Whitfield) Ulrich and Resser.

Physical Evidence of the Breaks Between the Paleozoic

Systems in Wisconsin

General discussion. — In Wisconsin, as indeed is the rule else-

v/here on our own and other continents, the physical evidence in¬

dicating long interruptions of the process of marine sedimentation

consists mainly of phenomena of stratigraphic overlap. By careful

comparison of the fossil contents of the successive beds and the

identification of the fossiliferous zones in other regions we finally

arrive at a fairly reliable conception of which stages of the com¬

posite geological time scale are represented by marine deposits in

a given area and which are not. The most striking and important

of the conclusions thus arrived at is that within the always rela¬

tively flat interior areas of the continents submergence by marine

waters and the consequent marine depositional record is greatly

inferior in volume of deposits and ages represented by these de¬

posits than in the submarginal Appalachian, Ouachita and Cordil-

leran geosynclines. The structure of the latter regions with their

long troughs and broader surface depressions naturally permitted

more frequent and longer-enduring submergences than could have

obtained within the more extensive, relatively stable and hence

much less folded interior areas. But we must not overlook the fact

that even in these submarginal troughs the stratigraphic record

of geologic events is far from complete. In these also the record

is broken at the usual horizons. Indeed, and despite the fact that

the record in these troughs is more complete — or rather less inter¬

rupted, the stratigraphic planes at which conclusive evidence of

retreat of the seas and emergence of the troughs is found are

only much more numerous and each no less clearly indicated than

in the geologic sections of the flatter interior areas. Consequently

Ulrich — Paleozoic Systems in Wisconsin,

95

we can not escape the conviction that complete emergence of the

area embraced in the present continent occurred at many times dur¬

ing the almost unbelievably long course of geologic history.

Though, as said, the evidence on which we base our conclusions

respecting the frequency and relative durance of periods when

the sea was withdrawn and land prevailed consists mainly of

phenomena indicating alternating retreat and advance of the

strandline and consequent absence of deposits found elsewhere and

overlapping structure of those present, it is yet true that even in

Wisconsin the sedimentary record is not wholly devoid of such

more convincing criteria of preceding land conditions as basal

conglomerates. These conglomerates vary greatly in composition

of material and character of the enclosed pebbles. Naturally the

character of both depends entirely upon the material available and

relative nearness to the shore. Obviously, under usual conditions

pebbles of large size can be distributed over the bottom of the new

sea only for a mile or less out from the shore. Here and there a

river may transport them for longer distances. The latter con¬

dition is suggested in some instances in Wisconsin ; and in every one

of these instances the pebbles consist of rolled quartz or quartzite

derived from distant exposures of pre-Cambrian rocks. Moreover,

their distribution is decidedly local.

Under more usual conditions the basal conglomerate consists of

material washed out of the subaerially decomposed top of the under¬

lying formation. If the latter consists of sand then the average

size of the grains of quartz is larger than in the undisturbed part

of the contributing formation. If the residual mantle of the re¬

submerged area comprises plates, blocks or concretionary masses

of respectively limy sandstone, limestone or dolomite, or chert, then

the character of the basal conglomerate, or perhaps more properly

the initial deposit, of the succeeding formation is modified accord¬

ingly. Nearly always the initial deposit includes laminae or may

be quite unstratified fine material transported by temporary sus¬

pension in the invading waters.

However, in many places the initial deposit contains nothing

having any connection with the weathered residual mantle of a

preceding land age. An excellent illustration of this condition is

found in the large limestone quarry at Darlington. Here there is

unquestionable evidence of a land stage that broke the continuity

of Black River limestone deposition. The break was long enough

96

Wisconsin Academy of Sciences, Arts, and Letters.

to permit subterranean solution and excavation of a cavern at

least 25 feet wide and 20 feet deep in the limestone previously laid

down here. Then the cavern was filled with fossilifrous shale and

limestone. Not a trace of this filling or of its fossils is notable in

the quarry face 30 feet to either side of the cavern. Beyond that

£ro5m c/iamneh in basal Gale/ra and top cf F/atteyi/fe

or Seloit /mestone. Dar///7jtan, JVisco/jsift

distance the layers of limestone seem so perfectly conformable to

each other that none would suspect that one of the bedding planes —

indeed it is one of the least evident- — marks a time when this area

was land and subject to surficial and subterranean erosion. Cases

like this should make one pause before declaring in error another

who thinks he sees a stratigraphic hiatus between two apparently

conformable layers of rock.

Nor is the relative conspicuousness of the break any reliable in¬

dication of the time represented by the hiatus. The time may be

immeasurably greater than in the case just described, and yet it

may require very close scrutiny of the purely physical evidence to

determine precisely where the break occurs or, indeed, whether any

at all occurred. Of course, with the aid of the fossil evidence not

only the actual presence and precise location of the break often is

quickly established, but the fossils also give the best measure of

the time represented by it.

The Bevono -Silurian hreak. — So far as known this contact is

everywhere unconformable, meaning by that only that the sea was

. Ulrich — Paleozoic Systems in Wisconsin.

97

withdrawn at the close of the last of the Silurian deposits and

that resubmergence of parts of the continent at the beginning of

Devonian time left a bounding break of undetermined time value

between the youngest of the Silurian deposits and the oldest of the

succeeding Devonian sediments. The plane of separation between

these is sharp and slightly undulating even when cementation has

taken place so as to make it appear as passing through a single

layer of rock — in such cases usually a limestone. In many places,

among them east Wisconsin, the Devonian overlap was delayed to

Middle Devonian time. Beds of the latter age there rest on some

late or middle Silurian formation. At Louisville, Ky., the adjacent

rocks of both systems are of limestone and the line of contact be¬

tween the two is sometimes so obscure that it is scarcely determin¬

able in a weathered slab two inches in thickness. However, the

lower side of this slab contains Niagaran species of corals, the upper

side Middle Devonian species. And yet nearly 2,000 feet of mainly

limestone deposits were laid down in the northern Appalachian

Valley during the time of the hiatus indicated by the fossils in the

upper inch of this slab and those imbedded in its lower inch.

In Wisconsin the Middle Devonian Milwaukee formation follows

the Waubakee dolomite, which is regarded as of late Silurian age.

No outcrop showing the contact of the two formations in which the

lower bed was positively identified as belonging to the Waubakee

has been observed. But it is quite probable that this contact is

exposed in a small quarry three-quarters of a mile northwest of

Port Washington. Whatever age may finally be assigned to the

lower formation at this place it is certain that the contact between

it and the uneven, clearly overlapping base of the overlying Mil¬

waukee dolomite is unconformable, with at least 1,000 feet of beds

missing that were laid down elsewhere.

The Siluro-Ordovician break. — That the contact of the Silurian

and Ordovician formations in Wisconsin and adjoining States

marks a stratigraphic hiatus of considerable magnitude and intro¬

duced great changes in the geography of the time is also shown

mainly by absence of certain elsewhere important late Ordovician

formations and by early Silurian sea transgressions of extraordi¬

nary extent and correspondingly great overlaps of deposits left by

them. These earliest Silurian transgressions occurred during the

Richmond or Lower Medina stage. In the Upper Mississippi Val¬

ley the first of the Richmond formations lies on the Galena dolomite

98 Wisconsin Academy of Sciences, Arts, and Letters,

which is of Trenton age. Accordingly the section here lacks the

important Eden and Maysville groups of the Cincinnatian series.

Farther south in this great valley, as in southeastern Missouri and

northern Arkansas, the hiatus beneath the Eichmond is increased

by elimination of the Galena so that the former lies on a limestone

of either early Trenton or late Black Eiver age. In the western

half of the continent it is still further increased, the first of the

Eichmond deposits there being in contact with beds ranging in age

from Middle or Lower Ordovician to Canadian, Ozarkian, or even

Upper Cambrian.

In the Appalachian Valley region the hiatus between the Ordo¬

vician and the Silurian is of lesser time value than in the Missis¬

sippi Valley. But here, locally, as in the vicinity of Lewistown,

Pa., the red Juniata sandstone, which is of Eichmond or Lower

Medina age and there rests on a thick mass of the latest Ordovician

gray Oswego sandstone, is not only much thicker than usual but

contains in its lower 400 feet many often thick beds filled with

rounded quartz pebbles. In size the pebbles vary from very small

to three inches in diameter. In central Pennsylvania, therefore,

we find all the physical evidence that even the most critical ob¬

server could demand on which we may confidently base the con¬

viction that the boundary between the base of the Eichmond (or of

formations corresponding in age to it) and the deposits beneath

its base is of systematic value; (1) great thickness of red muddy

sandstone indicating preceding, long enduring land-surface decay,

(2) thick beds of quartz conglomerate, and (3) extensive trans¬

gressions of new seas over previously long emerged areas of much

older rocks. It is important further to note that in Pennsylvania

where the first and second of these conditions are more clearly in¬

dicated than usual the boundary between the Juniata and the over-

lying Tuscarora sandstone (Upper Medina with which Schuchert,

Grabau, and other paleontologists propose to begin the Silurian) is

practically indeterminable. And yet the part of the section in

which it must lie is perfectly exposed in the section between Lewis-

town and Eeedsville. Aside from the Oswego- Juniata contact

above discussed the only other reasonably possible plane at which

the boundary between the two systems might be drawn is at the

locally clearly defined contact between the base of the Oswego

sandstone and the top of the Eeedsville shale (~Eden plus Lower

Ulrich — Paleozoic Systems in Wisconsin,

99

Maysville) where Butts/® following the long-continued practice of

the State geologists of New York, recently drew it. But it is im-

possible to locate this plane in New York. There, as in Pennsyl¬

vania, the boundary must be placed above the Oswego.^^

The Ordovician-Canadian break. — The Ordovician system as de¬

fined and restricted by Ulrich and given in the preceding table

begins in the Upper Mississippi Valley with the St. Peter sand¬

stone. That the surface upon which this sandstone was laid down

is decidedly uneven because of preceding weathering and erosion

has been long recognized in the geological reports of the State.

The evidence in hand 40 years ago was already sufficient to estab¬

lish much of the importance of the break between the St. Peter and

the underlying formation which then was supposed to be always

some part of the ^‘Magnesian limestone.” However, in more re¬

cent years much information bearing on this break has been ac¬

quired. We now know, especially from the evidence of deep wells,

reported by Thwaites, that the St. Peter may rest on any forma¬

tion between the top of the Shakopee and the eroded top of the

Eau Claire formation. We have learned further that the Sf. Peter

represents merely a late and perhaps the latest stage of an alter¬

nating series of limestones and sandstones that so far as known is

best developed in northern Arkansas. There, moreover, it is clearlj

shown^® that the deposition of this series was interrupted at least

once by emergence and deep erosional wear of the Everton lime¬

stone. At the bottom of the series in Arkansas there is often a

chert conglomerate which sometimes reaches a thickness of 8 or 10

feet. Under this lies the Powell limestone, which corresponds ap¬

proximately in age to the Shakopee dolomite of Wisconsin and ad¬

joining States to the west and south. Both of these formations are

assigned to the upper division of the Canadian system.

But we have learned something more, too, about the occurrence

of thick deposits of clastic material at' the base of the St. Peter in

southern Wisconsin and northern Illinois. This information has

been acquired in limited amount from study of outcrops by the

writer and his associates and in greater amount and details through

the study of well records by Thwaites. This clastic material or

conglomerate consists of great accumulations of chert boulders

Geologic section of Blair and Huntingdon counties, central Pennsylvania. Amer.

Jour. Sci., vol. 46, p. 536, 1918.

This systemic break is discussed at length by E. O. Ulrich, Ordovician- Silurian

Boundary, Int. Geol. Cong., XII, Canada, 1913, C. R., pp. 593-667, 1914.

Geol. Atlas U. S., Eureka Springs-Harrison Polio, 202, 1916, p. 7.

100 Wisconsin Academy of Sciences, Arts, and Letters.

and siliceous rock meal interbedded or irregularly mixed with

quartz sand and fine siliceous clay-like deposits, locally attaining an

aggregate thickness of nearly 200 feet and all derived from the sur¬

face wear of older formations to the north. The distribution of

this basal St. Peter conglomerate suggests long continued work of

streams of considerable size. In short, the physical data now in

hand clearly establish the propriety of Ulrich’s proposal to draw

a systematic boundary between the top of the Shakopee and the

base of the St. Peter in the Upper Mississippi Valley and at the

corresponding everywhere easily identified plane elsewhere in

America. Out of the hundreds of species of fossils found on the

two sides of this stratigraphic break not a single one passes across

the line ; and of the Canadian genera many less than half their num¬

ber are recognized in the succeeding Ordovician faunas. Besides,

this restriction of the Ordovician system still leaves it with an

aggregate thickness of deposits exceeding the average for most of

the systems now recognized.

The Canadian-0 zarhian break. — In the Upper Mississippi Valley

this break commonly lies within the mass of dolomitic limestone

that up to ten or fifteen years ago was generally known as the

Lower Magnesian limestone. More recently the locality term

Prairie du Chien limestone or formation was ill-advisedly proposed

to replace the lithologic name. Before that McGee^^ working in

northeastern Iowa proposed the name Oneota for the lower part of

the mass and WinchelP® gave the name Shakopee to what we now

know as its upper part. A sandstone believed to lie between the

two was called New Richmond sandstone by Wooster.-^ Because of

the formerly prevailing belief that these three divisions are coex¬

tensive much confusion has attended their identification from place

to place in Iowa and Minnesota. In Wisconsin, following publica¬

tion of Wooster’s work in 1882, practically no published attempt'

has been made to distinguish the divisions.

The field work of the writer in Wisconsin and adjoining States,

particularly during the past ten years, has demonstrated that' the

Shakopee is unquestionably distinguishable from the Oneota dolo¬

mite throughout their extent in the Upper Mississippi Valley. The

evidence respecting the New Richmond sandstone is much less sat¬

isfactory. It is evident not only that this dolomitic series embraces

U. S. Geol. Surv., 11th Ann. Rept., pt. i, p. 331, 1891.

>2® Minn. Geol. and Nat. Hist. Surv., 2d Ann. Rept., p. 138, 1874.

Geology of Wisconsin, vol. IV, pp. 106 and 127, 1882.

Ulrich — Paleozoic Systems in Wisconsin.

101

more than one sandstone but also that the two dolomite formations

are not by any means always separated by a sandstone. Moreover,

it seems that the sandstone beds are in the form of geographically

limited lenses lying at varying horizons within either of the dolo-

mitic formations and passing laterally into sand-free dolomites.

It has been established also that even the Shakopee and the Oneota

are not coextensive. In some places the Oneota is absent, as at

Jordan, Minn., and probably also at Eipon and Butte des Morts, in

eastern Wisconsin. At many other places in Wisconsin the Shak¬

opee is absent. In the former cases the absence of the Oneota is

most probably due mainly to nondeposition. In the latter instances

we can not be sure that the absence of the Shakopee is not caused en¬

tirely by the unusual activity of surface eroding agencies during

and preceding deposition of the St. Peter sandstone.

Where both of the dolomitic formations are present and clearly

exposed it has usually been found possible to point out precisely

the plane of contact. But the contact zone is a very likely one to

be affected and correspondingly obscured by secondary dolomitiza-

tion. In such unfavorable places the boundary was located only

approximately though with more time at our disposal than was

available better results might well be expected. In yet other places

the boundary proved to be quite irregular and was further empha¬

sized by mineralization where the hollows contained original car¬

bonaceous mud deposits and rather large pebbles of chert.

Good exposures of the contact were observed in the bluffs and

quarries at Stillwater, Minn. Here it lies about 60 feet above the

base of the Oneota which rests unconformably on the Jordan

sandstone. Above the top of the Oneota the Stillwater section

shows about 50 feet of Shakopee with usual characters. There is

no sandstone worth mentioning between the two formations.

That the surface of the Oneota was eroded here before the Shak¬

opee was laid on it is indicated clearly enough by (1) the rela¬

tively slight thickness of the lower formation, (2) the absence of

the fossiliferous cherty zone that is commonly present in the upper

part of Oneota sections at and to the south of Trempealeau, (3) the

unevenness of the contact plane which shows irregularities of con¬

tour of a foot or more and corresponding dissection across sedi¬

mentary planes in distances of less than 10 feet; and (4) the pres¬

ence of one to three inches of conglomerate with limestone and

chert pebbles in a matrix of coarse quartz sand and grains of glau¬

conite.

102 Wisconsin Academy of Sciences, Arts, and Letters,

In the section at Prairie du Chien the Oneota is at least 150 feet

thick, and the first of the fossiliferous cherts comes in about 65 feet

above the bottom. The formation therefore comprises at least 90

feet of more or less cherty beds at the top that are wanting at

Stillwater. At Dresbach, Minn., which lies about midway between

Prairie du Chien and Stillwater, the Oneota is approximately 100

feet thick. The fossiliferous cherty zone referred to in this and the

preceding paragraph occupies the upper 30 feet. The thinning of

the Oneota to the northward, apparently by erosion of the upper

beds, is suggested in similar manner by other sections in the valley

north of Prairie du Chien.

A sandstone, which may or may not correlate with the lower of

the two beds of sandstone at New Kichmond, Wis., is locally present

at the base of the Shakopee. A four to five-foot bed of sandstone

occupies this position at Prairie du Chien. Five miles to the north

of this city the same bed has thickened to about 10 feet. As re¬

marked in a preceding paragraph a bed of sandstone may occur at

the Shakopee-Oneota contact, but it is very unsafe to rely on its

presence.

It has been shown that good physical evidence of the break be¬

tween the Shakopee and the Oneota dolomites — in other words, be¬

tween the representatives respectively of the Canadian and

Ozarkian systems — is generally procurable in the numerous and

often excellent exposures of the “Lower Magnesian limestone’’ in

the Upper Mississippi Valley. But we must go elsewhere for the

data required to appreciate the true significance of the break. By

study of the sections in Wisconsin and Minnesota alone it would

be impossible to reach the absolutely established conclusion that

the Shakopee represents only the concluding stage of a great series

of limestones and dolomites having an aggregate thickness of over

5,000 feet. In northern Arkansas the Powell limestone carries the

same peculiar gastropod fauna that is found to the north in the

Shakopee dolomite. In Arkansas and Missouri the Powell is under¬

lain by three Canadian formations (Cotter, Jefferson City, and

Eoubidoux) each with its own characteristic fauna of which not a

trace has been observed north of Illinois. On the other hand, all

three of these faunal zones are clearly recognized in the Canadian

limestones in the Appalachian Valley, particularly in northeastern

Tennessee and western Virginia and also in central Pennsylvania.

In Virginia the Eoubidoux fauna occurs at the base of over 2,000

feet of dolomitic limestone; and the younger Canadian faunas

Ulrich — Paleozoic Systems in Wisconsin,

103

occur at appropriate horizons above it. In one of the best of the

sections in Pennsylvania (at Belief onte) the Eoubidoux fauna lies

over 3,000 feet beneath the base of the overlying Ordovician forma¬

tions. Moreover, this section shows more than 1,000 feet of yet

older Canadian limestones before reaching the top of the underlying

Ozarkian system.

Without facts like these one could scarcely believe that the break

between the Shakopee and Oneota in Wisconsin represents time so

long that it sufficed for the slow marine deposition elsewhere of not

less than 4,500 feet of limestone and dolomite. Nor is this all, for

even in the thickest of the Appalachian sections the physical evi¬

dence of the break between the Canadian and Ozarkian systems

appears quite as distinct and impressive as it is in Minnesota and

Wisconsin between the Shakopee and Oneota dolomites.

Striking and conclusive evidence of subaerial erosion of the

surface of the Ozarkian system during the interval between that

and the succeeding Canadian period was procured recently in south¬

eastern Missouri. Namely, in traveling northward through this

state from Van Buren to Potosi and thence westward toward Mera-

mec River the Gasconade dolomite, which in Missouri lies at the

top of the Ozarkian and is always easily recognized by its peculiar

chert and highly characteristic fossils, w^as found to be generally

rather thin and in places entirely absent. Indeed the areas in

which the Gasconade cherts and fossils were not observed must

aggregate at least 50 square miles. At first this local absence of

the Gasconade seemed most probably due to nondeposition but

further investigation, which showed an altogether unusual amount

of quartz sand and chert pebbles in the overlying Roubidoux forma¬

tion, soon tended to change the first suggestion to the belief and

finally to the conviction that the Gasconade after having been laid

down over the entire region was then locally removed by erosion

during the emergent interval between the two periods. The most

convincing evidence favoring the latter view was the discovery of a

ring or funnel of steeply inclined and highly fossilif erous Gasconade

chert in the midst of horizontal beds of an older Ozarkian forma¬

tion. As this older formation extended nearly to the tops of the

surrounding hills and was succeeded in the section by conglomeratic

Roubidoux sandstone and chert without intervention of the Gas¬

conade, no other explanation of the mentioned funnel-shaped out¬

crop of Gasconade chert than that it is a remnant of the original

104 Wisconsin Academy of Sciences, Arts, and Letters,

sheet of its formation which owes its preservation to collapse of the

roof of an Eopaleozoic cavern seemed to fit the case.

The Oneota is of Upper Ozarkian age. The sea of its epoch

lasted a long time in the southern Appalachian region but finally

spread very widely over southeastern North America. Its rocks

and characteristic fossils are recognized also in Missouri (Gas¬

conade formation), in Alabama and east Tennessee (Chepultepec

formation), in Pennsylvania, and in eastern New York. At the

close of its stage this great inland sea was withdrawn perhaps en¬

tirely. The next succeeding invasion laid down the limestone and

graptolitic shale deposits with which the accessible record of the

history of the Canadian period begins. Whereas the preceding

Oneota-Gasconade-Chepultepec stage spread northeastwardly from

the south and west, the succeeding Lower Canadian stage had its

development in North America mainly or entirely in Pennsylvania

and New York. Judging from the distribution of its deposits this

new sea must have differed from the preceding also in the location

of its inlet and the direction of its invasion. It came in from the

Atlantic side of the continent, whereas the preceding late Ozarkian

sea invaded from the south. Evidently then the physical changes

that occurred in passing from the Ozarkian to the Canadian period

involved very considerable local warping and also more generally

effective differential vertical movements of the lithosphere. The

geographic changes at this time — meaning particularly changes in

arrangement and outlining of water and land areas — also the

faunal modifications, were greater and more important than at any

other time between the close of the Cambrian and the beginning of

the Ordovician period.

The Ozarkian-Cambrian break. — To get a true conception of the

diastrophic changes that' occurred in America during the passage

from the Cambrian to the Ozarkian period we must study the

stratigraphic records preserved in the great Appalachian and

Cordilleran geosynclines. However, as we found in discussing the

younger Paleozoic breaks, the record in Wisconsin and adjoining

States regarding the Ozarkian-Cambrian break also is far from

blank. In fact, the Devils Lake sandstone, which is the oldest of

the Ozarkian formations in Wisconsin, affords a more impressive

development and display of conglomerate than has . been observed

at this horizon anywhere else in America. This sandstone attains

a thickness of at least 100 feet and at' many places includes — as in

Parfrey Glen, east of Baraboo, and at the northern end of the

Ulrich — Paleozoic Systems in Wisconsin.

105

gorge at Ableman- — at its base and at various higher positions in

the formation beds of almost perfectly rounded quartzite pebbles.

These range in size from less than an inch in diameter to masses

three or four feet across. The typical outcrop of the Devils Lake

sandstone occurring^ as they do, on the flanks of old quartzite

ridges, were deposited under conditions admirabily adapted to the

production of such coarse clastic material.

Conglomerate of similar kind and probably derived from the

same or similar sources occurs in southern Wisconsin somewhat

rarely at the bases of preceding Cambrian formations. But the

volume of such material in these older occurrences is far inferior

to that found in the Devils Lake sandstone. This disparity in

quantity suggests not only intervening diastrophic movements that

made more of such material available but also the lapse of consid¬

erable time during which such large quantities might be produced.

Eeally, in view of the demonstrable fact that the Devils Lake sand¬

stone is of quite early Ozarkian age and the deposits beneath it

very late Cambrian, the abundance and character of the conglom¬

erates in the former prove to be the best objective evidence we have

in establishing the verity of the break between the Cambrian and

the Ozarkian.

That warping of the surface did occur in the Upper Mississippi

Valley during the otherwise unrecorded interval between the Cam¬

brian and Ozarkian is further clearly proven by the altogether dif¬

ferent distribution of the Cambrian and early Ozarkian marine

sediments. Thus, whereas the Cambrian formations are for the

most part common to both the east and west sides of Wisconsin the

first two of the Lower Ozarkian formations are confined to small

areas in the eastern half of the State. The succeeding Madison

sandstone seems to extend from the most southern of these areas

across the State to the Mississippi, but even this appears to be con¬

fined to much narrower limits in north and south directions than

were the Cambrian formations.

Another interesting and in this connection important fact is that

whereas the organic remains in the Cambrian formations, except¬

ing perhaps those of the Mazomanie, become fewer and finally dis¬

appear entirely when the formations are traced toward the east

and especially the northeast angle of the State, the Lower Ozarkian

formations, on the contrary, become barren in a westerly direction.

This fact suggests at once that the seas in which the more impor¬

tant of the Cambrian formations were deposited terminated on the

106 Wisconsin Academy of Sciences, Arts, and Letters,

northeast side against the beaches and low coastal plains of land

areas then prevailing over and beyond the area of Michigan. The

Lower Ozarkian seas, on the other hand, shallowed to beaches on

the west side where seas with abundant life had prevailed in the

preceding period. These suggestions are raised to the status of

practically demonstrated facts when the distribution and origin of

their respective faunas is carefully taken into account. Unpub¬

lished studies of Cambrian faunas have reached the point where

we can say definitely that direct marine connections existed re¬

peatedly and perhaps continuously between the Cambrian seas of

the Upper Mississippi Valley and those of Wyoming and Montana

and Oklahoma and central Texas. Indeed, there is no reason to

doubt that the most characteristic genera and species of the Cam¬

brian faunas found in the Upper Mississippi region migrated here

from the west and southwest. But the case with respect to the

Lower Ozarkian faunas of Wisconsin is totally different. These

are like and must have been in communication with the Potsdam

and Hoyt faunas of New York and, by extension of colonies south¬

wardly, with Missouri. Moreover, the Lower Ozarkian faunas in

the Mississippi Valley and New York are almost totally different

from those of similar age in the Cordilleran province. The latter

evidently invaded from the Arctic side of the continent.

The facts here presented in brief outline in support of the re¬

puted high taxonomic importance of the break between the Cam¬

brian and Ozarkian systems might by themselves be accepted as

sufficient proof of the author’s contention regarding the actual dis¬

tinctness of the two systems. But it is unnecessary to be satisfied

with the data supplied by Wisconsin localities alone. Other and

many of them mucn more convincing data have been procured from

other parts of North America. In fact, the Ozarkian is very in¬

completely represented in Wisconsin. A fuller sequence of its de¬

posits is found in the Ozark Uplift of Missouri and Arkansas. But

very much thicker deposits of this period have been observed and

carefully studied in the Appalachian Valley region. There, par¬

ticularly in central Pennsylvania and central Alabama, Upper

Cambrian deposits consisting largely of limestone and aggregating

thousands of feet in thickness are succeeded unconformably by

Lower Ozarkian dolomites and relatively pure limestones that

measure as much as 2,000 feet or more in a single completely ex¬

posed section. On these, then, rest thousands of feet of Middle and

Upper Ozarkian limestones and dolomites giving a total volume of

Ulrich — Paleozoic Systems in Wisconsin.

107

limy deposits that must, on the basis of thickness alone, rank the

Ozarkian among the most important of the Paleozoic systems.

In North America the Ozarkian system has as good or a better

foundation in its diastrophic history, volume and character of de¬

posits, and in its distinctive faunas than have the Silurian, De¬

vonian, Permian, Triassic, and Jurassic systems, most of which are

generally accepted by stratigraphers.

In the matter of areal distribution also, especially when we con¬

sider the fact that as one of the oldest of the Paleozoic systems its

rocks are likely to be largely buried beneath younger sediments,

the Ozarkian compares favorably with other Paleozoic systems.

Kocks of its age are now definitely recognized as outcropping in

New York, Vermont, New Jersey, Pennsylvania, Maryland, Vir¬

ginia, Tennessee, Georgia, Alabama, Missouri, Iowa, Wisconsin,

Minnesota, Oklahoma, central and western Texas, Colorado, Idaho,

Nevada, and, through the explorations of Walcott, in British Co¬

lumbia. By means of deep wells this already very widely demon¬

strated areal distribution doubtelss will be extended to other

States. Moreover, there are good reasons for believing that the

system is represented also in Quebec and Newfoundland and on the

nther side of the Atlantic by the Tremadoc in Great Britain and

perhaps in other European countries. For the present, however,

certain very real difficulties, which always are encountered in the

endeavor to correlate fossil shore and bottom faunas of widely sepa¬

rated provinces, forbid more definite statements regarding the possi¬

ble and probable representatives of the Ozarkian on other than the

North American continent.

THE FAUNA OF THE LAKE WINNEBAGO REGION*

A Quantitative and Qualitative Survey with Special

Reference to the Mollusca

Frank Collins Baker

During the summer of 1920 (July and August) a somewhat de¬

tailed survey was made of the Lake Winnebago region, including

Lake Winnebago, Lake Butte des Morts, the Fox River at Omro,

and the small swales and pools bordering the shores of these places.

The work was carried on under the auspices of the Wisconsin

Geological and Natural History Survey, by which the writer was

commissioned to make a survey of the molluscan fauna of the State

preparatory to the completion of a monograph of the Mollusca.

The absence of any literature, excepting scattered records, con¬

cerning the mollusk fauna of the Winnebago region led to its se¬

lection as a profitable area for statistical study for comparison

with other lakes which have been similarly treated.

Headquarters were established at the Wisconsin State Fish

Hatchery at Oshkosh, where motor and row boats were placed at

the writer’s disposal, as well as suitable laboratory facilities for

sorting and otherwise caring for the collections. Collections were

made at definite stations along the west shore of Winnebago Lake

from Asylum Bay to Long Point Island, a distance of about 12

miles. Shore material was also gathered at other places on the lake

shore. In Lake Butte des Morts, collections were made at places

covering the greater part of this body of water. Over 450 dredg¬

ings were made and, in addition, mollusks were collected from many

places along the shore and in inland habitats. The dredgings were

made with an Ekman bottom sampler. The results shown in the

tables indicate the number of individuals per square meter of

bottom.

The material upon which the study is based is preserved in the

Zoological Museum of the University of Wisconsin (Mollusca) and

Contribution from the Museum of Natural History, University of Illinois, No. 32.

110 Wisconsin Academy of Sciences, Arts, and Letters.

in the Museum of Natural History of the University of Illinois

(Mollusca and associated animals).

My thanks are due a large number of people for assistance in

the field work or in the subsequent study of the material. Chief

among these are Professor Chancey Juday of the Wisconsin

Geological and Natural History Survey, who made the study pos¬

sible; and the Conservation Commissioners of the State, through

Mr. I. H. Bloomer, in charge of the Pish Hatchery, who gave every

possible facility for advancing the work. Mr. Hall and Mr. Apel,

hatchery assistants, also granted many courtesies which greatly

assisted in the success of the undertaking. To Mr. Clyde B. Terrell,

of Oshkosh, the owner of the interesting aquatic farms on Lake

Butte des Morts, the writer is greatly indebted for assistance in

studying the fauna of that lake.

The sincere gratitude of the writer is especially due those spe¬

cialists who cheerfully gave of their time for the identification of

groups of animals in which they are experts. The large amount of

material sent could not but have been somewhat of a burden to

classify, and I am sure that the feelings of other workers in

ecological subjects are expressed in the statement that the value

of their assistance cannot be overestimated. The following persons

assisted in this work of identification: Dr. Bryant Walker, De¬

troit, Mich., various mollusks; Dr. V. Sterki, New Philadelphia,

Ohio, Sphaeriidae; Mr. Calvin Goodrich, Detroit, Pleurocera; Dr.

Kuth Marshall, Rockford College, Rockford, Ill., Acarina; Dr. J.

Percy Moore, University of Pennsylvania, leeches; Dr. J. G. Need¬

ham, Cornell University, acquatic insects; Dr. Cornelius Betten,

Cornell University, Trichoptera; Dr. Paul S. Welch, University of

Michigan, aquatic Lepidoptera ; Professor Frank Smith, University

of Illinois, Oligochaete worms and sponges; Dr. C. P. Alexander,

LTniversity of Illinois, aquatic insects; Dr. Wm. Trelease, Univer¬

sity of Illinois, certain plants; Miss Ada L. Weckel, Oak Park, Ill.,

Amphipoda; Mr. Waldo L. Schmitt, U. S. Nat. Mus., Washington,

D. C., Isopoda; Prof. Chancey Juday, University of Wisconsin,

Cladocera.

Baker — The Fauna of The Lake Winnebago Region. Ill

PHYSIOGRAPHY AND GEOLOGY

A. Description of the Lake

Lake Winnebago lies in the eastern part of the State of Wis¬

consin in latitude 44° N. and between longitude 88° and 89° W.

It is 33 miles west of Lake Michigan and 30 miles southwest of

Green Bay. The lake is oriented north and south, which is its

greatest length, and is about 28 miles (45 km.) in length by 10.4

miles (16.6 km.) in greatest width. The greatest depth is 20%

feet (6.38 m). It lies at an elevation of 745 feet above the sea and

165 feet above Lake Michigan. The lake has a present area of about

215 square miles, which is about 5.5 per cent greater than its

original area, the increase being due to the dams in the outlet

channels which have raised the level of the lake several feet.

The eastern and western shores of Lake Winnebago differ greatly

in character. The eastern short presents an almost continuous

margin broken by no bays or points of notable size. The western

shore, on the contrary, is made up of a succession of bays and

points with many shoals extending well into the lake. On the

eastern side, the land rises rather abruptly forming cliffs or bluffs

of a more or less bold character. In most places the rise is gradual

(50 feet) for a quarter of a mile and then abrupt (100-150 feet)

in a distance of a quarter or half a mile. Elevations of 1,000 feet,

or 255 feet above the lake, occur at a distance of less than two miles

from the lake margin.

The western margin of the lake is very low and lacks entirely the

bold character of the eastern shore. Elevations of more than six¬

teen feet above the lake are not encountered within a distance

of a mile west of the lake. Marsh areas occur at the south end

near the City of Fond du Lac. West of the City of Oshkosh, low,

marshy areas extend westward for a distance of nearly twenty

miles, where smaller lakes, Butte des Morts, Poygon, and Winne-

conne, occupy wide areas.

1

Fig. 1. Map of Lake Winnebago showing 3, 5, and 6 meter contour lines.

The circles indicate the stations where collections were made.

Baker — The Fauna of The Lake Winnebago Region. 113

Lake Winnebago is fed by a number of large streams. Fox River

enters the northwest end of Lake Butte des Morts and forms a

channel which extends through this lake, emptying into Lake Win¬

nebago at Oshkosh. In fact, these lakes are really widened-out

portions of the Fox River. Pine Creek and Willow Creek enter

the western end of Lake Poygon and Wolf River enters the same

lake from the north. Fond du Lac River, a small stream, enters

Lake Winnebago at the southern end. The water shed of Lake

Winnebago covers a wide area in east-central Wisconsin, approxi¬

mating 6,200 square miles (see Whitbeck, 1915, pi. iv). The out¬

let of Lake Winnebago is by way of the Lower Fox River into

Green Bay. Just below the dams the river widens to form Little

Lake Butte des Morts.

The basin of Lake Winnebago is somewhat platter-shaped, the

bottom descending more or less abruptly to a depth of three meters

and then more gradually to five and six meters. The greater part

of the lake bottom forms a subaqueous plain which varies but

slightly in contour (see Juday, 1914, pi. 26, and Lake Survey

map). Subaqueous terraces are rare on the eastern shore but

common on the western shore, where there are many shallow bays,

at the points of which bars and shoals extend into deeper water. In

some places the slopes of these terraces are very steep. The bot¬

tom of the lake is of glacial drift of great thickness.

B. Origin of Lake Winnebago

Lake Winnebago is the result of changes which occurred during

the last glacial period — the Late Wisconsin. When the Green

Bay lobe of the glacier receded it left a morainic dam in the Fox

River Valley at the present site of the City of Menasha. The lake

was at first much larger than at present and extended westward

and southward in the valley now occupied by lakes Butte des

Morts, Poygon, Winneconne, and a part of the Fox River Valley.

Fond du Lac was also submerged. Due to readvances of the ice,

the lake fluctuated in size before reaching its present level. The

red till, so conspicuous in the wave-cut cliffs near South Asylum

Bay, and elsewhere, was laid down during one of these advances

of the Green Bay lobe.

The glacial outlet was at the present site of Menasha, at the

northwest corner of the lake, where the bed rock comes to the sur¬

face and forms a rock-sill which held the water at different heights

114 Wisconsin Academy of Sciences, Arts, and Letters.

as it was gradually cut down. The present level of the lake is

controlled by the government dam at Menasha. The preglacial

channel of the Fox Eiver lies to the east of the present outlet,

toward Clifton, but it was filled with glacial material, thus block¬

ing the old outlet ; this compelled the lake to find the lowest avail¬

able col or notch which chanced to be at the present sites of Neenah

and Menasha (see Whitbeck, 1915; Martin, 1916; Goldthwait, 1907 ;

Alden, 1918).

C. Lake Physiology

Many agents, which may be called physiological, affect the habi¬

tats in Lake Winnebago. Among these, winds and waves are

potent factors in shaping the physiography of the shores and con¬

sequently the character of the animal inhabitants. The great sur¬

face area of Lake Winnebago provides a fertile field for the action

of winds, which disturb the surface of the lake and often descend

to considerable depths. A severe gale from the northeast causes

waves of great length and height and provides an undertow which

disturbs the bottom to a depth of two meters or more. These heavy

waves have made the peculiar rocky shores seen ,on Doemel Point,

Asylum Point, Stony Point, and other places. Sand, gravel, and

cobbles are constantly being carried from one point to another,

profoundly affecting these habitats. In many places the waves and

undertow have formed off-shore bars or shoals composed mostly of

sand, the water being a third of a meter in depth on the bars and

a meter or more between the bars. Gravel and cobble bars have

also been formed off every point, in several places in water as deep

as three meters. Plant zones are profoundly affected by wave

action, being absent or scanty where this action is heaviest and

most abundant where it is lightest.

Temperature. The temperature of a shallow lake like Winne¬

bago is high in summer and low in winter; ice is formed in Lake

Winnebago much earlier than in deeper lakes, which do not lose

their heat as quickly. On account of its shallowness, there is no

thermal stratification of the water in summer.

Transparency. The degree of transparency of the water of

Lake Winnebago varies with the season and with the climatic con¬

ditions. During July and August, 1920, the turbidity of the

water was great and transparency was reduced to a minimum. A

Baker — The Fauna of The Lake Winnebago Region. 115

white disc could be seen only to a depth of a few inches in the open

lake. This was due largely to the presence of vast quantities of

phytoplankton which was universally distributed and unusually

abundant during these months. The wind moved this mass from

shore to shore. Occasionally some of the bays were free of the

plankton and the water became fairly clear so that the bottom could

be plainly seen at a depth of a meter. Almost everywhere the

water was filled with fine sediment held in suspension. Compared

with some other lakes the degree of transparency in Lake Winne¬

bago is very low. Bkman (1915) gives 18 jneters for Lake Vaet-

tern in Sweden ; Muttkowski gives three meters for Lake Mendota ;

and Baker found between four and five meters in Oneida Lake,

N. Y.

D. Pollution

Lake Winnebago is suffering from some degree of sewage pollu¬

tion. Sewage from the City of Oshkosh enters the Fox River and

is discharged into the lake south of the city. Dredgings made about

half a mile to a mile and a half south and east of Oshkosh showed

a bottom of black mud at a depth of 4.6 meters, in which there was

a large quantity of oil that came to the surface of the water in the

pail containing the dredgings, forming a thick film. No animal

life of any kind was found in these dredgings, but near by, outside

of the channel of the Fox River, bottom life was fairly abundant

at similar depths. The same conditions that compelled Chicago

to divert its sewage from the source of water supply is now con¬

fronting Oshkosh and epidemics of typhoid or other water-borne

diseases are liable to occur as the sewage increases in volume and

becomes mixed with the water supply, which is drawn from the

lake east of the city, about half a mile from shore, in water less

than four meters in depth. It is probable that some manufactur¬

ing wastes also enter the lake. The sewage of Fond du Lac also

contaminates the waters of this lake.

THE MACROFAUNA OF THE LAKE WINNEBAGO REGION

The region in which lie lakes Winnebago, Butte des Morts, and

the Fox River provides unusual opportunities for the study of

ecological variation. The Fox and Wolf rivers, flowing for many

miles as typical rivers, expand to form large lakes west of Oshkosh.

Lake Butte des Morts outflows through a wide river at Oshkosh,

116 Wisconsin Academy of Sciences, Arts, and Letters.

which, after flowing for a distance of about two miles, enters Lake

Winnebago. The waters of this lake find outfiow by way of the

lower Fox Kiver which empties into Green Bay of Lake Michigan.

The change in the physical environment from the comparatively

quiet waters of a river to the rough and turbulent waters of a

lake, has been strikingly refiected in at least one group of animals,

the mollusks, the lake species being mostly different from the river

species. As would be expected. Lake Butte des Morts, which is

but the widening out of the lower part of the Fox River, shows

on the whole less change from the river type than does Lake Win¬

nebago, which is larger, with all the features of a true lake.

As has been shown elsewhere (Baker, 1916, 1918) the larger

part of the macrofauna of a lake is found within the two meter

contour (about six feet) and consists of animals in intimate con¬

tact with the substratum of the lake — the bottom and the plants

growing on the bottom — where are found the optimum conditions

for successful continuance of life — food and oxygen. This is the

littoral (or eulittoral) area comparable to the great areas of the

sea shore which teem with life.

The shallowness of Lake Winnebago, with a maximum depth of

6.38 meters, makes an ecological division of the lake into depth

zones impossible ; there is no deep area, or even a true aphytal area,

as in the deeper lakes, such as Lake Mendota and Green Lake. An

area possibly comparable to the aphytal region of the deeper lakes,

where large aquatic plants are absent, may be represented in Lake

Winnebago by the central area of maximum depth, although this

may equally well be placed in the sublittoral region. The littoral

(eulittoral) area extends from the shoreline to a depth of three

meters, forming a narrow shelf bordering the shores of the lake;

life is most abundant in this region. The sublittoral (or aphytal)

area includes the remainder of the lake bottom. In this region the

bottom consists of soft, black mud with a considerable amount of

organic material.

Nearly all of the bottom of the lake is covered with a detritus

which is made up of small fragments of plants, minute pieces of

wood, crustacean skeletons, caddis-worm cases, and fragments of

molluscan shells. This material is absent only where the waves

are strong enough to wash it away. The plant fragments probably

constitute an important source of food for the bottom-feeding

forms.

Baker — The Fauna of The Lake Winnebago Region, 117

Quantitative studies of the composition of the bottom fauna of

the sea have been carried on for a number of years (see Petersen,

1911, 1915, etc.) but similar studies on the biota of inland lakes

are of comparatively recent date (Baker, 1916, 1918 ; Ekman, 1915 ;

Muttkowski, 1918). Petersen (1911, p. 71) has said that we must

know the main points concerning the productibility of a body of

water before that body can be exploited for fishery purposes. This

statement applies even more forcefully to the inland waters be¬

cause here the problems are in a measure simpler and fish culture

is more easily controlled. Studies of this kind have been carried

on by both counts of the number of animals found in a measured

unit area and by weighing the dry animal matter of a unit area.

Both methods are useful and should be used wherever practicable.

In the Lake Winnebago work, which was conducted primarily to

ascertain the number and kind of mollusks in Lake Winnebago and

surrounding waters, only the counts of unit areas were made.

Table 1 shows the numerical results for the Mollusca, together

with the associated animals, in the different physiographic regions.

The depth of the water is given in meters and the character of the

bottom is indicated. The figures indicate the number of individ¬

uals per square meter of bottom. In the mollusks, the identifica¬

tion has been carried to all varieties ; in the associated animals, only

the larger groups are recorded.

Lake Winnebago

The Littoral Region, This is the most profoundly affected of

any part of the lake area, including in its territory the shore with

its breaker line where the physical forces are constantly at work.

The littoral area may be divided into three quite characteristic

subdivisions; 1, the general shore line; 2, the breaker line; and

3, the plant areas. The first two divisions often overlap, and in

the present paper will be considered under the several headings

of boulders, sand, etc. The shore line is subject to great physio¬

graphic changes owing to the molar activity which causes the bot¬

tom to constantly shift more or less. Ecological conditions are

here severe, the shifting bottom, the exposure to wide changes in

temperature, and the pounding of the waves compelling the biota

to make frequent readjustments to the environment. It is here

that the greatest diversity of molluscan life occurs and the species

118 Wisconsin Academy of Sciences, Arts, and Letters.

differ more or less markedly from the same type living in the qniet

parts of the Fox River.

Life on Boulder Shores. All of the points facing the lake and

all unprotected shores are covered with boulders, which form a

typical breaker line; gravel, followed by sand, being distributed

above the boulders (shoreward) and the same material occurring

downward (lakeward). Typical rocky shores composed of large

boulders seldom descend below two meters in depth. On long, ex¬

posed shores, as at Doemel Point, Par Rockaway Point, Asylum

Point, and near Stony Beach, the boulders are frequently of large

size, indicating that wave action has removed all of the finer par¬

ticles, leaving the heavier material as a breaker line. The typical

boulder shore (rachion) is usually not over three meters wide on

steep shores and descends to a depth of about a meter. Beyond

this depth the bottom changes to coarse gravel, then to sand, and

finally to mud.

The plant life of this region is scanty, as would be expected,

consisting chiefiy of filamentous algae (principally Cladophora)

the plume-like fronds of which hang from the upper surface or

sides of the rocks. A few emergent and submergent plants brave

this inhospitable habitat, Scirpus occidentalis being found in one

place, two species of Potamogeton in two places, and Vallisneria in

another place. The last two plants occur in some abundance in

several habitats, but Scirpus is rare. The gravel in these habitats

made growth for these plants possible.

Animal life is not abundant in species although certain species

may be abundant in individuals. Among the Mollusca eleven

species of Unionidae occur, all of them modified in form to meet

the rigorous conditions of this unstable environment. The shells

are smaller and thicker, on the average, than those of their river

representatives. They are not numerous in individuals, occurring

scattered among the boulders, between which they burrow. The

gastropods include such species as cling to rocks and feed upon

algae. Pleurocera, Physa, and Lymnaea are typical rock in¬

habitants and occur in some abundance. Planorhis parvus and

Somatogyrus are stragglers from other shores where they live

among the algae. All of the gastropods found on rocks (Pleuro¬

cera, Physa, Lymnaea) have a large, wide, ventral surface (the

foot) which enables them to cling to the support and so prevents

them from being dislodged by the waves. Species in this kind of

Baker — The Fauna of The Lake Winnebago Region. 119

a habitat often have the foot unusually developed for this pur¬

pose. Three species make up 73 per cent of the total population

of the boulder habitat : Pleurocera acuta 19 per cent ; Physa sayii

43 per cent; and Lymnaea winnebagoensis 11 per cent. Physa is

here the dominant species.

Associated animals are about six times as numerous in individ¬

uals as the Mollusca (Mollusca 17 per cent, associated animals 83

per cent). Fifteen higher groups of aquatic animals are repre¬

sented. The conspicuous forms on the rocks are the caddie-flies

(Trichoptera) the elongated or flattened tubes of which are fas¬

tened to the rocks by filmy anchors. The snail-like case of Heli-

copsyche and the flattened tubes of Leptocella are the most con¬

spicuous. Hydroptilidae and Agraylea are also represented. Poly-

centropid larvae, with their funnel-shaped tubes attached to the

rocks by broad bands of a fine cementing substance, are common.

Chironomid larvae and pupae are abundant, their tubes being at¬

tached to the under side of the rocks. The Bphemerids, repre¬

sented by the nymphs of Heptagenia, are occasionally seen on the

under side of rocks, but these are not as numerous as was observed

in Oneida Lake. Caenis occurred at one station. An Enallagma

nymph was caught at another station. Coleoptera, represented

principally by the flat larva of Psephenus lecontei, also included

Stenelmus bicarinatus, Oyrinus ventralis, and an unknown larva

of Dascyllidae.

In some places the smaller cobbles mixed with the larger rocks

are covered with life, especially associated animals. Thus at sta¬

tion 12, 1,225 individuals, representing twelve higher groups, were

picked from 24 rocks with a maximum surface area of 1,350 square

centimeters.

Cladophora harbors certain species which cannot be strictly

called petrophilus. Small Nematodes, Planaria, Cladocera, Ostra-

coda, the mites Limnesiopsis, Hygrobates, and Lebertia ; two

Hemiptera, Belostoma and a Corixid, occurred at two stations, in¬

cluding both adults and nymphs. The Amphipod, Hyalella knick-

erbockeri, was abundant where the alga was most plentiful. A few

Chironomid larvae as well as Trichopterid larvae also live among

the algae.

Burrowers, including small leeches and Cambarus propinquus,

were observed beneath the boulders, but they were not abundant

in the purely boulder habitats. The disparity in numbers between

the Mollusca and the associated animals in the boulder habitats is

120 Wisconsin Academy of Sciences, Arts, and Letters.

probably due to the better clinging powers of the latter, many of

which are especially modified to meet these conditions, as for ex¬

ample the flattened bodies of Psephenus and Heptagenia. Three

groups of associated animals make up 85 per cent of the entire pop¬

ulation of this kind of habitat, Amphipoda with 23 per cent, Tri-

choptera with 38 per cent, and Diptera with 24 per cent.

Ecological conditions in the boulder habitats are usually rather

rigorous. Feeding can only be done during periods of comparative

calm. The snails are all phytophagous, eating the algae. The

clams may secure their sustenance from the microorganisms that

are brought to them by the waves and currents. The difference in

shape and size between the clams of the river and lake may be due

in part to the greater difficulty of obtaining a sufficient quantity of

food in this rougher environment. The .same laws govern many

of the associated animals as regards their food supply.

Life on Gravel Shores (Table 1). Gravel habitats occupy a

large part of the shore line around Lake Winnebago. Gravel is

often found in association with boulders but is more usually found

occupying the place of the boulders on some exposed shore. It

varies in character from cobble stones to fine gravel, with which

there is usually more or less sand, often in spots. The gravel or

cobble is generally smooth and polished by the continuous wave

action. This kind of habitat is such that the inhabitants must be

good dingers or burrowers to escape being destroyed by the shift¬

ing character of the environment. Gravel beds may occur from

shore to a depth of nearly two meters. In one place it was found

at a depth of 3.4 meters, and subaqueous bars of gravel extend

lakeward from the points for a long distance. Vegetation consists

of the same species noted in the boulder habitats, with the addition

of Potamogeton lucens and Castalia odorata.

Among the Mollusca the same kind of naiades occur but the num¬

ber of species is reduced to six. Sphaerium appears sparingly and

Pisidium increases in both species and individuals. A number of

new gastropods appear and several of the boulder species (Pleu-

rocera, Physa) are less numerous, while Lymnaea has disappeared.

Amnicola and Yalvata occur in some abundance owing probably to

the presence of filamentous algae which is attached to the larger

stones. Three species of gastropods are common and abundant on

gravel bottom ; Amnicola limosa porata, Amnicola emarginata, and

Valvata tricarinata and its varieties. Habitat number 40 was the

Baker — The Fauna of The Lake Winnebago Region. 121

most prolific; the bottom was mixed with sand and the depth of

water and the distance from shore provided a habitat unusually

favorable to the development of an abundant molluscan fauna.

The associated animals were equally numerous. This station might

equally well, perhaps, be included among the sand shore stations.

Valvata makes up 46 per cent of the molluscan life of the gravel

habitats. The number of species of Mollusca has increased from

22 on the boulder shore to 35 on the gravel shore, an increase of

63 per cent.

Among the associated animals some of the rock-loving species

have disappeared or become less numerous. Heptagenia and

Psephenus are absent ; Leptocella and Polycentropidae are reduced

in numbers. Agraylea, Oecetis, and Molanna are scarce. Hemip-

tera is represented by a few Plea striola, adult, and Belostoma

nymphs. Among Coleoptera the larvae of an unknown Dascyllid,

common on the boulder bottom, is rare, but Gyrinus ventralis,

adult, is common. Of Hirudinea, Glossiphonia complanata, neph-

eloidea and stagnalis, Expohdella punctata and Nephelopsis ob-

scura are common. The filamentous algae harbor a number of

species, which occur sparingly in individuals, except in the Amphi-

pods. Planaria, a few oligochaete worms, and a number of Chiro-

nomid larvae occupy this algal habitat.

The groups of associated animals most numerous on the gravel

shores of Winnebago Lake are Amphipods, forming 31 per cent,

and Chironomid larvae, forming 41 per cent. Compared with the

boulder shores the gravel shores have about the same average popu¬

lation but there is less disparity between the mollusks and asso¬

ciated animals in the gravel habitats (Mollusca about 40 per cent).

Animal Life on Sand Shores (Table 1). Sand occurs on many

parts of the shore of the lake, either in connection with gravel or

boulders, or as large sand flats and shallows in more or less pro¬

tected bays. Such areas are found in Miller Bay, Asylum Bay,

and particularly along the shore south of the mouth of Fox Eiver

near Oshkosh, where sand borders the rocky shore for a distance of

several miles. Fahney Bay and the bays north of Moreley and

Black Wolf points, have sand beaches and shallows that are espe¬

cially well developed. South of Long Point Island sand bottom

occurs for upwards of a mile. The shore from Fox Eiver to Stony

Beach is perhaps characteristic of many parts of Winnebago Lake,

This shore is bordered by boulders with some gravel. Sand begins

122 Wisconsin Academy of Sciences, Arts, and Letters.

within a meter of the shore. At a distance of 50 or 60 meters from

shore there is a succession of sand bars representing present and

former breaker lines, which run parallel with the shore.

The transition from gravel to sand shore is in some places well

marked and in others is almost imperceptible. Usually, pebbles

and large stones gradually become less in number until they finally

disappear. In the coves and bays, especially in the area between

Fox River and Long Point Island, there is little fine gravel and no

coarse gravel, all is fine sand of even texture. Sand bottom areas

are usually shallow near shore (out to 150 meters) the depth sel¬

dom exceeding one meter. In the protected bays, and even on the

more exposed shores, vegetation is abundant and varied, including

both submerged and emergent species. In two habitats the sand

was mixed with marly clay and Chara was abundant. Filamentous

algae (Cladophora, etc.) are usually abundant. A sand bottom

unbound by vegetation is generally a poor faunal habitat, the

shifting character of the substratum burying the biota and smoth¬

ering it. Some Naiades, however, prefer this kind of a habitat and

are found abundantly in such places.

Mollusca are abundant in many of the habitats. Curiously

enough, the Naiades, which are fairly common on gravel and

boulder bottoms, occur sparingly on the sand bottom area,

Sphaerium is rare and Pisidium not as abundant as on the gravel

bottom. Of gastropods, Somatogyrus forms 22 per cent, Amnicola

limosa porata 18 per cent, and Planorhis parvus and Valvata tri-

carinata each 8 per cent. Pisidium constitutes 12 per cent. As a

whole, the molluscan fauna of the sand bottom is richer in species

than the gravel bottom, increasing from 35 to 40. The presence of

filamentous algae on the higher plants accounts in large measure

for the abundance of some of the mollusks, as the Amnicolidae and

Valvata.

The associated animals outnumber the mollusks about two to

one. The most abundant being Hyalella (about 18 per cent) and

Chironomid larvae (25 per cent). Oligochaete worms (mostly

Stylaria) make up about 14 per cent. Trichoptera comprise six

groups, Agraylea, Molanna, Oecetis, Helicopsyche, Polycentropidae,

and a few Leptocerids. Ephemeridae includes nymphs of Caenis,

Ephemera, and Ephemerella. Acarina is represented by five

genera, Limnesia, Limnesiopsis, Piona, Unionicola, and Hydrachna,

Piona being the most numerous. Hemiptera includes nymphs of

Belostoma, adults of Plea striola, and a Corixid nymph which was

Baker — The Fauna of The Lake Winnebago Region. 123

very abundant at station 28 in filamentous algae. Tubes of

Chironomid larvae are very numerous in the sand habitats. Leeches

are rare, Glossiphonia fusca, stagnalis, and nepheloidea and Dina

fervida occurring as single individuals.

Animal Life on Mud Bottom (Table 1).

Mud bottom areas in shallow water near shore are not common

in Lake Winnebago, occurring in protected bays and behind bars

or points. This kind of bottom is found in Miller Bay, behind

Asylum Point, in marsh portions of Fahney Bay, and in other small

protected bays south of the mouth of the Fox River, as well as in

several small beach pools behind the shore line. In parts of Miller

Bay a marl or marly-clay bottom occurs on which no life was found.

About the same species of plants are found here as on the sand

bottom.

Among mollusks the Naiades are entirely absent from mud bot¬

tom in shallow water and but one specimen occurred on this kind

of bottom in deeper water (5.2 meters). Pisidium is common in

places. Of the gastropods, which form 78 per cent of the mullusks,

Valvata includes 17 per cent, Physa sayii 11 per cent, and Planor-

Ms parvus 17 per cent. The mollusks are fewer in number of

species on a mud bottom than on any other kind of substratum ex¬

cept boulder. Habitat number 32, a stagnant pool behind the

beach south of Roe Point, contained the largest number of indi¬

viduals. Habitat number 42, a small bay on the south side of Long

Point Island, contained the next largest number per unit area, due

principally to the presence of filamentous algae. Taken as a whole,

the mud bottom habitats in shallow water are poor in molluscan

inhabitants.

Associated anim.als are 52 per cent more numerous in individuals

than the mollusks. Three groups make up 68 per cent of the asso¬

ciated animal life ; Hyalella knickerbockeri 28 per cent, Asellus in-

termedius 19 per cent, and Chironomid larvae 39 per cent. The

other eleven groups are poorly represented. Among Ephemerids,

Caenis, Heptagenia, and Ephemera occur rarely. Of Odonata,

nymphs of Enallagma, Anax junius, and Tetragoneuria semiaequa

are represented by a few individuals; Sialis infumata was found

only in habitat number 36, in a small bay south of Fahney Bay.

Of Hemiptera, the large Banatra fusca and a Belostoma occurred

in one habitat, the mouth of a small creek near Eweco Park, south

124 Wisconsin Academy of Sciences, Arts, and Letters.

of Oshkosh; Coleoptera is represented by adults of Hydrovatus

pustulatus, Haliplus ruficollis, Stenelmis hicarinatus, and Bidessus

flavicollis. Acarina includes Limnesiopsis, Fiona, Limnesia, and

Arrhenurus, all rare and occurring mostly at station 36, the small

pool behind the beach at Roe Point. Leptocerids among Trichoptera

are uncommon ; and a single larva of the Lepidopterid Nymphula

was found at station 23, the marsh behind Asylum Point. Leeches

occur at five stations and include Glossiphonia stagnalis, G. neph-

eloides, G. fusca, Erpohdella punctata, and Dina parva.

The Vegetation Areas. Plants, owing to the shallowness of the

water, are very abundant in Lake Winnebago, and occur com¬

monly to a depth of two meters. Beyond this depth they decrease

very rapidly. No plants (excepting algae and microscopic forms)

were found below 2.5 meters and no filamentous algae below 3

meters. The great plant areas are in shallow water, 0.3 to 1.5

meters in depth, in bays and along the margins of shores. Plant

zones of greater or less size extend entirely around the lake, bor¬

dering the shore. All of the bays contain an extensive flora which

supports a large and varied fauna. Many individuals collected

from the bottom are migrants, either by intention or accident, from

these plant areas. All kinds of bottom, except boulder, support an

extensive flora, and some plants occur on this inhospitable sub¬

stratum, as at habitat 43 where two species of Potamogeton and a

Vallisneria occurred.

Plants serve three very useful purposes for the bottom fauna.

First, they form a binding medium which prevents the bottom

from shifting ; second, they form a means of attachment and sup¬

port for the crawling and clinging members of the fauna; and

third, they provide a foraging ground, either directly by their own

material, or indirectly by harboring many small animals used as

food by other members of the fauna or by providing support for

filamentous algae, the principal food supply of many forms of

animal life. The statement so often made by biologists — that an

abundant fauna is dependent upon an abundant flora — is strik¬

ingly emphasized in Lake Winnebago, for the animal population is

very large per unit area and the flora is equally luxuriant. The

lake’s large and varied fauna is due to the shallow depth which

enables a rich flora to become established.

Baker — The Fauna of The Lake Winnebago Region. 125

The aquatic flora may be divided broadly into two groups,

emergent and submerged. These may be differentiated as follows :

Emergent

Submerged

Zizania aquatica

Scirpus validus

Scirpus occidentalis

Castalia odorata

Nymphaea advena

Lemna trisulca

Potamogeton natans

Vallisneria spiralis

Elodea canadensis

Ceratophyllum demersum

Myriophyllum verticillatum

Potamogeton pectinatus

Potamogeton richardsoni

Potamogeton zosterifolius

Potamogetom lucens

Chara

Cladophora

No attempt was made to obtain a list of all the species of aquatic

plants, only the common and conspicuous ones being included. The

lake offers a wide field for botanists and a complete list of the flora

would be quite extensive.

Animal life was four times as abundant in vegetation as on the

sand bottom areas, which was the richest region of the bottom

areas. The number of species represented, however, was not as

great (sand 40, vegetation 25). Among mollusks, four species

make up 91 per cent of the total population, Amnicola limosa

porata 21 per cent, Valvata tricarinata 48 per cent, Physa sayii 13

per cent, and Planorhis parvus 9 per cent. Among associated ani¬

mals, which are more than four times as abundant in individuals

as the mollusks, three groups make up 87 per cent of the entire

population ; Hyalella knickerhockeri 41 per cent, Chironomid

larvae 31 per cent and Oligachaete worms 15 per cent.

Several animals prefer certain species of plants as a habitat.

Thus the broad leaves of Nymphaea and Castalia, principally on

the under side, are tenanted by Physa sayii, Planorhis parvus,

Pldnorhis antrosus, Planorbis campanulatus, Amnicola limosa

porata, Amnicola walkeri, Valvata tricarinata, young Bythinia

tentaculata, and Lymnaea winnebagoensis. Among associated ani¬

mals such species as Donacia proxima, Planaria maculata, the

larvae of Nymphula, and the eggs of Donacia and Gyrinus are

common. Some Hyalella are always found on these leaves. Scirpus

is the natural home of Ferrissia parallela and Bythinia tentaculata.

126 Wisconsin Academy of Sciences, Arts, and Letters.

as well as the bryozoan Plumatella polymorpha. Elodea is used

by Physa sayii, Bythinia tentaculata and Planorbis parvus among

snails, and by Anax junius, Enallagma, and other insects. The

Potamogeton leaves serve as a resting place or foraging ground

for many mollusks and associated animals. Vallisneria, with its

long, narrow leaves, is a favorite resort of Amnicola and young

Lymnaea and Planorbis.

But over and above all the filamentous algae are the great forag¬

ing grounds of both mollusks and associated animals. Algae, in

many places, cover all upright plants like a huge blanket (hence

often called blanket algae) and among them many animals occur

in great abundance. No less than 19 higher groups of animals

have representatives that live among the tangled masses of Clado-

phora. Planaria maculata, leaches of several species, Glossiphonia

fusca, stagnalis, complanata, nepheloidea, Dina fervida, and Oli-

gochaete worms, Stylaria, are at times very abundant; minute

Cladocera of several species are common; the Amphipod, Hyalella

knickerhockeri, is the most abundant animal and with this is asso¬

ciated the Isopod, Asellus intermedins ; among Ephemerids, Caenis

is common; the nymphs of Corixa, Belostoma, Notonecta, and Plea

striola are more or less abundant, with adults of Plea and Ranatra

fusca; Trichoptera larvae, including Agraylea, Hydropsyche,

Oecetis, Leptocella, Phryganea, Polycentropidae, and a few Helico-

psyche, are common or abundant; Chironomids, next to Hyalella,

are the most abundant, their larvae and pupae occurring in count¬

less numbers in the mass of algae; Coleoptera include Bidessus

affinis and B. flavicollis, as well as the larvae of Dytiscids and

Dascyllidae; the mites (Acarina) are the best represented as re¬

gards genera, of which eleven have been identified, including

Limnesia, Hydrachna (common), Tayas, Piona (not common),

Hygrobates, Lehertia porosa, Torrenticola, Limnesiopsis, Eylais,

Unionicola, and Arrhenurus (rare).

Nearly all aquatic mollusks frequent algal communities. By¬

thinia, young Amnicola, Valvata, and Planorbis, especially the

smaller species, browse among the stringy filaments. As already

noted in Oneida Lake (Baker 1918, p. 158) some species live in a

plant habitat when young and later migrate to a different kind of

a habitat. Pleurocera acuta when young is found in algae but

later migrates to the boulder or gravel shores, where algal food

may be gleaned from rocks. Lymnaea winnehagoensis also lives

in algae when young and later occupies a sand, gravel or boulder

Baker — The Fauna of The Lake Winnebago Region. 127

habitat. As observed in Oneida Lake (Baker, 1918, p. 151) many

of the animals were of a peculiar green color due to the algae

they had eaten. Even the shells of mollusks appeared green when

containing the living animal.

Habitats in Water Deeper than Two Meters (Table 1.) Be¬

yond the two meter contour both plant and animal life becomes

greatly reduced in both number and kind. Upright plants were

not found in Lake Winnebago below three meters. The fauna cor¬

respondingly decreases. Among Naiades only one species was

found deeper than 3.4 meters, Lampsilis luteola rosacea at station

69 in water 5.2 meters deep on a mud bottom. Sphaerium de¬

scends to four meters, and Pisidium alone among Pelecypods fre¬

quents the deepest parts of the lake, about six meters, where it is

abundant. Most of the gastropods disappear between three and

four meters. Amnicola limosa porata descends to 5.5 meters and

is abundant; Valvata tricarinata is rare at 6.1 meters.

There is a rapid decrease in number of species as the depth of

the water increases. Thus, between 2 and 3 meters, 63 species

occur ; between 3 and 4 meters, 32 species ; 4 to 5 meters, 29 species ;

and 5 to 6 meters, 13 species. This decrease is greater for Lake

Winnebago than for Oneida Lake, the percentages being as fol¬

lows :

Lake Winnebago

Oneida Lake

86 per cent.

60 per cent.

44 per cent.

40 per cent.

2 meters, 85 per cent.

3 meters, 43 per cent.

4 meters, 39 per cent.

5 meters, 17 per cent.

The depth areas seem divisible into three subregions: Littoral

to a depth of three meters where rooted plants cease to grow; sub¬

littoral to a depth of four meters where Cladophora and other

' algae cease to grow; and aphytal from four to six meters where

plant life (except plankton algae) ceases to grow. The decrease

in plant and in animal life with depth thus appears to be co¬

incident.

The associated animals show aboi^t the same decrease with

depth as the mollusks, although a greater variety inhabit deeper

water. Chironomid larvae descend to a depth of over six meters

and appear to bear the same relation to the associated animals that

Pisidium does to the Mollusca. Leeches ( Glossiphonia and Dina)

128 Wisconsin Academy of Sciences, Arts, and Letters.

and Oligochaete worms also descend to six meters or more. This

fauna is not large. Hydra oligactis occurred at two habitats on

a gravel bottom and Planaria maculata was found in the same

places; Hyalella was rare in the deeper habitats; Ephemera oc¬

curred at 2.5 meters, Caenis at 4 meters, and Hexagenia at 5.5

meters ; Sialis infumata was rare ; Corixa and Plea striola, nymphs,

were rare; Trichoptera (Molanna, Agraylea, Leptocella, Hydro¬

psyche and Helicopsyche) disappeared at 3.1 meters. A single

larva of Nymphula was dredged at a depth of 4 meters on a mud

bottom. Coleoptera included Psephenus lecontei at 2.8 meters,

Hydrovatus pustulatus at 4 meters, and Dascyllid larvae down to

3.4 meters, none common. Hygrobates was found at 2.5 meters,

and Limnesiopsis at 3.1 meters.

Lake Butte des Morts (Table 1).

Lake Butte des Morts is about five miles long and two and a half

miles wide with a maximum depth of 4.6 meters. It is but a wid-

ened-out lower portion of the Fox Eiver, though carrying ^so the

waters of Wolf Kiver and several creeks. Most of the ecological

conditions of a lake environment are present. Boulder shores are

rare, occurring on such places as Plummers Point and Sunset

Point, where, however, sand and gravel are quite as common and

are usually closely associated with boulders. The shores in the

bays are usually marshy with an abundant growth of sedge, cat¬

tails, and other aquatic plants. In many places this kind of shore

is fully a quarter of a mile in width. The bottom of the bays is

usually of mud, clay, or fine sand covered with a thick layer of

plant debris. Sand occurs along some shores, as west of Plummers.

Point, as well as on the bottom of the marsh areas at the west end

near the Pox River channel. The same vegetation occurs as in

Lake Winnebago.

About the same kind of mollusks occur in this lake as in Lake

Winnebago. A few species, as Nephronaias carinata, Amhlema

costata, and Amnicola limosa, indicate the influence of the Fox

River fauna. As in Lake Winnebago, the number of species de¬

creases with increased depth, 51 species being found between shore

and 2 meters, 12 species between 2 and 3 meters, and but 3 species

below 3 meters. Three species make up 53 per cent of the total

molluscan population, Amnicola limosa 22 per cent, Bythinia 24

per cent and Valvata 7 per cent.

Baker — The Fauna of The Lake Winnebago Region. 129'

Associated animals a little more than equal the mollusks in

abundance. They include a number of genera divided among

fourteen higher groups. Two groups make up 65 per cent of the

total, Hyalella, 43 per cent and Chironomid larvae 22 per cent.

Caenis occurred at 2 and 3.4 meters and Hexagenia was found on

a mud bottom in 1 to 3.4 meters. Enallagma and Anax junius

were included among Odonata, both rare. Nymphula was rare at

all but one station. Plea striola, adult and nymph, was not un¬

common. Trichoptera included Agraylea, Leptocella, Helico-

psyche, Molanna, and Polycentropidae, which were not found

deeper than 1.5 meters. Among Coleoptera, Bidessus flavicollis,

Stenelmus hicarimatus, Donacia proxima, and a Dascyllid larva

occurred uncommonly in shallow water (1 meter) excepting

Donacia which was found at 3 meters. The mites were very rare

and included Hydrachna, Limnesiopsis, and Unionicola, mostly in¬

habitants of shallow water. Pour species of leeches occurred in

the deepest water on mud and sand bottom, Glossiphonia stagnalis,.

nepheloidea, fusca, Erpohdella punctata.

The vegetation population of this lake is large but not particu¬

larly varied. Only thirteen species of mollusks, of which Bythinia,,

Amnicola limosa, and Ferrissia parallela form about 79 per cent,,

were collected from plants as compared with 25 species in Lake

Winnebago.

Among associated animals, Hyalella and Chironomid larvae

make up 56 and 36 per cent respectively, but are otherwise poorly

represented. At one station (93) the filamentous algae were fairly

alive with Hyalella and Chironomid larvae. The genera repre¬

sented in the vegetation include Plea striola and Belostoma

nymphs; Hexagenia nymphs; Enallagma nymph; Agraylea, Heli-

copsyche, Leptoceridae larvae ; Nymphula larva ; Donacia proxima^

Bidessus fiavicollis, B. affinis, Dytiscid larva; Hydracna, Lim-

nesia ; Asellus intermedius ; Plumatella polymorpha; Hydra oligac-

tis, and Glossiphonia and Erpobdella.

Fox River (Table 1).

Dredgings were made near Omro, the maximum depth found be¬

ing 2.9 meters. The bottom is of mud, sand, clay, or gravel, mud

occurring in all protected places. Gravel occurred near Omro in

water 2.9 meters deep in which three species of Naiades were

found. The shores of the river are swampy for the most part, low,.

130 Wisconsin Academy of Sciences, Arts, and Letters,

and bordered by marsh vegetation. Aquatic plants grow in the

shallower parts of the river. Mollusks were most numerous near

the shore on a sand or mud bottom in water 0.5 to 1 meter in depth.

The Naiades are mostly different from the lake species as noted

under Lake Winnebago species. The gastropods are mostly of

species living in quiet habitats and there is not the variety found

in the lakes, the latter being 50 per cent richer in species. Of the

population of the river habitats Sphaerium striatinum makes up

67 per cent, forming great beds bordering the shore in many places.

Quantitative Analyses of the Fauna

On rocky shores, a number of boulders were measured, the total

animal life removed, and the area computed to the square meter

unit. In the vegetation stations the total area covered by the plants

was computed to the same unit. In several cases, as in Scirpus

habitats, the unit included a column from surface to bottom.

The population of Lake Winnebago compares favorably with

that of any lake studied, being exceeded only by Oneida Lake

among American inland lakes. The average number of animals

per unit area on the different kinds of bottom in the two lakes

mentioned is shown in table 2.

It will be seen that among the Mollusca, Oneida Lake is 59 per

cent richer, while among associated animals. Lake Winnebago is

9 per cent richer in individual population. The small population

of the vegetation habitats in Oneida Lake is due to the fact that

much of the population is included in the bottom areas which had

an algal covering.

Table 3 shows that Oneida Lake has a greater population per

unit of area in the shallower water, but that Lake Winnebago has

a greater population in the deeper water. Lake Mendota has a very

small population in the shallow water as compared with the other

lakes. Lake Butte des Morts has the largest population in the

deeper water as well as the largest total population per unit area.

The molluscan fauna of the Winnebago region is one of the

most extensive and varied of any similar area yet studied. A total

of 114 species and varieties of fluviatile and lacustrine forms were

found, including three forms believed to be new to science. Table

4 shows the relative abundance of the molluscan fauna in the dif¬

ferent parts of this region.

Baker — The Fauna of The Lake Winnebago Region, 131

Table 5 shows the relation of the number of species of Mollusca

to the depth of the water. The greatest variety is found at depths

not exceeding two meters.

Table 6 shows the relation of the number of species of Mollusca

to the character of the bottom in Lake Winnebago. The maximum

number was found on sand bottom.

Table 7 shows the variety of the molluscan faunas of three lakes,

namely, Oneida Lake, New York ; Maxinkuckee Lake, Indiana ; and

Lake Winnebago, Wisconsin.

One of the interesting features brought out by the study of the

molluscan fauna of the Lake Winnebago region is the difference

in size and shape between the Unionidae of the Fox River and

those of the lakes, a difference which appears to be comparable to

that noted by Grier (1919) between the Naiads of Lake Erie and

the upper drainage of the Ohio River. Grier states that “if we put

a shell in the lake environment we may expect it will change its

morphological features, not at random, but in a distinct, deter¬

minate, or orthogenetic direction. ’ ’ This change in the morphology

of shells that have migrated from a river to a lake is strikingly

shown in the Lake Winnebago fauna, and a study of the two areas

by the methods of Grier would produce the same results as attained

by the study of the Lake Erie shells. It is a significant fact that

the same varietal forms inhabit both Lake Winnebago and Lake

Erie, indicating that the law holds good under similar conditions

in widely separated areas. Dr. Sterki notes that almost all of the

Pisidium of the lake are small and slight, some even depauperate,

and Sphaerium and Musculium are similarly affected. The gastro¬

pods of the lake are in the main different from those of the river.

Thus the entire molluscan fauna is affected by the same law of

variation produced by river and lake environment, clearly indi¬

cating that ecological station plays a large part in the evolution of

species. A study of the tables and of the systematic list which fol¬

lows will bring out additional features of this ecological character¬

istic. Just what factors have been potent in producing these

changes does not seem to be definitely known. It is probable that

variation in food supply, in the chemical character of the fluid

medium in which they live, as well as in the general physical

environment, plays a large part in these changes of form.

132 Wisconsin Academy of Sciences, Arts, and Letters.

LIST OF MOLLUSCA

Unionidae

Dysnomia triquetra Eaf. A single small individual was dredged from the

Fox Eiver at Omro. This is the first record for Wisconsin, and the northern¬

most record for any locality. It occurs in southern Michigan.

Lampsilis ventricosa (Barnes). Common in the Fox Eiver at Omro and

other places.

Lampsilis ventricosa canadensis (Lea). Lakes Winnebago and Butte des

Morts, abundant. Not before reported from Wisconsin.

Lampsilis siliquoidea (Barnes). Common in the Fox Eiver at Omro and

other places.

Lampsilis siliqudidea rosacea (DeKay). Lakes Butte des Morts and Winne¬

bago, abundant. First authentic records from Wisconsin.

Ligumia recta (Lamarck). Lake Winnebago, common. The typical form

from Lake Erie seems to be represented in the lake, where the individuals

are smaller and more brightly colored and rayed than the large, black speci¬

mens from the rivers.

Ligumia recta latissima (Eaf.). Fox Eiver at Omro and other places. The

large river form. Abundant.

Proptera alata (Say). Common in Lake Butte des Morts and Lake Win¬

nebago. As noted by Ortmann (1920) and others the Lake Erie form of alata

is smaller and more swollen than the river forms and the epidermis is usually

browner. The Winnebago Lake shells are of this form.

Proptera alata megaptera (Eaf.). Fox Eiver and other places, common.

A much larger shell with more compressed valves.

Leptodea fragilis (Raf.). Omro and other places on the Fox Eiver,

abundant.

Leptodea fragilis lacustris (Baker). Lakes Winnebago and Butte des Morts,

common. The lake shells differ markedly from the river shells in color and

form. (Baker, 1922, p. 131.)

Truncilla trunoata (Eaf.). Omro, Fox Eiver, common. Lake Butte des

Morts, rare.

Truncilla truncata (Eaf.) variety. A form of truncata occurs in Lake

Winnebago which is somewhat smaller than the river type of this species.

The umbones are less elevated, the valves are rounder and more compressed and

the posterior ridge is much less developed. Not enough material is at hand

to differentiate this from the typical species.

Actinonaias carinata (Barnes). Omro, Fox Eiver, common; Lake Butte

des Morts, rare.

Strophitus rugosus (Swainson). Omro, Fox Eiver, rare.

Strophitus rugosus (Swainson). Var. Lakes Winnebago and Butte des Morts,

not common.

Anodonta imljecillis Say. Omro, Fox River, common.

Anodonta grandis gigantea Lea. Omro, Fox Eiver, common.

Anodonta grandis footiana Lea. Lakes Winnebago and Butte des Morts,

abundant. This is the type locality for footiana.

Baker — The Fauna of The Lake Winnebago Region. 133

Lasmigona complanata (Barnes). Omro, Fox Eiver. The type locality is

the Fox Eiver.

Lasmigona costata (Eaf.). Omro, common and typical.

ElUptio dilatatus (Eaf.). Omro, not common.

Elliptio dilatatus delicatus (Simpson). Omro, not common.

Elliptio dililatus sterTcii Grier. Lakes Winnebago and Butte des Morts, com¬

mon. The lake forms are all smaller and seem in every way like the Lake Erie

forms described by Grier (1918).

Pleurobema coccineum (Conrad). Omro, not common.

Quadrula quadrula (Eaf.). Omro, not common.

Quadrula verrucosa (Eaf.). Omro, not common.

, Amblema costata (Raf.). Omro, Fox Eiver, Lake Butte des Morts, rare.

Fusconaia flava (Eaf.). Omro, not common.

Fusconaia flava parvula Grier. Lake Winnebago, not common. There is

great variation in the diameter of the shells of this form.

Sphaeriidae •

Sphaerium sulcatum Lam. Common in Winnebago and Butte des Morts.

Sphaerium sulcatum planatum St. Found only in Lake Winnebago.

Sphaerium lineatum St. In debris on Doemel Point, Lake Winnebago. Only

one specimen found of this species, which seems rare in the lake.

Sphaerium striatinum Lam. Lakes Winnebago, Butte des Morts and Fox

Eiver. The species shows great variation and there may be a variety that

can ultimately be separated when additional material is available.

Sphaerium lilycashense Baker. Fox Eiver, rare.

Sphaerium ohioense St. Lake Winnebago, not common.

Sphaerium flavum Prime. Lake Butte des Morts, rare.

Sphaerium solidulum Prime. Lakes Winnebago and Butte des Morts, not

common.

Sphaerium stamineum Conrad. Lake Winnebago and Fox Eiver, not com¬

mon.

Sphaerium occidentale amphibium St. Swale in woods on Plummers Point,

Lake Butte des Morts. Abundant.

Musculium transversum Say. Lakes Winnebago and Butte des Morts, not

common.

Musculium truncatum Linsley. Lake Buttes des Morts, not common.

Musculium jayense Prime. Lakes Winnebago and Butte des Morts, not

common.

Psidium virginicum Gmelin. Lake Butte des Morts, one specimen, juv.

Pisidium compressum Prime.

Pisidium compressum pellucidum St. Lakes Winnebago and Butte des Morts,

common, the variety pellucidum rarer.

Pisidium fallax St. Lake Winnebago, not common.

Pisidium punctatum simplex St. Lake Winnebago, rare.

Pisidium variabile Prime. Lakes' Winnebago and Butte des Morts, common.

Pisidium pauperculum St. Same as above, not common.

Pisidium glabellum St. Lake Butte des Morts, very rare.

134 Wisconsin Academy of Sciences, Arts, and Letters.

Pisidium minusculum St. Lake Butte des Morts, rare. Types of this species

were from the Fox Eiver.

Pisidium sargenti St. Lakes Winnebago and Butte des Morts, not common.

Pisidium adamsi affine St. Lakes Winnebago and Butte des Morts, rare.

Pisidium strengi St. Same localities as above, not common. Pond in Ter¬

rel’s gravel pit.

Pisidium decorum St. Lake Butte des Morts, rare.

Pisidium griseolum St. MS. Lake Winnebago, very rare.

Pisidium milium Held. Lake Winnebago, rare.

Pisidium walTceri St. Lake Winnebago, not common.

Pisidum tenuissimum St. Lakes Winnebago and Butte des Morts, common.

Pisidium scutellatum cristatum St. Lake Winnebago, not common.

Pisidium vesiculare St. Lake Winnebago, not common..

Pisidium medianum St. Lake Winnebago, common.

Pisidium clavatum St. MS. Lake Winnebago, not common.

Pisidium splendidulum Sterki. Lake Winnebago, rare.

Pleuroceridae

Pleurocera acuta Eaf. Very abundant in Lake Winnebago.

Amnicolidae

Somatogyrus suhglobosus (Say). Omro, Fox Eiver, rare. Lakes Winnebago

and Butte des Morts, common.

Amnicola limosa (Say). Omro, Fox Eiver, common. Lake Butte des

Morts, common.

Amnicola limosa porata (Say). Lake Winnebago, very abundant and the

dominant form.

Amnicola judayi Baker. Lakes Winnebago and Butte des Morts. Not com¬

mon.

Amnicola walkeri Pilsbry. Lakes Winnebago and Butte des Morts. Common.

Amnicola lustrica Pilsbry. Lake Winnebago, rare.

Amnicola emarginata Kiister. Common in Lakes Butte des Morts and Win¬

nebago.

Bytliinia tenaculata (Lin.) Lakes Winnebago and Butte des Morts, very

common, especially in vegetation.

Valvatidae

Valvata tricarinata (Say). Lakes Winnebago and Butte des Morts. One of

the most abundant mollusks in these lakes. Five varieties have been noted in

the lake material; their relative abundance is shown in table 1.

F. tricarinata simplex Gould

F. tricarinata hasalis Vanatta

F. tricarinata unicarinata DeKay

F. tricarinata supracarinata Baker

F. tricarinata perconfusa Walker

Baker — The Fauna of The Lake Winnebago Region, 135

ViVAPARIDAE

Liliplax subcarinata (Say). Lakes Winnebago and Butte des Morts, and

Omro, Fox Eiver, common.

Campeloma rufum (Haldeman). Same localities as above.

Physidae

Fhysa sayii Tappan. Lake Winnebago and Butte des Morts, common.

These Physas are smaller than sayii from other localities, especially river

habitats, and may be the result of a lake environment, as has been the case

with so many other lake forms herein listed.

Fhysa integra Haldeman. Terrell’s gravel pit, near Lake Butte des Morts,

in small stream flowing from artiflcial pond. Bare.

Fhysa integra hillingsi Heron. The Lake Winnebago and Butte des Morts

specimens of integra seem referable to hillingsi. Common.

Fhysa gyrina hildrethiana Say. Swale half mile north of Oshkosh.

Fhysa gyrina oleacea Tryon. Summer-dry pond in woods on Plummers

Point, near Lake Butte des Morts. Abundant.

Aplexa hypnorum (Linn.). Swale half mile north of Oshkosh.

Planorbidae

Flanorhis trivolvis Say. Omro, Fox Eiver, among vegetation, common. Lakes

Butte des Morts and Winnebago in protected situations.

Flanorhis trivolvis fallax Haldeman. Lakes Winnebago and Butte des Morts.

In more exposed situations than typical trivolvis.

Flanorhis trivolvis pseudotrivolvis Baker. Lake Winnebago and Butte des

Morts in marshy places. Listed as Flanorhis glahratus by Andrews (1915,

p. 200).

Flanorhis truncatus Miles. Lakes Winnebago and Butte des Morts. The

most abundant Planorbis in the lake.

Flanorhis campanulatus Say. Lakes Winnebago and Butte des Morts, com¬

mon.

Flanorhis antrosus Conrad. Same localities as above. Abundant only in

places.

Flanorhis antrosus striatus Baker. Same localities. The majority of the

antrosus are referable to the striate form.

Flanorhis deflectus Say. Lakes Butte des Morts and Winnebago, common.

Flanorhis exaeuous Say. Same localities as above, not common.

Flanorhis parvus Say. Common in the two lakes. Eare at Omro, Fox Eiver.

Flanorhis umhilicatellus Ckll. Shallow pool under railroad bridge on shore

of Lake Butte des Morts, north of Oshkosh, rarej swale north of Oshkosh, rare;

pool on Plummers point, in woods. Lake Butte des Morts, common. The first

record for Wisconsin.

Segmentina armigera (Say). Small pool beneath railroad tracks north of

Oshkosh on shore of Lake Butte des Morts, not common.

136 Wisconsin Academy of Sciences, Arts, and Letters.

Lymnaeidae

Lymnaea stagnalis appressa Say. Fox Eiver, Omro, abundant. Lakes Win¬

nebago and Butte des Morts in protected places, common.

Lymnaea winnebagoensis Baker. Common in the two lakes. (Baker, 1922,

p. 22.)

Lymnaea elodes Say. Omro, Fox River. On vegetation in quiet, pond-like

stretches of the river, associated with stagnalis. Common. Recorded by An¬

drews from Lake Butte des Morts (1915, p. 200).

Lymnaea palustris (Muller). Hatchery Bay among algae; Doemel Point in

cat-tail pond behind point. Recorded by Andrews from marsh bordering Lake

Butte des Morts (1915, p. 200).

Lymnaea reflexa Say. Swale north of Oshkosh; swale in woods on Plum¬

mers Point, Lake Butte des Morts; artificial pond in Terrell’s gravel pit, Lake

Butte des Morts.

Lymnaea obrussa Say. Small stream flowing from sand machine near Ter¬

rell’s gravel pit, emptying into Lake Butte des Morts.

Lymnaea obrussa exigua Lea. Pond in Terrell’s gravel pit.

Lymnaea humilis modicella Say. Hatchery Bay, Oshkosh, on leaves of

water lily (Castalia). Rare.

Lymnaea parva Lea. Marshy border of Lake Butte des Morts, north of

Oshkosh. Rare.

Lymnaea caperata Say. Shallow pool beneath railroad track north of Osh¬

kosh; small stream flowing into Miller Bay, Lake Winnebago; pool in woods

on Plummers Point, Lake Butte des Morts.

i Ancylidae

Ferrissia parallela (Haldeman). Lake Butte des Morts, near Plummers

Point on leaves of Scirpus and Nymphaea; Lake Winnebago, near Long Point

Island, on Scirpus leaves.

Terrestrial Pulmonata

Helicidae

Polygyra multilineata algonquinensis Nason. Long Point Island, Lake Win¬

nebago, and Plummers Point, Lake Butte des Morts, common.

Polygyra profunda (Say). Long Point Island, Asylum Point, and Dormel

Point, Lake Winnebago, not common.

Polygyra monodon (Rackett). Doemel Point, Lake Winnebago; Plummers

Point, Lake Butte des Morts, not common.

Polygyra fraterna (Say). Doemel Point, Lake Winnebago, rare.

ZONITIDAE

Vitrea rhoadsi Pilsbry. Doemel Point, Lake Winnebago; woods on Plum¬

mers Point, Lake Butte des Morts, rare.

Zonitoides arborea (Say). Woods on Plummers Point, Lake Butte des

Morts, abundant.

Baker — The Fauna of The Lake Winnebago Region, 137

Zonitoides nitida (Muller). Doemel Point, Roe Point, lake shore near Osh¬

kosh, Lake Winnebago, common.

Endodontidae

Pyramidula alternata (Say). Doemel Point, Lake Winnebago; woods on

Plummers Point, Lake Butte des Morts, abundant.

Pyro/midula cronTchitei anthonyi Pilsbry. Woods on Plummers Point, com¬

mon.

Helicodiscus lineatus (Say). Woods on Plummers Point, common.

SuCCINEIDAE

Succinea ovalis Say. Woods on Plummers Point, common.

Succinea retusa Lea. Swale north of Oshkosh; Par Eockaway Point, shore

near fish hatchery, Oshkosh; shore of Lake Butte des Morts, north of Oshkosh,

abundant.

Succinea avara Say. Common near shores of both lakes.

Succinea avar major W. G. Binney. Far Eockaway Point, Lake Winnebago,

specimens 12 mm. in length.

PUPILLroAE

Strobilops virgo (Pilsbry). Woods on Plummers Point, Lake Butte des

Morts.

StroMlope ajfinis (Pilsbry). Common, associated with above.

Gastrocopta contracta (Say). Doemel Point, Lake Winnebago; woods on

Plummers Point.

Vertigo ovata Say. Doemel Point, Lake Winnebago.

Auriculidae

Carychium exiguum canadense Clapp. Common in woods on Plummers Point.

ANIMALS ASSOCIATED WITH THE MOLLUSCA

The following list includes the species or genera of the invertebrate animals

found associated with the Mollusca. The list does not include all species that

inhabit the territory under consideration, only those taken in dredgings. The

names of those specialists who determined the groups are given in the intro¬

duction. All material is from the lakes.

Hydrozoa

Hydra oligactis Pallas

PORIFERA

Spongilla fragilis Leidy

138 Wisconsin Academy of Sciences, Arts, and Letters.

Turbellaria

Planaria maculata Leidy

Nematoda

Genera et species undetermined. Very small worms.

Stylaria lacustris (Linn).

Naias sp?

Dero sp? (Winnebago only)

Sparganophilus eiseni Smith (Lake Butte des Morts only).

Limnodrilus species, possibly hoffmeisteri Claparede.

Lumhriculus probably inconstans Smith.

Hirudinea

Glossiphonia complanata (Linn).

Glossiphonia nepheloidea (Graf).

Glossiphonia stagnalis (Linn).

Glossiphonia fusca Castle.

Variety lineata,

Variety papillata.

"Dina fervida (Verrill).

Dina parva Moore.

Dina macrostoma Moore.

JErpohdella punctata Leidy.

Nephelopsis obscura Verrill.

Placobdella phalera (Graf).

Cladocera

Eurycercus lamellatus Muller.

Simocephalus serrulatus Koch.

Simocephalus vet ulus Muller.

Daphnia retrocurva Forbes.

Sida crystallina Muller.

Diaphanosoma brachyurum (Lieven),

Bryozoa

Plumatella polymorpha Kraepelin.

OSTRACODA

Cypria species. Cypridopsis species.

Amphipoda

Eyalella TcnicTcerbocTceri Bate. Gammarus fasciatus Say.

ISOPODA

Asellus intermedius Forbes.

Decapoda

Cambarus propinquus Girard.

Baker— The Fauna of The Lake Winnebago Region. 139

Acarina (Hydracarina)

Eylais.

Hydrachna.

Thyas eatapJiracta Koenike.

Arrhenurus americanus Marshall.

Torrenticola.

Lebertia.

Lehertia porosa Thor.

Limnesiopsis sp. nov.

Limnesia histrionica wolcotti Piersig.

Limnesia paucispina Wolcott.

Unionicola crassipes (Miill.) 9

Fiona reighardi Wolcott.

Fiona turgidus (Wolcott).

Hygrobates.

Ephemera

Ephemerella

Gomphus

Tramea

Anax junius Drury.

Ephemerida (Nymphs)

Callibaetis Hexagenia

Caenis Heptagenia

Odanata (Nymphs)

Tetragoneuria semiaqua Burm.

Plathemis

Enallagma

Hemiptera

Coriza, adults and nymphs.

Belostoma, adults and nymphs.

Gerris marginatus Say.

(Adults and nymphs)

Flea striola Eieber.

Notonecta undulata Say.

Banatra fuSoa Beauv.

Neuroptera (Larvae)

Sialis infumata Newman.

Trichoptera (Larvae)

Hydroptila Agraylea Hydropshche

Molanna Phryganea Polycentropus

Leptocella Leptocerus Oecetis

Helicopsyche 'borealis Hagen.

Calaclysta

Lepidoptera (Larvae)

Nymphula

COLEOPTERA

Gyrinus ventralis Kirby.

Cnemidotus edentulus Lee.

Ealiplus ruficollis DeGeer.

Bidessus flavicollis Lee.

Bidessus affinis Say.

Hydrovatus pustulatus Melsh.

Comptotomus interrogatus Pabr.

Laceophilus maculosus Germar.

(Mostly adult)

Agabus gagates Aube.

Berosus peregrinus Herbst.

Fsephenus leeontei Hald. Larva.

Stenelmis bicarinatus Lee.

JDonacia proxima Kirby.

Dascyllidae. Larva of unknown spe¬

cies.

Chironomus

Diptera (Larvae)

Palpomyia

Psyehodid pupa

Tanypus

140 Wisconsin Academy of Sciences^ Arts, and Letters,

LITEEATUEE CITED

Alden, W, C. 1918, The quaternary geology of southeastern Wisconsin.

Professional Paper No. 106, TJ. S. Geol. Survey.

Andrews, Olive V. 1915. An ecological survey of the Lake Butte des

Morts bog, Oshkosh, Wisconsin. Bui. Wis. Nat. Hist. Soc. 13: 196-211.

Baker, Frank C. 1916. The relation of mollusks to fish in Oneida Lake.

N. Y. State Coll. For. Syracuse Univ. Tech. Pub. 4: 15-366.

- . 1918, The productivity of invertebrate fish food on the bottom of

Oneida Lake, with special reference to mollusks. N. Y. State Coll.

For. Syracuse Univ. Tech, Pub. 9: 11-233.

- -. 1920. The effect of sewage and other pollution on animal life of

rivers and streams. Trans. Ill. Acad. Sci. 13: 271-279.

- . 1922. a. New species and varieties of Mollusca from Lake Win¬

nebago, Wisconsin, with new records from this state. Nautilus 36:

19-21.

- . 1922. b. New Lymnaeas from Wisconsin and Minnesota, with

notes on shells from the latter state. Nautilus 36: 22-25.

Chadwick, G. H. 1902. Notes on Wisconsin Mollusca. Bui. Wis. Nat.

Hist. Soc. 4: 67-99.

Ekman, Sven. 1915. Die Bodenfauna des Vattern. Internal. Eev. 7:

146-204, 275-425.

Evermann, B. W. and Clark, H. W. 1920. Lake Maxinkuckee. A physical

and biological survey. Department of Conservation, State of Indiana.

Pub. No. 7, 2 vols. Indianapolis.

Grier, N. M. 1918. New varieties of Naiades from Lake Erie. Nautilus

32: 9-15.

- . 1919. Morphological features of certain mussel-shells found in

Lake Erie, Ann. Carnegie Mus. 13: 145-182.

Goldthwait, J. W. 1907. The abandoned shore lines of eastern Wisconsin.

Bui. 17, Wis. Geol. and Nat. Hist. Survey. 134 pp. Madison.

Juday, Chancey. 1914. The inland lakes of Wisconsin. — Hydrography and

morphometry, Bui. 27, Wis. Geol. and Nat. Hist. Survey. 137 pp.

Madison.

Martin, Lawrence. 1916. The physical geography of Wisconsin. Bui. 36,

Wis. Geol. and Nat. Hist. Survey. 549 pp. Madison.

Muttkowski, E. A. 1918. The fauna of Lake Mendota. Trans. Wis. Acad.

Sci., Arts, and Let. 19: 374-482.

Ortmann, A. E. 1919. A monograph of the Naiades of Pennsylvania.

Part III. Mem. Carnegie Mus. 8: No. 1.

Ortmann, A. E. and Walker, Bryant. 1922. On the nomenclature of certain

North American Naiades. Occ. Pa. Mus. Zool. Univ. Mich. No. 112.

Pearse, A. S. and Terrell, Clyde B, 1920. Aquatic preserves, Nat. Hist. 20:

103-106.

Petersen, C. G. J. and Jensen, P. B. 1911. Valuation of the sea. Animal

life of the sea bottom, its food and quantity. Eept. Danish Biol. Sta.

20: 1-76.

Whitbeck, R. W. 1915. The geography of the Fox-Winnebago valley. Bui,

42, Wis. Geol. and Nat. Hist. Survey. 105 pp. Madison.

Baker — The Fauna of The Lake Winnebago Region, 141

Table 1. This table shows the average number of individuals per square

meter in the bottom fauna of Lake Winnebago, Lake Butte des Morts and the

Fox River at Omro in the summer of 1920. The bottom has been divided into

areas according to depth and character as indicated in the headings of the

different columns.

142 Wisconsin Academy of Sciences, Arts, and Letters,

Table 1. — Continued.

Baker — The Fauna of The Lake Winnebago Region, 143

Table 1. — Continued.

144 Wisconsin Academy of Sciences, Arts, and Letters.

Table 1. — Continued.

Table 2. Average number of individuals per square meter on bottoms of

Lake Winnebago and Oneida Lake.

Baker—The Fauna of The Lake Winnebago Region, 145

Table 3. delation of Mollusca to the depth of the water.

Table 4. Distribution of the Mollusca by families in the Winnebago region.

Table 5. Delation of the number of species of Mollusca to the depth of the

water.

146 Wisconsin Academy of Sciences, Arts, and Letters,

Table 6. Belation of the number of species of Mollusca to the character of

the bottom in LaTce Winnebago.

Table 7, Comparison of the molluscan faunas of three different lakes.

*Baker, 1918.

tEverman and Olark, 1920.

OBSERVATIONS ON PARASITIC WORMS

FROM WISCONSIN FISHES

A. S. Pearse

The following descriptions of trematodes and nematodes from

Wisconsin fishes relate partly to new species and partly to those

that have been inadequately described or confused by previous

writers.

TREMATODA

AGETODEXTRA, new genus

Flattened Distomata having the acetabulum on the right of the

median line. Genital aperture between the acetabulum and the

junction of the intestinal rami ; situated somewhat toward the right

side. Cirrus sac very small. Two more or less linear testes lie on

either side at the posterior end of the body, between the intestinal

rami and the median excretory duct. Ovary, somewhat lobate,

elongated, near the center of the body. Uterus with many coils;

extending across the body from the acetabulum nearly to the pos¬

terior end. Intestinal rami reaching nearly to the posterior end of

the body.

Type species: Acetodextra amiuri (Stafford).

Acetodextra amiuri (Stafford)

(Fig. 6)

Stafford (1900) assigned this species to the genus Monostomum,

believing that no acetabulum was present, though his figure plainly

shows one. He was doubtless deceived by the fact that the acetabu¬

lum always lies to the right of the median line. The following

description supplements his :

Length: 3.6 mm.; width: 1.8 mm.; pharynx, .14 by .10 mm.;

length of esophagus : 25. mm. ; diameter of oral sucker : .185 mm. ;

diameter of acetabulum : .277 ; egg : .035 by .026 mm.

148 Wisconsin Academy of Sciences, Arts, and Letters.

The oral sucker is small; the pharynx is almost 1% times as

long as wide; esophagus, very slender, about three times as long

as pharynx; intestinal rami, thick, extending to posterior end of

the body.

Vitelline glands are in contact with the lateral surfaces of the

intestinal rami, extending from the acetabulum to the tips of the

rami; a duct from each group of glands extends directly across

the body to the middle where it unites with that from the opposite

side and enters a rounded yolk reservoir.

The testes are somewhat lobate and elongated antero-posteriorly.

They lie between the intestinal rami and the posterior, median

excretory duct. Cirrus sac : very small and ovate. The ovary lies

just anterior to the vitelline ducts. It is lobate and elongated.

The uterus is filled with eggs and occupies the space between the

intestinal rami, from the acetabulum to the posterior end of the

body.

The bullheads in Lake Pepin were often infected with this

trematode. Figure 6 was drawn from a specimen taken from the

swim bladder of Ameiurus melas, July 12, 1920 (U. S. Nat. Mus.,

Cat. No. 7618). Other specimens were found in the swim bladders

of A. natalis and A. nehulosus. Young specimens were found

encysted in the liver peritoneum of a Schilbeodes gyrinns, June

23, 1920.

MACRODEEOIDES, new genus

Elongated Plagiorchiidae with the two suckers of nearly equal

size. The genital opening is at the anterior margin of the aceta¬

bulum. A slender prepharynx and a longer esophagus are present.

The intestinal rami arise from the esophagus some distance anterior

to the acetabulum. The body is covered with sharp spines which

decrease in size posteriorly. The vitelline glands extend from a

short distance behind the acetabulum to the posterior testis. The

genus shows resemblances to Macrodera Loos, Haplometra Loos,

and Glypthelmins Stafford.

Type species : Macroderoides spiniferus, Pearse.

Macroderoides spiniferus, new species

(Pig. 9)

Type: Cat. No. 7619, U. S. National Museum; Lake Pepin,

Wisconsin; July 11, 1920; collector, A. S. Pearse.

Pearse — Parasitic Worms From Wisconsin Fishes. 149

Host: the short-nosed gar, Lepisosteus platostomus Rafinesque.

Description: Body slender; length 2.4 mm.; width, .25 mm.;

covered with sharp spines, which decrease in size posteriorly. In

the region of the pharynx there are about fifty spines on the cir¬

cumference of a cross section of the body. The diameter of the

acetabulum is slightly greater than that of the oral sucker, which

measures .08 mm. The acetabulum is at the posterior end of the

anterior seventh of the body.

A slender prepharynx is present. The pharynx measures

.012 mm. in length and .088 in diameter. The slender esophagus is

about twice as long as the pharynx. The intestinal rami are slen¬

der, join the esophagus more than the length of the esophagus

anterior to the acetabulum, and extend nearly to the posterior

end of the body.

The genital pore lies on the median line, just anterior to the

acetabulum. The testes are ellipsoidal, their longest axis being in

the same direction as that of the body. They are about equal

in size, measuring .18 mm. in length. The anterior one is situated

on the left side at the beginning of the posterior third of the body.

The posterior one is slightly toward the right side, just in front

of the posterior sixth of the body. The cirrus sac is long, thick,

and somewhat sinuous, extending from some distance posterior to

the acetabulum to the genital pore.

The ovary is ellipsoidal and about half as long as a testis. It is

situated near the middle of the body, somewhat toward the left

side. The vitelline glands are small, irregular in form but gen¬

erally spherical; and distributed along each side of the body from

a little posterior to the acetabulum to the posterior testis. The

uterus is coiled from the genital pore to the posterior end of

the body. It usually contains a couple of hundred eggs which are

elliptical, without a distinct cap, and measure .04 mm. in length.

Eleven specimens, including the type, were taken from the in¬

testine of a short-nosed gar on July 11, 1920. Four other speci¬

mens were found in bullheads caught in the slews at the outlet of

Lake Pepin : three in an Ameiurus natalis on July 6, 1920 ; one in

an Ameiurus nehulosus, July 7, 1920.

Crepidostomum illinoiense Faust

(Fig. 7)

Specimens studied: Cat. No. 7626, U. S. National Museum; June

25, 1920; Lake Pepin, Wisconsin; Collector, A. S. Pearse.

150 Wisconsin Academy of Sciences, Arts, and Letters.

Sixteen specimens of this trematode were taken from the intes¬

tine of a mooneye canght in Lake Pepin, June 25, 1920. Two

other mooneyes were captured at the same time. One contained

twenty-one of the parasites; the other, none.

Allocreadium armatum MacCallum

(Fig. 8)

Length of a large contracted specimen, 3.2 mm. (U. S. Nat.

Mus., Cat. No. 7620); width, 1.08 mm.; diameter of oral sucker,

.35 mm.; diameter of acetabulum, .6 mm.; egg, .11 by .07 mm.

Length of a smaller expanded specimen, 2.8 mm.; width, .53 mm.

Cuticle minutely denticulate, especially toward the anterior end.

The diameter of the oral sucker is nearly two-thirds that of

the acetabulum, which is situated at the posterior end of the an¬

terior third of the body. A short prepharynx is present. The

pharynx is about half the diameter of the oral sucker in length, and

it is about as wide as long. The intestinal rami are slender and

reach nearly to the posterior end of the body.

The ovary is about half as long as the anterior testis and oval,

with the long axis extending across the body. It lies on the right

side of the body just behind the acetabulum. The uterus is coiled

between the acetabulum and the anterior testis, lying mostly on

the left side. It may contain as many as eleven eggs. The genital

pore is on the median line anterior to the acetabulum. The vitel¬

line glands surround the intestinal rami in the posterior third of

the body and extend forward outside the rami nearly to the

middle of the acetabulum.

The two testes are somewhat lobate and are slightly elongated;

their long axes lying across the body. They lie close together

immediately posterior to the ovary between the intestinal rami,

one being directly behind the other. The posterior testis is slightly

larger than the anterior. The cirrus pouch is oval, about a third

as wide as long, and is about as long as the diameter of the

acetabulum.

Sixty-five specimens were taken from a sheepshead, Aplodinotus

grunniens Kafinesque, caught in Lake Pepin, Wisconsin, July 9,

1920. Other specimens were taken from the same locality and

species on July 10, 13; from sunfishes Eupomotis gihhosus (Lin¬

naeus) at Sturgeon Bay, Wisconsin, July 19, 23.

Pearse — Parasitic Worms From Wisconsin Fishes. 151

Allocreadium ictaluri, new species

(Fig. 5)

Type: Cat. No. 7621, U. S. National Museum; Lake Pepin, Wis¬

consin; July 3, 1920; collector, A. S. Pearse. Other specimens

were found in the same host at the same locality July 8, 10.

Host: let alums punctatus (Rafinesque), the channel catfish.

Length of body, 5.9 mm. ; width, 1.85 mm. Diameter of aceta¬

bulum, .63 mm. ; oral sucker, .5 mm. ; pharynx, .3 mm. Cirrus sac,

.45 by .25 mm. Egg, .08 by .045 mm.

Cuticula smooth except on the suckers, where it is rough. The

acetabulum is slightly larger than the oral sucker. The latter is

situated on the ventral surface close to the anterior end. The

acetabulum is in the anterior end of the middle third of the body.

A very short and wide prepharynx connects the oral sucker and

the pharynx, which is nearly spherical in form. There is appar¬

ently no esophagus. The intestinal rami are about .37 mm. in

diameter and reach nearly to the posterior end of the body.

The genital aperture is on the median line at a point nearly half

the distance from the acetabulum to the pharynx. The cirrus sac

is broad, ovate, and reaches to the anterior margin of the aceta¬

bulum. The two testes lie in the anterior portion of the posterior

third of the body. They lie on the median line. The posterior

one is somewhat lobate and the anterior one is elongated with its

axis across the body.

The vitelline glands extend from the pharynx to the posterior

end of the body. They are absent for a space of about .25 mm. on

either side of the acetabulum. They stain most heavily near the

lateral margins of the body but a few occur throughout the middle

portion also. The uterus is coiled between the posterior border of

the acetabulum and the anterior testis. It contains about six

hundred eggs. The ovary is spherical and lies about .1 mm. pos¬

terior to the acetabulum.

Allocreadium boleosomi, new species

(Fig. 4)

Type. Cat. No. 7622, U. S. National Museum; Lake Pepin, Wis¬

consin; July 9, 1920; collector, A. S. Pearse.

Host: the Johnny darter; Boleosoma nigrum; intestine.

152 Wisconsin Academy of Sciences y Arts, and Letters.

Length of body, 1.33 mm.; width, .37 mm.; diameter of oral

sucker, .14 mm. ; diameter of acetabulum, .22. Eggs, .04 by .16 mm.

Cuticula minutely denticulate.

Diameter of oral sucker slightly more than half that of ace¬

tabulum. No prepharynx. Pharynx, a little more than half the

diameter of the oral sucker in length, spindle-shaped, about two-

thirds as wide as long. Esophagus very slender, about two-thirds

as long as the pharynx. Intestinal rami, slender, extending nearly

to posterior end of body.

Genital pore, median, a short distance anterior to the acetabulum.

Ovary, ellyptical with the long axis across the body ; .09 by .175 mm.

Uterus between the ovary and the acetabulum, more on the right

than the left side, containing as many as fourteen eggs. Vitelline

glands forming large rounded follicles, about .05 mm. in diameter,

which are distributed along margins of the body behind the

acetabulum and across the posterior end.

The testes are elongated, median, and immediately behind the

ovary. The anterior one measures .09 by .17 mm. and its long

axis extends across the body. The posterior testis measures .13

by .21 mm. and its long axis is nearly longitudinal. The cirrus

pouch is pyriform; length, .15 mm. It extends back to a little

beyond the center of the acetabulum.

The type was one of seven specimens taken from the intestines

of four Johnny darters collected at the lower end of Lake Pepin,

July 9, 1920. Three other specimens were found in the intestine

of a log perch, Percina caprodes (Rafinesque), collected at the same

place July 5, 1920.

Phyllodistomum superhum Stafford

(Fig. 2)

The writer is convinced that Stafford (1904) confused two

species of the genus Phyllodistomum. Stafford’s name is therefore

retained for the species which most nearly fits his description and

a new name, P. staff ordi, is given to the other.

Length of body, 2.57 mm.; width, 1.5 mm. Diameter of ace¬

tabulum, .29 mm.; of oral sucker, .24 mm. Length of anterior

testis, .38 mm.; of posterior testis, :52 mm.; of ovary, .22 mm.

Size of egg, .022 by .033 mm. (Cat. No. 7623, U. S. Nat. Museum.)

Anterior end of body narrow (.33 mm. wide) as far back as

acetabulum, which is about 1.2 mm. from the anterior end. ^ Ex-

Pearse — Parasitic Worms From Wisconsin Fishes. 153

panded posterior portion is crinkled on margins. Posterior end,

notched.

The esophagus is slender, and about .35 mm. in length; the

intestinal rami are slightly lobate and reach to or a little beyond

the posterior end of the posterior testis ; diameter, about .15 mm.

The genital pore is about midway between the anterior margin

of the acetabulum and the junction of the intestinal rami. The

testes are lobate. They are of about the same width but the pos¬

terior one is longer than the anterior. Though the testes lie in

about the same relative positions as in P. fausti, they are never in

contact, but are separated by a distance of about .1 mm. The

space between them is occupied by about two loops of the uterus.

The ovary is elongated and irregularly lobate. It lies on the

right side, about .1 mm. posterior to the vitelline gland. The two

vitelline glands, though elongated and pyriform, are rather irregu¬

lar in outline. The uterus is filled with eggs and clearly arranged

in coils. It is coiled throughout the posterior portion of the body

posterior to the acetabulum. A few loops extend forward between

the vitelline, glands to about the middle of the acetabulum.

This trematode is not uncommon in the urinary bladder of the

yellow perch, Perea flavescens Mitchill, from two Wisconsin lakes —

Green Lake and Lake Michigan. Stafford (1904) apparently found

this species at Montreal, Canada. The specimens from which the

writer’s description was made were collected at Sturgeon Bay,

July 30, 1921.

Phyllodistomum fausti, new species

(Fig. 1)

Type: Cat. No. 7625, U. S. National Museum; Lake Pepin, Wis¬

consin; June 26, 1920; collector, A. S. Pearse. Fifteen specimens

were taken from a single sheepshead.

Host: Aplodinotns grunniens Rafinesque; urinary bladder.

Length, 4.77 mm.; width, 2 mm.; diameter of oral sucker,

.45 mm.; diameter of acetabulum, .7 mm.; egg, .037 by .028 mm.

Body very flat, widest at junction of middle and posterior thirds ;

anterior third narrow, tapering toward anterior end. Middle por¬

tions of lateral margins crinkled.

Esophagus, .7 mm. in length; about .05 mm. in diameter. In¬

testinal rami about .1 mm. in diameter; reaching to the posterior

fifth of the body.

154 Wisconsin Academy of Sciences, Arts, and Letters.

Ovary .42 by .32 mm., lobate, on left side of body. Vitelline

glands, ovate, slightly lobate, .25 by .13 mm. Uterus, slender,

coiled throughout the body posterior to the acetabulum and on

each side of the acetabulum through about half its length; filled

with eggs ; the coiled terminal duct passes dorsal to the acetabulum

to the genital pore, which lies about midway between the anterior

margin of the acetabulum and the junction of the intestinal rami.

The two testes lie between the intestinal rami, posterior to the

vitelline glands, usually in contact with each other ; the anterior one

is on the right side, the posterior one is in the middle of the body ;

both have deeply incised lobes; the right one measures .35 by

.65 mm., the median one .45 by .8 mm. Seminal vesicle about .3 mm.

long and .17 wide.

Phyllodistomum Stafford!, new species

(Pig. 3)

Type: Cat. No. 7624, U. S. National Museum; Lake Michigan,

Sturgeon Bay, Wisconsin; July 21, 1920. Other specimens were

found in Ameinrus melas (Rafinesque).

Host: the speckled bullhead, Ameiurus nedulosus (Le Seur) ;

urinary bladder.

Length of body, 4.1 mm. ; width, 2.98 mm. Diameter of oral

sucker, .38 mm. ; acetabulum, .52 mm. Egg, .027 by .021 mm.

Anterior quarter of body narrow, flattened, sharply set off from the

posterior portion by a groove. Behind the middle of the acetabu¬

lum the body is discoidal, being nearly circular in outline; the

margins are thin, but not crinkled, and are covered with minute

papillae. The esophagus is short — a little less than the diameter

of the oral sucker in length. The intestinal rami are quite wide

(about .3 mm.) and extend to the posterior eighth of the body.

The genital pore is nearer the anterior margin of the acetabulum

than the junction of the intestinal rami, and is median. The testes

are nearly equal in size, the left being a little larger, irregularly

lobate, and lie on either side of the body; latero-posterior to the

vitelline glands. The left testis is a little more posterior than the

right, and the ovary lies between it and the nearest vitelline gland.

The testes are small, measuring about .42 mm. in diameter. There

is a seminal vesicle, about .5 by .7 mm.

The ovary is elongated, lobate, and measures about .25 mm. in

length. The two vitelline glands are each composed of about eight

Pearse — Parasitic Worms From Wisconsin Fishes. 155

oval lobules. They are .15 mm. apart and about the same distance

from the posterior margin of the acetabulum. The uterus is coiled

between and posterior to the testes, largely between the intestinal

rami. It does not come close to the lateral or posterior margins;

its coils do not lie very close together and do not contain a con¬

tinuous series of eggs, but discontinuous groups.

In writing descriptions of the three species of Phyllodistomum,

the writer has examined specimens as follows: the first figure in¬

dicating the number of phyllodistomes and the second the number

of fishes from which they came : P. fausti, 14-1 ; P. superhum,

11-7 ; P. staffordi, 18-5. The following key separates the three

species :

Key to Species of Phyllodistomum in United States Fishes

1 (2) Anterior, narrow, portion of body separated from discoidal portion

by a distinct groove; testes not extending into posterior quarter of

of body; esophagus not longer than oral sucker; uterus containing

scattered groups of eggs, not reaching near margins of body ;

vitelline glands lobate . . . P. staffordi

2 (1) Anterior, narrow, portion not separated from the posterior portion

by a groove; posterior portion not discoidal, crinkled more or less

along margins; esophagus longer than oral sucker; eggs close to¬

gether in uterus; vitelline glands pyriform . 3

3 (4) Post-acetabular portion of body widest just behind the acetabulum;

testes somewhat separated; posterior margin of body with a distinct

median notch . P. superhum

4 (3) Post-acetabular portion of body widest in middle; testes close to¬

gether; posterior margin of body without a distinct notch . P. fausti

NEMATODA

Ascaris lucii, new species

(Fig. 12)

Type: Cat. No. 7613, U. S. National Museum; Lake Michigan,

Sturgeon Bay, Wisconsin; July 23, 1920. Other specimens were

collected from the same host at the same place on July 20, 21, 1920.

Host : the pickerel, Esox lucius Linnaeus ; intestine.

No males of this species were obtained, but eleven females were

examined.

156 Wisconsin Academy of Sciences^ Arts^ and Letters,

Length of body^ 36 mm.; diameter, 1.1 mm. Lip lobes: length,

.1 mm.; width, .1 mm.; with thin, auriculate, proximal angles;

interlabia absent. The cuticula bears about 48 annulations per

millimeter. The anus is about 1.8 mm. from the posterior end of

the body.

Ascaris scaphirh3mclii, new species

(Fig 11)

Type : Cat. No. 7611, 7612, U. S. National Museum ; Lake Pepin,

Wisconsin; June 20, 1920; collector, A. S. Pearse. Another speci¬

men was taken from the stomach of another sturgeon on the same

date. On June 23 six specimens were taken from the stomach of

a sand sturgeon.

Host : the sand sturgeon, Scaphirhynchus platorhynchus

(Rafinesque) ; intestine.

Length of females, 41 mm.; width, 1 mm.; length of male, 33;

width, .9 mm, Cuticula with about 105 annulations per millimeter.

Anus of female .4 mm. from posterior end; of male, .2 mm.

Lips, .17 mm. long and .15 mm. wide; rounded at tip; base re¬

curved toward tip so that lateral angles are auriculate; two small

papillae on either side of inner surface near distal end. Inter¬

labia conical, rounded at tip, .05 mm. in length.

Posterior end of female, conical, rounded at tip; at .08 mm.

from the end it tapers at a more acute angle and beyond this point

is somewhat bent ventrally. Spicules of male .025 in diameter and

,3 mm. long; somewhat curved, obliquely truncate at tip.

Cystidicola serrata (Wright)

(Fig. 13)

Sixteen specimens of this species were taken from the intestine

of a sheepshead, Aplodinotus grunniens Rafinesque, caught in Lake

Pepin, Wisconsin; July 13, 1920.

The largest female (Cat. No. 7609, U.S.Nat.Museum) is 12.25 mm,

long, and .098 mm. wide; male, 7.53 mm.; .051 mm. In the fe¬

male the anterior portion of pharynx is .4 mm. long; posterior

portion, 2 mm. Ulva, 6.1 mm. from anterior end; anus, .2 mm.

from posterior end. There are ten conical teeth surrounding the

Tear 86 — Parasitic Worms From Wisconsin Fishes. 157

mouth, and inside it. The posterior end of the male is coiled about

three times. There is one short conical spicule, .025 by .015 mm.,

and one long signate spicule, .3 by .02 mm.

Capillaria catostomi, new species

(Fig. 10)

Type: Cat. No. 7627, U. S. National Museum; Lake Michigan;

Sturgeon Bay, Wisconsin ; August 2, 1920 ; collector, A. S. Pearse.

One female was examined and males are unknown.

Host: the common sucker, Catostomus commersonii (Lacepede) ;

intestine.

Length of body, 8 mm.; width, .06 mm.; ulva, 2.55 mm. from

anterior end ; eggs .055 by .025 mm.

The cuticle is smooth. There are one hundred eighty-seven cells

in the esophagus and about seventy eggs in the uterus. The eggs

are typically trichurid; lemon-shaped, with a plug at each end.

Near the ulva they are arranged in a single row, but are more

crowded together further back.

Bibliography

Cooper, A. H. 1915. Trematodes from marine and fresh-water fishes, in¬

cluding one species of extoparasitic turbellarian. Trans. Roy. Soc.

Canada, (3) 9: 181-206.

Faust, E. C. 1918. Studies on American Stephanophialinae. Trans. Amer.

Microscopical Soc. 37: 183-198.

GrOldberger, J. 1911. Some known and three new endoparasitic trematodes

from American fresh-water fish. U. S. Publ. Health and Mar. Hospt.

Serv., Hygienic Lab., Bull. 71: 7-35.

MacCallum, W. G-. 1915. On the anatomy of two distome parasites of

fresh- water fish. Veterinary Mag. 2: 1-12.

Marshall, W. S. and Gilbert, N. 0. 1905. Three new trematodes found

principally in black bass. Zool. Jahrb. Syst. 22: 479-488.

Osborn, H. L. 1903. Bunodera cornuta sp. nov.: a new parasite from the

crayfish and certain fishes of Lake Chautauqua, N. Y. Biol. Bull. 5;

63-73.

Stafford, J. 1900. Some undescribed trematodes. Zool. Jahrb. Syst. 13:

399-414.

- 1904. Trematodes from Canadian fishes. Zool. Anzeig. 27:

481-495.

Wallin, I. E. 1909. A new species of the trematode genus Allocreadium.

Trans. Amer. Mier. Soc. 29: 50-66.

Ward, H. B. and Magath, T. B. 1916. Notes on some nematodes from

fresh-water fishes. J. Parasitol. 3: 57-64.

Ward, H. B. and Whipple, G. C. 1918. Fresh-water biology. New York,

x-1-1111.

158 Wisconsin Academy of Sciences, Arts, and Letters.

TRANS. WIS. ACAD., VOL. XXI PLATE I

1. Phyllodistomum fausti new species.

2. Phyllodistomum superhum Stafford. ,

3. Phyllodistomum staffordi new species.

3 a. Phyllodistomum staff or di; anterior end showing oral sucker, esophagus,

and intestinal rami.

4. Allocreadium holeosomi new species.

5. Allocreadium ictaluri new species.

6. Acetodextra amiuri (Stafford).

Pearse — Parasitic Worms From Wisconsin Fishes. 159

TRANS. WIS. ACAD., VOL. XXI PLATE 7.1

7. Crepidostomum illinoiense Faust.

8. Allocreadium urmatum (MacCallum).

9. Macroderoides spiniferus new species.

10. Capillaria catostomi new species; portion of body showing esophagus,

intestine, ulva, and uterus.

10a. Capillaria catostomi new species; egg.

11. Ascaris scaphirhynchi new species, anterior end.

160 Wisconsin Academy of Sciences , Arts, and Letters.

TEANS. WIS. ACAD., VOL. XXI PLATE IH

12. Ascaris lucii new species; anterior end.

12a. Ascaris lucii new species; posterior end.

13. Cystidicola serrata (Wright) ; anterior end.

13a. Cystidicola serrata (Wright) ; posterior end of male.

13b. Cystidicola serrata (Wright) ; posterier end of female.

13c. Cystidicola serrata (Wright) ; egg.

THE PARASITES OF LAKE FISHES

A. S. Pearse

Introduction

Since the times of Van Beneden and Leuckilart animal parasites

have been intensively studied, but little effort has been made to

determine the amount or frequency of parasitic infection under

natural conditions; except for those species directly related to

man and his domestic animals. Many species of parasites that in¬

fest fishes have been, and are being, described, but few accurate

observations that relate to their abundance and the factors which

make them numerous or few have been made.

Van Cleave (1919) found that half the species of fishes that he

examined from Douglas Lake, Michigan, were infected with acan-

thocephalans, and he determined the percentage of infection for

sixteen species. La Rue (1914), Marshall and Gilbert (1905),

Smallwood (1914), and Ward (1910) made incidental observations

concerning the number of parasites present in certain fishes.

Surber (1913) remarks on the small percentage of natural infec¬

tions with glochidia. The white crappie, which carries more species

of glochidia than any other fish, he found to show an infection of

only 0.7 per cent, and the sheepshead, known to carry two species

of glochidia, had 3.7 per cent.

Zschokke (1902) found that salmon lost a large number of their

parasites while migrating up the Rhine. But Ward (1909) points

out that such migrations are not always conducive to parasitic

losses, for the Alaskan salmon during its journey inland acquires

a copepod which is never found in salt water.

Little is known of the effects of seasonal succession on the life

cycles of the parasites of fishes. Van Cleve (1916) states that

acanthocephalans vary greatly in this respect and cites two species

in one genus which, though occurring in the same host, mature at

different seasons. Hausmann (1897) found that perch had very

few trematodes in the spring. In studying frogs. Ward (1909)

found the lowest percentage of infection in late spring or early

162 Wisconsin Academy of Sciences, Arts, and Letters.

summer, and a maximum was reached during hibernation. There

is a great need of more information concerning the seasonal prev¬

alence of the parasites of all aquatic animals.

Of the factors that control the occurrence of fish parasites there

is also a dearth of knowledge. Hausmann (1897) states that when

fishes eat little on account of cold or heat, parasites are few; and

he assigns an important role to temperature as a factor in parasitic

infection. He also points out that most parasites enter fishes with

food. Ward (1909) stresses the fact that parasites respond to

changes in the habits of their hosts to such a degree that their

presence or absence furnishes evidence of particular habits. Pratt

(1919) affirms that epidemics of fish parasites are apt to occur

when the water is warm and that small inclosed bodies of water

harbor more parasites than those of larger size because fishes can¬

not escape by migration.

The present paper describes the results of statistical studies on

the occurrence of fish parasites in different types of lakes. The

writer w^as led to make such studies in attempting to discover why

fishes fail to grow much in certain bodies of water while they may

attain large size in other bodies near by. It seemed desirable to

learn whether particular lakes showed specificities in regard to the

numbers and kinds of parasites present and whether there is cor¬

relation between the presence and size of particular fishes and

the presence or absence of parasites. The work began in 1917 and

was at first confined to the yellow perch. Observations were made

on specimens from sixteen lakes on three different river systems.

Later, more extensive observations were made on five different

types of lakes where the parasites of all available species of fishes

were studied.

In studying fishes for parasites they were always examined

while fresh, as it was found that results from old or preserved

fishes were of little value. The skin, fins, mouth and gills were

first scrutinized ; then the specimen was opened from vent to throat,

and the visceral organs were examined. The contents of the ali¬

mentary canal were stripped out on a glass plate and the canal

itself was opened from end to end with scissors. The food and

faecal matter were carefully teased across under a binocular micro¬

scope. The number and location of all parasites was entered on a

form sheet, one sheet being used for each fish examined. Parasites

were placed in corrosive sublimate solution and alcohol. Later

they were stained and mounted. In this paper all measurements

Pearse — The Parasites of Lake Fishes,

163

of fishes are given in millimeters and do not include the tail fin.

In the tables indicates less than 0.05.

The work could never have been completed without the cheerful

and excellent assistance rendered by Drs. George R. LaRue and

H. J. Van Cleave, who identified Proteocephalidae and Acanthoce-

phala, respectively. Others who deserve thanks for identifications

are Dr. A. R. Cooper, tapeworms; Prof. H. S. Davis, Sporozoa;

Dr. A. D. Howard, glochidia ; Prof. J. P. Moore, leeches ; and Prof.

C. B. Wilson, copepods. Mr. Leslie Tasche assisted the writer

in the field; Misses Henrietta Achtenberg and Marion E. Lamont,

and Mr. J. C. Stucki mounted slides.

Before all the factors which influence parasitism in fishes are

known, if they ever are, parasitologists and ecologists will have to

labor for several generations. This paper is, of course, only a

beginning in the ecology of fish parasites and is concerned par¬

ticularly with certain Wisconsin lakes.

PARASITES OP THE YELLOW PERCH

From June, 1917, to May, 1918, the yellow perch from the deep

waters of Lake Mendota were examined each month, except dur¬

ing December. During the summer and at intervals throughout

the year perch from four other lakes on the Yahara River and

from the shallow waters of Lake Mendota were also examined.

The results of these studies are given in table 1. The parasites

found in the perch in these lakes were as follows:

Trematoda

Diplostomum cuticola Van Nordmann

This trematode was common, occurring in small cysts in the

skin and fins, and sometimes on the gills. The cysts are easily

seen on account of the black pigment that surrounds them and

gives infected fishes a speckled appearance. Their distribution

on the bodies of 335 perch from all the lakes studied averaged as

follows : pectoral fins 1 ; pelvic fins 0.'9 ; anal fin 0.5 ; caudal fin 1.3 ;

dorsal fins 1; anterior dorsal 0.4; posterior dorsal 0.5; gills +>

opercles 1.3; ventrum 1.4; tail +; dorsum 2.4; sides 3.3.

164 Wisconsin Academy of Sciences, Arts, and Letters,

Bunodera Inciopercae 0. F. Muller

A parasite that was found only in the intestine and the in¬

testinal caeca.

Clinostomum sp.?

Large white cysts enclosing this parasite occurred in the flesh

just beneath the skin ; sometimes on the gills and in the eye sockets.

Acanthocephala

Echinorhynchus thecatus Linton

This hook-headed worm occurred as an intestinal parasite and

in cysts in the peritoneum.

NeoecJiinorJiynchus cylindratus (Yan Cleave)

An intestinal parasite.

Acanthocephalan cysts

These cysts were mostly those of Echinorhynchus thecatus Lin¬

ton, but as it is not certain that they all belonged to that species,

they are not definitely assigned to it.

Nematoidea

Bacnitoides cotylophora Ward and Magath

This nematode was an intestinal parasite in the perch.

Icthyonema cylindraceum Ward and Magath

A filarial worm that occurred in cysts in the peritoneum and

liver.

Cestoidea

Proteocephalus sp.?

Most of the intestinal proteocephalids were P. pearsei La Rue.

Perhaps all belonged to this species.

Proteocephalid cysts

Encysted proteocephalids were commonly encountered; usually

in the liver, but often in the peritoneum. Those identified were

the cysts of Proteocephalus amhloplitis (Leidy).

Pearse — The Parasites of Lake Fishes,

165

Bothriocephalus cuspidatus Cooper

This tapeworm was found in the intestine of the perch.

Glochidia

Probably all the glochidia observed were Lampsilis luteola

Lamark, which is the most abundant species in the lakes along

the Yahara River.

Piscicolaria sp. ?

This leech was usually attached about the bases of the fins.

Unknown cysts

Very often cysts were encountered among the viscera that could

not be identified.

The four lakes along the Yahara River decrease in size and

depth downstream and come thus in the following order: Men-

dota, Monana, Waubesa, Kegonsa (Table 3). Lake Wingra is

small, shallow, and is connected with Lake Monona through a nar¬

row, swampy stream. In general the perch increase in size down¬

stream, but average smaller in Lake Wingra than in Lake Men-

dota (Pearse and Achtenberg, 1920).

The results from the five Yahara lakes may properly be com¬

pared for the months of July, August, and September, when col¬

lections were made in all of them. During that time the total av¬

erage parasitism* for each lake was: Mendota, deep 66.7 (60.7) ;

Mendota, shallow 120.9 (67.7) ; Monona 84.8 (18.8) ; Wingra 38.8

(38.6) ; Waubesa 24.9 (16.4) ; Kegonsa 40.9 (13.4). The figures

in parenthesis represent the total average infection without the

averages for Diplostomum cuticola^ which, when it occurs in very

large numbers on even one fish, may change the total very much.

The figures in parenthesis are probably better as a basis for com¬

parison. In any case it is apparent that a decrease in infection

is generally correlated with larger size in these lakes. Before the

writer had studied lakes along other rivers he thought that a

greater infection might occur in deeper lakes, or in those nearer

the headwaters of a stream.

The different kinds of parasites varied greatly in abundance

in the lakes. Table 1 shows that the perch in the shallow water

*Average total number of parasites per fish.

166 Wisconsin Academy of Sciences, Arts, and Letters.

of Lake Mendota have more parasites than those at greater depths

and that Lake Mendota has a higher percentage of infection than

any of the other lakes. If the highest percentage of infection with

a particular parasite makes a lake ‘‘first/’ the lakes rank about

as follows:

Mendota, shallow: 6 firsts; 6 seconds; 2 thirds; all species pres¬

ent.

Mendota, deep : 5 firsts ; 1 second ; 2 thirds ; 4 species absent.

Wingra: 3 firsts; 1 third; 1 fourth; 1 fifth; 2 sixths; 6 species

absent.

Waubesa: 1 first; 2 seconds; 3 thirds; 2 fourths; 2 fifths; 4 spe¬

cies absent.

Kegonsa: 1 first; 2 seconds; 2 thirds; 1 fourth; 3 fifths; 3 spe¬

cies absent.

Monona: 1 first; 2 seconds; 1 third; 1 fourth; 2 sixths; 6 spe¬

cies absent.

That Monona ranks last is probably explained by the fact that

it is heavily contaminated with organic matter from the city of

Madison. It is on the whole the most barren of the lakes and

contains fewest fishes.

Glochidia, leeches, intestinal proteocephalids, and acanthoceph-

alans are apparently more abundant in shallow water. All en¬

cysted parasites are somewhat more abundant in deep water. Lake

Kegonsa has a much higher infection with bothriocephalids than

any other lake, and also has many Dacnitoides.

The data in table 1 are rearranged in table 3 to show the av¬

erage infection of the perch in the Yahara lakes by months. It

will be noted that the maximum infections were in spring. This

was due largely to the increase of acanthocephalans at that season.

Winter came next, autumn was third, and summer showed the

lowest infection. The different parasites reached their seasons

of greatest numbers as follows :

Spring: acanthocephalans; Bunodera.

Summer : glochidia — ^which were found at no other season.

Autumn: bothriocephalids, Diplostomum, Clinostomum, Icthy-

onema, and proteoeephalid cysts.

Pearse — The Parasites of Lake Fishes.

167

Winter: Dacnitoides; intestinal proteocephalids ; unknown vis¬

ceral cysts.

In order to compare other lakes with deep, fertile lakes along

the Yahara, eleven lakes on the Oconomowoc and Fox Kivers were

visited during August, 1917. The general characteristics of these

lakes are given in table 3. Those on the Oconomowoc were deep,

with sandy and pebbly shores. Those on the Fox (except Green

Lake, which was much like those on the Oconomowoc River) were

shallow with swampy shores.

Table 3 shows that the average infection was about equal on

the Oconomowoc (15.2) and Fox (14.4) Rivers; the latter perhaps

having a slight excess because eight parasites showed the highest

infection average on it — to five on the former. The Yahara lakes

showed about two-thirds the infection (9.3) of those on the othei

two rivers, and the infection average of only two parasites ex¬

ceeded those of the same parasites in the lakes on the other rivers.

In table 4 the lakes of each river system are arranged in order,

with that nearest the headwaters on the left. It will be seen

that only on the Yahara was infection greater toward the head¬

waters.

During August infection with all trematodes, intestinal pro¬

teocephalids, glochidia, and leeches was greater in the Fox River

lakes; in the Oconomowoc lakes acanthocephalans, Icthyonema

cysts, proteocephalid cysts, and unknown cysts were most abund¬

ant. The Yahara lakes showed the highest infections with acan-

thocephalan cysts and Bothriocephalus.

Except for Diplostomum, the trematodes were most abundant in

the shallow lakes. Acanthocephala occurred in largest numbers in

the deep, sandy, and rather barren Oconomowoc lakes ; but acantho-

cephalan cysts reached their greatest numbers in the deep, fertile

Yahara lakes. Dacnitoides was somewhat more abundant in the

shallow lakes than in the sandy, barren, deep lakes and was absent

from the deep, fertile lakes. Icthyonema cysts reached their maxi¬

mum in Lake Poygan (shallow, swampy) but on the whole were

slightly more abundant in the deep, sandy, Oconomowoc lakes.

Intestinal proteocephalids (P. pearsei La Rue) also were prevalent

in the shallow, swampy lakes, but encysted proteocephalids, mostly

P. amhloplitis (Leidy), reached their maximum in the sandy, deep

lakes. Bothriocephalids were most abundant in the deep fertile

lakes. Glochidia and leeches reached their maxima in the shallow.

168 Wisconsin Academy of Sciences, Arts, and Letters,

swampy lakes. Unknown visceral cysts were most common in the

sandy, deep lakes.

According to their degree of average infection the sixteen lakes

rank in the following order: Green 59.1; Monona 42.1; Beaver

32.7 ; Poygan 32.2 ; Mendota shallow 31.1 ; North 30 ; la Belle 30 ;

Oconomowoc 25.5 ; Butte des Morts 19.2 ; Pine 17 ; Mendota deep

15.3; Okauchee 14.5; Puckaway 14.1; Wingra 9.5; Wauhesa 6.5;

Winnebago 3 ; Kegonsa 0.5. If Diplostomum cysts are disregarded

the order is as follows : Beaver 32.1 ; Poygan 30.7 ; North 26 ; Men¬

dota, shallow 23.3 ; Butte des Morts 19.1 ; Mendota, deep 14.7 ;

Oconomowoc 14.2; Puckaway 14; Pine 10.8; Wingra 9.4; Monona

7.1; Green 7.1; la Belle 6.5; Okauchee 2.9; Wauhesa 2.5; Winne¬

bago 1 ; Kegonsa 0.2. There is no apparent relation between river

systems, depth, fertility, character of shore and the general abund¬

ance of parasites. This appears to indicate that all parasites are

not influenced by the same factors. There is also no apparent rela¬

tion between the size of the perch in various lakes and the degree

of parasitic infection, but when there are more fishes per unit of

area there is a heavier infection.

Phyllodistomum superhum Stafford was found in the urinary

bladders of many perch in the lakes along the Fox Kiver, but

was never observed in any of the two thousand perch that the

writer has examined in the lakes on the Mississippi drainage. This

parasite is apparently confined to the St. Lawrence drainage system.

The food of the perch examined was carefully recorded during

August, 1917. For the three river systems it may be summarized

as follows; the figures after the foods indicating the average per¬

centage eaten:

Number examined .

Average length .

Insect eggs .

Fish remains .

Chironomid larvae .

Caddis-fly larvae .

Unidentified insect larvae

Chironomid pupae .

Corixa .

Orasshoppers .

Sialis larvae .

Mites . . .

Crayfishes .

Amphipods, unidentified . .

fishes, snails, and leeches is associated with parasitic infection

more than chironomid larvae and cladocerans, but the observations

cover too limited a period to be of much significance.

COMPARISON OF THE FISH PARASITES OP FIVE

WISCONSIN LAKES

As the study of the yellow perch in sixteen lakes had not given

results of particular value, it was decided to study the parasites

of all the fishes available in several different types of lakes. Ac¬

cording to this plan five Wisconsin lakes were studied intensively

during the summers of 1919 and 1920. Accounts of the food and

distribution of the fishes in these lakes have already been pub¬

lished (Pearse, 1921, 1921a). All the lakes were of considerable

size and depth. Their general characteristics are given in table 5.

Lake Geneva is deep, clear, and its deepest parts are without

oxygen in summer. Mendota is deep, turbid and the water below

8 to 12 meters is without oxygen for three months during summer

and early autumn. Pepin is the shallowest of the lakes, but it has

the greatest area of any lake except Michigan. Its temperature is

nearly uniform from top to bottom and it forms a part of the

Mississippi River. Green Lake is deep and has a small surface

area ; it is sharply stratified thermally but has plenty of oxygen at

the bottom at all seasons. Lake Michigan is clear, cool, and of

course, has a very large volume of water compared to the other

lakes. In regard to “fertility,’^ as judged by the probable amount

of food for fishes per unit of area, the lakes rank in about the

following order : Mendota, Green, Geneva, Michigan, Pepin (Pearse,

170 Wisconsin Academy of Sciences, Arts, and Letters.

1921, pp. 19, 58). Detailed accounts of the general characteristics

of these lakes and of routine catches in them have been published

(Pearse, 1921, 1921a).

Green Lake and Lake Mendota were studied during August,

1919. The lakes investigated in 1920 were studied as follows:

Pepin: June 20 to July 25; Lake Michigan: July 27 to August 7;

Geneva: August 8 to 25.

Attention will now be directed to a detailed consideration of

the parasites that were found to infest Wisconsin lake fishes and

the parts of fishes’ bodies that they frequented. Although un¬

identified parasites of various groups occurred in a number of

fishes, they are not given in the following lists.

Protozoa

Myxobolus cysts were found once in Lake Mendota on the gills

of a yellow perch. Doubtless other protozoan parasites occurred,

but were not observed.

Trematoda

Acetodextra amiuri Pearse

Found in the swim bladders of the yellow, black, and speckled

bullheads in Lake Pepin; in the black and speckled bullheads in

Lake Michigan.

Acrolechanus petalosa (Lander)

Common in the sand sturgeon in Lake Pepin.

Allocanthocasmus varius Van Cleave

Occurred in the white bass in Lake Pepin; free in the intestine

and encysted in the liver.

Allocreadium armatum (MacCallum)

Found in intestine of the sheepshead in Lake Pepin and Lake

Michigan.

Allocreadium holeosomi Pearse

In intestines of the Johnny darter and log-perch in Lake Pepin.

Pearse — The Parasites of Lake Fishes.

171

Allocreadium ictaluri Pearse

In the intestine of the channel catfish in Lake Pepin.

Allocreadium lobatum Wallin

In the intestine of common sucker in Lake Mendota.

Azygia sp. ?

An unidentifiable Azygia was found in the intestine of a yellow

perch in Lake Michigan.

Azygia acuminata Goldberger

This species occurred in the stomachs and intestines of the dog¬

fish, white bass, and wall-eyed pike in Lake Pepin.

Azygia bulhosa Goldberger

In the intestine of the wall-eyed pike in Lake Pepin.

Azygia loosii Marshall & Gilbert

Was found in the stomach and intestine of the rock bass in

Lake Geneva; the pickerel in Lake Mendota; the pickerel and

small-mouth black bass in Lake Pepin ; the pickerel in Green Lake.

Bunodera luciopercae 0. P. Muller

In the intestine and caeca of the yellow perch in Lake Geneva,

Lake Mendota, and Green Lake.

Caecincola parvulus Marshall & Gilbert

In the intestine of the smallmouth black bass in Lake Pepin;

in the rock bass in Lake Michigan.

Centrovarium lobotes MacCallum

In the intestine of the shiner, Notropis hudsonius (De Witt Clin¬

ton), in Lake Michigan.

Clinostomum marginatum Osborn

Found in cuticular cysts in the yellow bullhead in Lake Pepin.

Cysts, probably of this species, were found on a yellow perch from

172 Wisconsin Academy of Sciences, Arts, and Letters.

Lake Michigan. Cnticular cysts were also found in Green Lake on

the speckled bullhead, J ohnny darter, smallmouth black bass, large-

mouth black bass, and yellow perch. In the same lake visceral

cysts were found in the speckled bullhead, pickerel, and yellow

perch.

Crepidostomum sp.?

A single unidentified Crepidostomum was found in the intestine

of a carp from Lake Pepin.

Crepidostomum cornutum (Osborn)

In the intestine of the channel cat in Lake Pepin ; the rock bass,

yellow and speckled bullheads, and one mud puppy in Lake Michi¬

gan.

Crepidostomum illinoiense Faust

In the intestine of the mooneye in Lake Pepin.

Cryptognomius chyli Osborn

Present in the intestine of the rock bass and smallmouth black

bass in Lake Geneva ; rock bass and pumpkinseed in Lake Michigan.

Diplostomum sp. ?

Black cuticular cysts, which probably were D. cuticola van Nord-

mann, were found on the rock bass, pumpkinseed, bluegill, small

and largemouth black bass, and wall-eyed pike in Lake Geneva;

rock bass, Johnny darter, sucker, carp, pickerel, pumpkinseed,

bluegill, both black bass, yellow perch, and wall-eyed pike in Lake

Mendota; black bullhead, carp, pickerel, pumpkinseed, log perch

in Lake Pepin ; rock bass, pickerel, bluegill, smallmouth black bass,

and yellow perch in Green Lake; on the yellow perch in Lake

Michigan.

Visceral Diplostomum cysts commonly occurred in the liver and

peritoneum; sometimes on the heart and in the muscles. In Lake

Geneva they were found in the rock bass, pumpkinseed, bluegill,

both species of black bass, yellow perch; in Lake Pepin in the

pumpkinseed and log perch ; Green Lake : rock bass, pumpkinseed,

top minnow, bluegill, cisco, smallmouth black bass, and blunt-

nosed minnow; Lake Michigan: rock bass, pumpkinseed, and log

perch.

Pearse — The Parasites of Lake Fishes,

173

Gasterostomum pusillum Stafford

In the intestine of the sauger in Lake Pepin.

Leucerthrus micropteri Marshall & Gilbert

In the intestine of the smallmouth black bass in Lake Geneva;

white bass in Lake Mendota and Lake Pepin.

Macroderoides spiniferus Pearse

In the intestine of the yellow and speckled bullheads, and short¬

billed gar in Lake Pepin.

Microphallus opacus Ward

In Lake Mendota this species occurred in the intestines of the

yellow bullhead, dogfish, silversides, smallmouth black bass.

Phyllodistomum fausti Pearse

Found in the urinary bladder of the sheepshead in Lake Pepin.

Phyllodistomum staff ordi Pearse

In the urinary bladder to the black and speckled bullheads, and

the mud cat in Lake Pepin; black and speckled bullheads in Lake

Michigan.

Phyllodistomum superhum Stafford

Found in the intestines of a bream and several perch in Lake

Michigan.

Plagiorchis corti Lament

In the intestine of the tadpole cat in Lake Mendota.

Stephanophiala farionis 0. F. Muller

In the intestine of the Johnny darter and a shiner, Notropis

heterodon (Cope) in Lake Mendota; the yellow perch in Lake

Michigan; the pumpkinseed in Lake Geneva.

174 Wisconsin Academy of Sciences^ Arts, and Letters.

Cestoidea

Ahothrium crassum (Bloch)

In the intestine of the largemouth black bass in Lake Geneva;

lake trout and lota in Lake Michigan.

Bothriocephalus claviceps (Goeze)

In the intestine of the pumpkinseed in Lake Geneva; eel and

pumpkinseed in Lake Pepin.

Bothriocephalus cnspidatus Cooper

In the intestine of the wall-eyed pike in Lake Mendota; sauger

and wall-eyed pike in Lake Pepin.

Corallohothrium sp.f

Perhaps more than one species is represented in the following

notes. Intestinal corallobothria were found in the speckled bull¬

head in Lake Mendota; black, yellow and speckled bullheads,

Johnny darter, channel cat, tadpole cat, and mud cat in Lake

Pepin; speckled bullhead in Green Lake and Lake Michigan.

Encysted corallobothria were found among the viscera of the

Johnny darter in Lake Mendota.

Cyathocephalus americanus Cooper

In the intestine of the cisco in Green Lake and in one of the

ciscoes, Leucichthys hoyi (Gill), in Lake Michigan.

Glariddcris catostomi Cooper

In the intestines of the sucker and pickerel in Lake Geneva;

sucker in Lake Mendota; three species of quillbacks, three species

of buffaloes, the white-nosed sucker, and log perch in Lake Pepin ;

sucker in Green Lake ; sucker, long-nosed sucker, and mud puppy

in Lake Michigan.

Ligula intestinalis Linnaeus

Immature ligulids were found in the peritoneal cavity of the

perch in Green Lake.

Pearse — The Parasites of Lake Fishes.

175

Marsipometra hastata Linton

In the intestine of the spoonbill in Lake Pepin.

Ophiotaenia lonnbergii (Fuhrmann)

In the intestine of the mud puppy in Lake Michigan.

Proteocephalus sp.

Unidentified proteocephalids were found in the intestines of

fishes in Lake Geneva as follows: bluegill and smallmouth black

bass; Lake Pepin: yellow bullhead, smallmouth black bass, two

species of shiners, yellow perch, log perch, white bass, sauger, and

wall-eyed pike; Green Lake: pickerel, bluegill, smallmouth black

bass, yellow perch, and blunt-nosed minnow ; Lake Michigan : rock

bass, sucker, three species of ciscoes, lota, perch, and cottid.

Proteocephalid cysts were found in the liver and peritoneum as

follows : Lake Mendota : yellow perch and white bass ; Lake Pepin :

black and speckled bullheads, eel, sheepshead, channel cat, small¬

mouth black bass, short-nosed red-horse, black and white crappies,

white bass, and wall-eyed pike; Green Lake: rock bass, bluegill,

cisco, small and largemouth black bass, and yellow perch.

Proteocephalus amhloplitis (Leidy)

Intestinal representatives occurred as follows: Lake Geneva:

both species of black bass; Lake Mendota: speckled bullhead,

sucker, pickerel, long-nosed gar, and largemouth black bass; Lake

Pepin : long-nosed gar and white bass ; Green Lake : rock bass,

pickerel, smallmouth black bass.

Visceral cysts were common in the liver and peritoneum. Their

occurrence was as follows: Lake Geneva: rock bass, both black

bass, and yellow perch; Lake Mendota: rock bass, yellow and

black bullheads, sucker, pickerel, top minnow, long-nosed gar,

both black bass, white bass, and wall-eyed pike ; Lake Pepin : black,

yellow and speckled bullheads, sheepshead, sucker, channel cat,

bluegill, mud cat, both black bass, a shiner (Notropis atherinoides

Rafinesque), yellow perch, black and white crappies, white bass,

sauger, and wall-eyed pike; Green Lake: speckled bullhead and

smallmouth black bass ; Lake Michigan : smallmouth black bass and

yellow perch.

176 Wisconsin Academy of ticiences, Arts, and Letters.

Proteocephalus exiguus La Rue

In the intestine of the cisco in Green Lake, and in two of the

ciscoes in Lake Michigan.

Proteocephalus macrocephalus (Creplin)

In the intestine of the eel in Lake Pepin.

Proteocephalus pearsei La Rue

In the intestines of fishes as follows : Lake Geneva : yellow perch ;

Lake Mendota: yellow perch and white bass; Lake Pepin: yellow

perch and log perch. Green Lake: pickerel and yellow perch;

Lake Michigan: yellow perch.

Proteocephalus perplexus La Rue

Occurred in the intestines of fishes. Lake Mendota : yellow bull¬

head, dogfish, and pickerel; Lake Pepin: rock bass; Green Lake:

blunt-nosed minnow.

Visceral cysts were found in the white bass in Lake Mendota;

in the speckled bullhead and blunt-nosed minnow in Lake Pepin.

Proteocephalus pinguis La Rue

Occurred in the intestines of the pickerel in all five of the lakes

studied. Visceral cysts were found in the rock bass in Green Lake.

Proteocephalus singularis La Rue

Pound in the intestine of the long-nosed gar in Lake Mendota

and in the short-nosed gar in Lake Pepin.

Triaenophorus nodulosus Pallas

Encysted visceral representatives of this species were found as

follows : Lake Pepin : speckled bullhead and short-headed redhorse ;

Lake Michigan: sucker, long-nosed sucker, smallmouth black bass,

and yellow perch.

Trypanorhyncha, gen.? sp.?

Visceral cysts occupied by representatives of this order were

found in the yellow bass and skipjack in Lake Pepin; and in the

top minnow in Lake Michigan.

Pearse — The Parasites of Lake Fishes,

177

Nematoda

Ascaris sp.?

An Ascaris that could not be identified was found in the in¬

testine of a rock bass in Lake Geneva.

Ascaris angulata Eudolphi?

An ascarid that perhaps belonged to this species was taken from

the intestine of a pumpkinseed caught in Lake Michigan.

Ascaris lahiata Eudolphi

Found in the intestine of the rock bass in Lake Mendota.

Ascaris lucii Pearse

This species was the commonest intestinal ascarid found. Lake

Geneva : pickerel ; Lake Mendota : rock bass, pickerel ; Green Lake :

pickerel; Lake Michigan: pickerel.

Ascaris scaphrhynchi Pearse

Pound in the intestine of the hackleback, or sand sturgeon, in

Lake Pepin.

Camallanus ancylodirus Ward & Magath

This species was found in the intestine of a lake carp ( Carpoides

thompsoni Agassiz) caught in Lake Pepin.

Camallanus oxycephalus Ward & Magath

This species was found in Lake Pepin only. It was usually

hanging from the intestine out through the anus. The color in

life was bright red. Occurred in the following fishes: mooneye,

mud cat, yellow bass, short-headed redhorse, log perch, black crap-

pie, white bass, wall-eyed pike.

Capillaria catostomi Pearse

This species was found in the intestine of a sucker caught in

Lake Michigan.

178 Wisconsin Academy of Sciences, Arts, and Letters.

Cystidicola serrata (Wright)

Sixteen specimens were found in the intestine of a sheepshead

caught in Lake Pepin.

Cystidicola sigmatura (Leidy)

A single specimen was found in a mooneye caught in Lake

Pepin. This species was abundant in the swim bladders of the

ciscoes in Lake Michigan, occurring in all the species except the

long jaw.

Dacnitoides cotylophora Ward & Magath

This species was often common in the yellow perch, occurring in

the intestine of that species in all of the five lakes studied except

Pepin. It was also found in the intestines of other fishes as follows :

Lake Pepin: channel cat; Green Lake: speckled bullhead; Lake

Michigan: black bullhead, sucker.

Cysts of this species were also found among the viscera of a cisco,

Leucichthys johan7iae (Wagner), in Lake Michigan.

Haplonema immutatum Ward & Magath

Found in the intestine of a dog fish in Lake Pepin.

Histerothylacium hracJiyurum Ward & Magath

Pound in the intestines of the largemouth black bass and yellow

perch in Lake Geneva.

Icthyonema cylindraceum Ward & Magath

Cysts of this species were common in the liver and peritoneum

of many fishes. Occurrence : Lake Geneva : smallmouth black bass,

yellow perch ; Lake Mendota : speckled bullhead, J ohnny darter, top

minnow, cisco, largemouth black bass, shiner — Notropis heterodon

(Cope), yellow perch, white bass; Lake Pepin: black bullhead;

Green Lake: rock bass, top minnow, yellow perch, blunt-nosed

minnow; Lake Michigan: Johnny darter, Iowa darter, pumpkin-

seed, yellow perch, blunt-nosed minnow.

Oxyuris sp.?

A mutilated specimen, apparently belonging to this genus, came

from a blue-gill caught in Lake Pepin.

Pearse — The Parasites of Lake Fishes.

179

Spinitectus gracilis Ward & Magath

This is an intestinal parasite of wide distribution. Occurrence:

Lake Geneva: pumpkinseed, bluegill, smallmouth black bass; Lake

Pepin : bluegill, smallmouth black bass ; Lake Michigan : rock bass,

pumpkinseed.

Acanthocephala

Echinorhynchus coregoni Linkins

In Lake Michigan in the intestines of long-nosed and common

suckers, whitefish, lake trout, pickerel, five species of ciscoes, lota,

yellow perch.

Echinorhynchus salvelini Linkins

In Lake Michigan in intestines of lake trout, four species of

ciscoes, smallmouth black bass, yellow perch.

Echinorhynchus thecatus Linton

This was found to be a common and widely distributed intes¬

tinal parasite. Occurrence: Lake Geneva: rock bass, pickerel,

bluegill, both black bass, yellow perch; Lake Mendota: rock bass,

speckled bullhead, pumpkinseed, long-nosed gar, bluegill, both

black bass, yellow perch ; Lake Pepin : dogfish, eel, J ohnny darter,

pickerel, pumpkinseed, bluegill, both black bass, yellow perch,

white bass, sauger; Green Lake: rock bass, speckled bullhead,

sucker, top minnow, bluegill, both black bass, yellow perch; Lake

Michigan: speckled bullhead, yellow perch.

Larval individuals were found in small white crappies in Lake

Pepin; Green Lake: pickerel, top minnow, bluegill, smallmouth

black bass, yellow perch.

Visceral cysts were found as follows: Lake Geneva: rock bass,

pumpkinseed, both black bass; Lake Mendota: rock bass, speckled

bullhead, pumpkinseed, largemouth black bass ; Lake Pepin : sheeps-

head, pumpkinseed, channel cat, largemouth black bass, tadpole

cat; Green Lake: rock bass, speckled bullhead, pumpkinseed, top

minnow, bluegill, largemouth black bass, yellow perch; Lake

Michigan: speckled bullhead, sucker, yellow perch.

180 Wisconsin Academy of Sciences ^ Arts, and Letters.

Neoechinorhynchus crassus Van Cleave

Found in the intestine of the sucker in Lake Mendota, Lake

Geneva, and Lake Michigan.

Neoechinorhynchus cylindratus (Van Cleave)

In the intestines of fishes as follows : Lake Mendota : largemouth

black bass; Lake Pepin: Johnny darter, pickerel, both black bass,

yellow perch, white bass, sauger, wall-eyed pike; Lake Michigan:

sucker.

Juvenile specimens occurred in the pickerel in Green Lake.

Visceral cysts were found as follows : Lake Mendota : rock bass ;

Lake Pepin: pickerel, smallmouth black bass, white bass, sauger,

wall-eyed pike; Green Lake: pickerel, smallmouth black bass.

Octospinifer macilentus Van Cleave

Occurred in the intestine of the sucker in Lake Geneva and

Lake Michigan.

Pomphorhynchus hulhocolli Linkins

In the intestine of fishes: Lake Geneva: rock bass, sucker, both

black bass; Lake Mendota: sucker, blunt-nosed minnow; Lake

Pepin: sucker, a buffalo — Ictiohus cyprinella (Cuvier & Valencien¬

nes), white-nosed and short-headed suckers, skipjack; Green Lake:

sucker; Lake Michigan: black bullhead, sucker, Iowa darter.

Juvenile individuals were found in Green Lake in the following

fishes: rock bass, speckled bullhead, top minnow, smallmouth

black bass.

Visceral cysts occurred as follows : Lake Geneva : rock bass,

sucker, both black bass ; Lake Mendota : yellow and black bullheads ;

Lake Michigan: speckled bullhead, sucker.

Hirudinea

Unidentified leeches were found as follows : Lake Mendota : rock

bass, speckled bullhead, largemouth black bass ; Lake Pepin : black

and yeUow bullheads; Green Lake: rock bass, speckled bullhead,

top minnow, largemouth black bass, yellow perch ; Lake Michigan :

lake trout, lota.

Pearse — The Parasites of Lake Fishes.

181

Piscicola punctata (Verrill)

Leeches, identified by the writer, occurred as follows: Lake

Mendota: carp, bluegill, largemouth black bass; Green Lake:

pumpkinseed.

Piscicola milneri (Yerrill)

Found in Lake Michigan on a cisco — Leucichthys hoyi (Gill),

and the lota.

Piscicolaria sp. ?

Professor J. P. Moore identified specimens from the following

sources : Lake Geneva : rock bass, yellow perch ; Lake Pepin : black

and yellow bullheads, channel cat; Lake Michigan: rock bass,

smallmouth black bass.

Placohdella montifera Moore

Occurrence : Lake Geneva : smallmouth black bass ; Lake Pepin :

carp and hackleback sturgeon.

Placohdella parasitica (Say)

Occurrence: Lake Mendota: bluegill; Green Lake: pickerel,

top minnow.

Placohdella picta (Verrill)

Occurred in Lake Michigan on the sucker and perch.

Lamellibranchiata

During all the studies described in this paper glochidia were

never found on any part of fishes except the gills. In Lake Men¬

dota the yellow perch was infected with Lampsilis luteola (La-

mark). In Lake Pepin the yellow perch was infected with Lamp¬

silis luteola (Lamark) and Quadrula plicata (Say) ; the sauger,

with Quadrula metaneura (Say) (‘‘probably”)? Lampsilis recta

(Lamark), L. ligamentina (Lamark) ; the mud cat, with unidenti¬

fied glochidia.

182 Wisconsin Academy of Sciences, Arts, and Letters.

COPEPODA

Achtheres ambloplitis Kellicott

In Lake Michigan on the gills of a cisco — ^Leucichthys harengus

(Richardson), the lota, and smallmouth black bass.

Achtheres coregoni (Smith)

On the giUs of the lake trout in Lake Michigan.

Achtheres corpulentus Kellicott

On the gills of a cisco — Leucichthys johannae (Wagner), in Lake

Michigan.

Achtheres micropteri Wright

On the gills of the smallmouth black bass in Lake Geneva.

Achtheres pimelodi Kroyer

On the gills of the channel cat in Lake Pepin.

Argulus sp. ?

An unidentified Argulus was found on a carp in Lake Mendota.

Argulus appendiculosus Wilson

Found in Lake Pepin on the mud cat.

Argulus catostomi Dana and Herrick

Under the opercula and on the gills of suckers in Lake Geneva

and Lake Mendota.

Argulus maculosus Wilson

Op the yellow bullhead in Lake Pepin.

Ergasilus caeruleus Wilson

On the gills of fishes : Lake Mendota : yellow perch ; Lake Michi¬

gan: rock bass, sucker, a cisco — Leucichthys harengus (Richard¬

son), and yellow perch.

Pearse — The Parasites of Lake Fishes.

183

Ergasilus centharchidium Wright

In Lake Michigan on the gills of the rock bass.

Pisces

Icthyomyzon concolor (Kirtland)

This lamprey was taken in Lake Pepin on a spoonbill.

Discussion and Conclusions

A summary of the parasites found in the five lakes studied is

given in table 4. The writer had expected to publish six additional

tables giving the number of fishes infected and total average in¬

fection for each parasite but the space available will not permit

this. Those interested in such detailed information may obtain

it by letter. Lake Mendota in all respects contained the smallest

number of parasites — the average number in each fish being 2.0;

the average number of species of fishes each parasite infected, 4.7 ;

and the average number of individual fishes infected by each para¬

site, 1.4. Lake Pepin had the largest average number of parasites

per fish (5.0) ; Green Lake the largest average number of species

of fishes infected by each species of parasite (8.1) ; and Lake Gen¬

eva the largest average number of individuals infected by each

parasite (2.2).

The lakes with the widest range of territory and opportunity

for fishes to invade the greatest variety of habitats have the high¬

est average infection per fish (Pepin, 5.0; Michigan, 3.9). How¬

ever, the fishes in Pepin, with its shifting, sandy bottom and lack

of thermal stratification, have 22 per cent more parasites than

Michigan, with its soft mud bottom and cold deeper water. The

two lakes with the largest average number of species of fishes in¬

fected per species of parasite include the one with a large number

of species of fishes and habitats '(Pepin, 8.1) and the one with the

smallest number of species of fishes and the least variety of habi¬

tats (Geneva, 6.6). The lakes having the largest number of in¬

dividuals infected by each parasite include one with the least range

and variety of habitats (Geneva, 2.2), one with wide range and

variety of habitats {Michigan, 2.1), and one in which there was

little variety in the shore habitats and a sharp distinction between

deep and shallow water habitats (Green, 2.1).

184 Wisconsin Academy of Sciences, Arts, and Letters.

The lake in which there was the least infection by parasites

(Mendota) had the largest number of fishes per unit of area

(Pearse, 1921, p. 24), the most abundant food supply, the greatest

degree of stagnation in the deeper water during summer. Table

4 shows, however, that this lake had the second largest number of

species of parasites which showed the largest number of highest

infection averages; and that it was excelled in this only by Lake

Pepin, which had nearly twice as many species of fishes, the

scantiest food supply, and the greatest parasitic infection. Taking

the number of species of parasites that showed the highest average

infections as a criterion, the lakes rank in the following order:

Pepin, Mendota, Michigan, Green, Geneva. This order indicates

that there is a direct relation between variety of habitat and amount

of infection.

Taking the total number of species of fish parasites present as

a criterion the lakes rank in the following order: Pepin 70, Michi¬

gan 60, Mendota and Green 44, Geneva 35. This indicates that

variety of habitat is correlated with a large number of species of

parasites (and fishes) as well as a large amount of infection. In

other words the lake with the largest variety of habitats has the

greatest variety of fishes and parasites.

Arranged in order of average infection the fishes in each of

the five lakes in which extensive observations were made rank as

follows :

Lake Geneva: Kock bass 9.4, pumpkinseed 5.0, smallmouth

black bass 4.0, sucker 2.2, cisco 1.0, largemouth black bass 0.8, wall¬

eyed pike 0.6, pickerel 0.5, bluegill 0.5, perch 0.5, brook trout 0,

shiner (Notropis hudsonius) 0.

Lake Mendota: White bass 13.2, wall-eyed pike 12.7, dog¬

fish 8.3, smallmouth black bass 6.6, sucker 4,0, long-billed gar 3.8,

tadpole cat 2.8, pumpkinseed 2.6, speckled bullhead 2.4, largemouth

black bass 2.4, Johnny darter 2.0, bluegill 1.6, perch 1.4, yellow

bullhead 1.3, rock bass 1.1, pickerel 1.1, cisco 0.8, black crappie

0.7, top minnow 0.3, shiner (Notropis heterodon) 0.2, silversides

0.1, carp 0.1, blunt-nosed minnow +, bream 0, buffalo 0, miller’s

thumb 0.

Lake Pepin: Mud cat 462, lake carp 116.6, eel 40, dogfish

37, spoonbill 19, smallmouth buffalo 17, yellow bass 16, quillback

13.2, river carp 5, sucker 4.7, black bullhead 3.7, skipjack 3.3,

Pearse — The Parasites of Lake Fishes.

185

pickerel 3.1, speckled bullhead 2.9, white bass 2.9, channel cat

2.7, tadpole cat 2.5, mongrel buffalo 2, hackleback sturgeon 1.8,

sauger 1.7, wall-eyed pike 1.6, pumpkinseed 1.3, white crappie 1,

yellow perch 0.9, shiner (Notropis heterodon) 0.8, short-headed

redhorse 0.7, long-billed gar 0.6, largemouth black bass 0.5, bluegill

0.5, mooneye 0.4, white-nosed sucker 0.4, black crappie 0.4, Johnny

darter 0.2, shiner (Notropis atherinoides ) 0.2, log perch 0.2, carp

0.1, shiners (Notropis jejunus and N. hudsonius) 0.1, pirate perch

+, gizzard shad 0, lamprey 0.

Green Lake : Pumpkinseed 29.7, blunt-nosed minnow 22.5,

sucker 21.4, rock bass 8.9, bluegill 4.2, yellow bullhead 2.5, small-

mouth black bass 2.5, cisco 2.4, carp 2, speckled bullhead 1.9, top

minnow 1.1, largemouth black bass 0.8, yellow perch 0.4, pickerel

0.4, Johnny darter 0.1, shiner (Notropis atherinoides) 0.

Lake Michigan: Whitefish 31.1, smallmouth black bass 28.9,

blackfin 27.8, pumpkinseed 18.9, carp 15, rock bass 12.6, lake

trout 9.2, speckled bullhead 6.5, chub 5.8, mud puppy 5, black bull¬

head 4.1, bloater (Leucichthys harengus) 3.9, lota 3.9, bloater

(Leucichthys hoyi) 2, log perch 1.5, pickerel 1.4, cottid 1.3, sucker

1.2, top minnow 1, Iowa darter 0.9, long-nosed sucker 0.8, Johnny

darter 0.8, yellow perch 0.5, bream shiners (Notropis atherin¬

oides and N. hudsonius) +, blunt-nosed minnow 0, yellow bull¬

head 0.

It would not be proper to add the average infections in different

lakes in order to compare infection in different species of fishes,

for the total infection in different lakes varies greatly. Each spe¬

cies should be given a rating which will compare it with all others

in each lake. The writer has therefore used the following formula

in order to give to each fish a relative percentage (P).

p=100— I^XR

N is the number of fishes examined in the lake ; B, the rank of

the particular species of fish in the lake according to its average

infection. For example twelve species were examined in Lake

Geneva (3^). The largemouth black bass ranks sixth (^^X6).

For this species P=100 — i^X6— 50. Using this method for each

lake and averaging the relative percentages for all species of fishes

that occurred in two or more lakes, the fishes rank in the follow¬

ing order: dogfish 90, smallmouth black bass 78, white bass 78,

186 Wisconsin Academy of Sciences, Arts, and Letters.

rock bass 73, pumpkinseed 71, sucker 68, black bullhead 66, tad¬

pole cat 64, wall-eyed pike 61, speckled bullhead 60, long-billed

gar 56, bluegill 55, yellow bullhead 55, all species of ciscoes 54,

largemouth black bass 45, pickerel 42, carp 40, buffalo (Ictiohus

cyprinella) 36, log perch 35, blunt-nosed minnow 32, top minnow

32, yellow perch 31, shiner (Notropis heterodon) 30, Johnny darter

30, black crappie 27, shiner (Notropis atherinoides) 14, bream 11,

shiner (Notropis hudsonius) 11.

In general the fishes that frequent vegetation show the highest

infection with parasites, those that frequent the bottom and open

water are intermediate, and the small fishes that live in shallow

water have fewest parasites. Doubtless many factors influence

the prevalence of fish parasites, and much is yet to be learned

before general laws that will enable one to predict the degree of

infection that will be probable in a particular locality are for¬

mulated.

There are many ways in which parasites may infect fishes. The

most important means of infection are: (1) food, (2) the active

migration of the parasite to its host, (3) and accidental contami¬

nation from bottom mud, vegetation, or other material. Van

Cleave (1920) and Mrazek (1891) have found parasites encysted

in amphipods. The observations described in this paper show that

the fishes in Lake Michigan that feed largely on amphipods are

heavily infected with the parasites these crustaceans are known

to carry. Several types of parasites are found encysted in fishes

and fish-eaters often have a heavy infection. Hausmann (1897)

believed that fishes acquired parasites chiefly through their food,

and he was doubtless right, as intestinal parasites are most abund¬

ant. In general the fishes that eat the greatest variety of food

have the most parasites, but there are some notable exceptions to

this. The dogfish, wall-eyed pike, and gar, for example, subsist

largely on fishes (Pearse 1918) and are heavily infected.

Little is known of the active migration of fish parasites to their

hosts. The distribution of the cuticular cysts of Diplostomum in¬

dicates that infection may occur in such a way. Faust (1918)

notes that Bunodera has been known to wander out of dead fishes.

Perhaps parasites may infect a second host after leaving the first.

The acquiring of parasites from the accidental ingestion of

eggs or other stages may be characteristic of certain parasites that

are erratic in their occurrence — like intestinal nematodes.

Pearse — The Parasites of Lake Fishes.

187

The susceptibility of the host is an important factor in deter¬

mining the degree and frequency of parasitic infection. Proteo-

cephalus pinguis was never found by the writer except in the pick-

eral ; P. ambloplitis was found in a number of hosts. Fasten (1913)

has studied a parasitic copepod which kills brook trout in great

numbers, but will not live on the German brown trout. Howard

(1914) cites similar instances among glochidia. Pish parasites

may show considerable specificity for certain hosts and hosts may

possess a varying degree of immunity. The black crappie, for

example, appears to be immune to many parasites that attack

other Cenfrarchidae. The largemouth and smallmouth black bass

are closely related, but during the present investigations the latter

always carried more parasites. The pumpkinseed was always

more heavily infected than the bluegill. Perhaps some of these

differences are due to differences in habitats, but some are un¬

doubtedly due to susceptibility. Furthermore, there is general

similarity between the parasites of the Siluridae and certain Per-

cidae which leads one to believe that infection may be limited

at least in part by certain chemical substances which are present

in or absent from the bodies of fishes.

Surber (1913) has remarked on the remarkably small percent¬

age of fishes that carry glochidian. parasites in nature. Van

Cleave (1919) showed that only half the species of fishes in a

Michigan lake were infected with ancanthocephalans. Infection

depends on so many factors and opportunities that it is to be ex¬

pected that parasites will frequently fail to reach their hosts.

Most fish parasites do little harm to their hosts. Pratt (1919)

states that nematodes are most injurious and that trematodes do

the least harm. While his generalizations may apply^ to the para¬

sites of certain marine fishes, like the ,cod, the writer does not be¬

lieve they are generally applicable to the fresh-water fishes in the

United States. The most injurious parasites (leaving out the

Protozoa, which the writer has not studied) appear to be the

larval tapeworms, which destroy liver tissue; aeanthocephalans,

which cause ulcers in the wall of the intestine; and copepods,

which suck blood from the gills. Glochidia, nematodes, and leeches

are usually too few in numbers to do serious damage. Tapeworms

and trematodes often occur in enormous numbers but do not appear

to do much injury. The writer has never examined a dogfish that

did not contain numerous tapeworms; yet all appeared to be in

good condition.

188 Wisconsin Academy of Sciences, Arts, and Letters.

The migrations of fresh-water fishes doubtless afford them op¬

portunity to acquire and shed parasites (Ward 1909) : The fishes

that travel most and invade the greatest variety of habitats in

general have the most parasites.

Seasonal changes doubtless have a marked effect on certain fish

parasites. It was found that perch have most parasites in spring,

although some species of parasites were more abundant at other

seasons. Ward (1909), Marshall and Gilbert (1905) and Van

Cleave (1916) have also made observations on the abundance of

parasites at various seasons, but such information is as yet too

limited for generalization.

Hausmann (1897) thought that perch had few parasites when

little food was eaten on account of low temperature. In the

writer’s experience perch have not been found to refrain from

eating during winter and they have more parasites in winter than

in autumn or summer. Pratt (1909) says that epidemics of

trematodes are likely to occur when the water is warm. The writer

has found no evidence that this is the case, at least in fresh-water

fishes.

Some parasites are apparently limited to particular drainage

systems. Phyllodistomum superhum Stafford was quite common

in the urinary bladder of perch in several lakes on the St. Law¬

rence drainage but was never observed in a single one of the sev¬

eral thousand perch examined from the lakes on the Mississippi

drainage. Perhaps this trematode entered the St. Lawrence drain¬

age after the Mississippi separated from it. Other cases of this

kind probably occur but too few specimens have yet been examined

to demonstrate them.

Contrary to Pratt’s (1919) assertion, the size of a lake does not

appear to be correlated with the degree of parasitic infection of

its fishes. The density of the population may sometimes be of im¬

portance. However, the fishes of Lake Mendota, which has the

densest population of any of the five lakes studied extensively by

the writer, has the fewest parasites and the fishes in the depth of

Lake Michigan, where the population is scanty, have many : There

is, in general, no direct relation between number of fishes and num¬

ber of parasites.

The habitats of fishes are of course important in their relation

to parasitic infection. While a variety of habitats is desirable for

the growth of fishes, it gives opportunity for acquiring more para¬

sites. A wide range also gives more opportunities for acquiring

Pearse — The Parasites of Lake Fishes.

189

parasites than a restricted one. Ward (1910) has pointed out that

“the parasitic fauna of any animal is primarily a function of its

habitat”. The observations reported in this paper indicate that

the fishes in comparatively barren lakes have an unusual number

of visceral cysts and acanthocephalans — perhaps because they

wander about more in search of food. Visceral cysts appear to be

more abundant in deep water fishes than in those from shallow

water. Linton (1910) found that marine fishes in shallow water

had fewer parasites than those at greater depths. He believed

this was because the former had less opportunity to wander about

and become infected. In fresh-water the writer has found infection

to be greater on the whole in shallow water — probably because

there is a greater variety of habitats and secondary hosts there.

The ecological factors that appear to be important in relation to

parasitic infection have now been discussed. Perhaps only one

thing has been made clear — that, “now we see through a glass

darkly”. The writer makes no apology for this final conclusion.

There is great need for more information in regard to the ecology

of parasitism, not only among fishes but among all animals. It is

hoped that this paper may help to interest fishermen, parasitolo¬

gists, geographers, and ecologists in the opportunities that are open

for research on the parasites of fishes. The fisherman must in the

future learn to increase his catch by the control of parasites. There

are many undescribed species and life histories that await discov¬

ery by the parasitologist. The zoogeographer has unusual oppor¬

tunities for study — parasites depend on one or more hosts for their

distribution and .are hence conservative in their migrations. The

ecologist may disclose the most intricate and interesting relations

to be found in nature.

Bibliography

Cooper, A. R. 1917. A morphological study of bothriocephalid cestodes

from fishes. Jour. Parasitol. 4: 33-39. TJrbana.

Fasten, N. 1913. The behavior of a parasitic copepod, Lernaeopoda ed-

wardsii Olsson. Jour. An. Behavior, 3: 36-60. Cambridge.

Faust, E. 0. 1918. Studies on American Stephanophialinae. Trans. Amer.

Micro. Soc. 37: 183-198. Menasha.

Goldberger, J. 1911. Some known and three new endoparasitic trematodes

from American fresh-water fish. Bulletin XT. S. Hygienic Laboratory,

No. 71: 7-35. Washington.

Hausmann, L. 1897. Ueber Trematoden der Susswasserfische. Revue suisse

die Zoologie, 6: 1-42, pi. 1. Geneva.

190 Wisconsin Academy of Sciences, Arts, and Letters.

Howard, A. D. 1914. Some cases of narrowly restricted parasitism among

commercial species of fresh-water mussels. Trans. Amer. Fish. Soc.

1914: 41-44.

La Eue, Gr. R. 1914. A revision of the cestode family Proteoeephalidae.

Bui., Univ. Ill. 12: 1-531. IJrbana.

Linton, E. 1910. Notes on the distribution of entozoa of North American

marine fishes. Proceedings, Seventh International Zoological Congress,

1907: 1-11. Cambridge.

- . 1911. Trematode parasites in the skin and flesh of fish and the

agency of birds in their occurrence. Trans. Amer. Fish. Soc. 1911:

245-259.

Marshall, W. S. and Gilbert, N. C. 1905. Three new trematodes found

principally in black bass. Zoolog. Jahrb. Syst., 22: 447-488.

Mrazek, A. 1891. Pfispevky k vyvojezpytu nekterych tasemnic ptacich.

Zvalstni otisk z Vestnika kralovske ceske spolecnosti nauk, 1891: 97-

131, tab. V. Prag.

Osborn, H. L. 1911. On the distribution and mode of occurrence in the

United States and in Canada of Clinostomum marginatum, a trematode

parasite in fish, frogs. Biolog. Bui. 20: 350-364. Woods Hole.

Pearse, A. S. 1918. The food of the shore fishes of certain Wisconsin lakes.

Bulletin, U. S. Bureau of Fisheries. 35: 247-292. Washington.

- . 1920. The fishes of Lake Valencia, Venezuela. University of Wis¬

consin Studies in Science, No. 1: 1-51. Madison.

- . 1921. The distribution and food of the fishes of three Wisconsin

lakes in summer. Ibid., No. 3: 1-61. Madison.

- . 1921a. Distribution and food of the fishes of Green Lake, Wis¬

consin, in summer. Bulletin, U. S. Bureau of Fisheries, 37: 255-272.

Washington.

Pearse, A. S., and Achtenberg, H. 1920. Habits of yellow perch in Wis¬

consin lakes. Bulletin, U. S. Bureau of Fisheries, 36: 293-366. Wash¬

ington.

Pratt, H. S. 1919. Parasites of fresh-water fishes. Economic Circular, U.

S. Bureau of Fisheries, No. 42: 1-8. Washington.

Smallwood, W. M. 1914. Preliminary report on the diseases of fish in the

Adirondacks, a contribution to the life history of Clinostomum marginatum.

Tech. Pub. N. Y. State Col. For., Syracuse University, No. 1: 1-27.

Syracuse.

Surber, T. 1912. Identification of the glochidia of fresh-water mussels.

U. S. Bureau of Fisheries, Doc. 771: 1-10, pis. 1-3. Washington.

- . 1915. The identification of the glochidia of fresh- water mussels.

Report, U. S. Commissioner of Fisheries, 1914: 1-16. Washington.

Van Cleave, H. J. 1916. Seasonal distribution of some Acanthocephala

from fresh-water hosts. Jour. Parasitol. 2: 106-110. Urbana.

- . 1919. Acanthocephala from fishes of Douglas Lake, Michigan.

Occasional Papers of the Museum of Zoology, University of Michigan,

No. 72: 1-12. Ann Arbor.

- . 1919a. Acanthocephala from the Illinois River, with descriptions

of species and a synopsis of the family Neoechinorhynchidae. Bui.

Ill. Nat. Hist. Sur. 12: 225-271. Urbana.

Pearse — The Parasites of Lake Fishes.

191

- . 1920. Notes on the life cycle of two species of acanthocephalans

from fresh-water fishes. Jour. Parasitol. 6: 167-172. XJrbana.

Ward, H. B. 1909. The influence of hibernation and migration on animal

parasites. Proceedings, Seventh International Zoological Congress,

1907: 1-12. Cambridge.

- . 1910. Internal parasites of the Sebago salmon. Bulletin, U. S.

Bureau of Fisheries, 28: 1151-1194, pi. CXXI. Washington.

Ward, H. B. and Magath, T. B. 1916. Notes on some nematodes from

fresh-water fishes. Jour. Parasitol. 3: 57-64. Urbana.

Ward, H. B. and Whipple, G. 0. 1918. Fresh-water biology, pp. x+llll.

New York.

Wilson, C. B. 1915. North American parasitic copepods belonging to the

Lernaeopodidae, with a revision of the entire family. Proceedings of

the U. S. National Museum, 47: 565-729; pis. 25-56. Washington.

Zschokke, F. 1902. Marine Schmarotzer in Susswasserfischen. Verhandl.

Naturforschen. Gesellsch. in Basel, 16: 118-157.

Table. 1. Total average infection of the yellow perch in the lakes along the

Yahara Biver during July, August, September, 1917.

192 Wisconsin Academy of Sciences, Arts, and Letters.

Table 2. Showing average infection hy months in the Yahara Biver laTces,

1917-1918. After October, perch were not examined from any lake except

Mendota, with the exception of seven specimens from Kegonsa in November.

three drainage systems are summarized.

Pearse — The Parasites of Lake Fishes,

193

o

oi

CO

$5

A

S

I

§“

S>

e

I-

§

o

rs;

CQ

194 Wisconsin Academy of Sciences, Arts, and Letters,

Table 4. Summary of the average infection with fish parasites in five Wis¬

consin laJces during summer. + indicates less than 0.01 per cent.

THE ANATOMY OF TKOCTES DIVINATORIUS MUELL

Euth Chase Noland

Introduction

The internal anatomy of Troctes divinatorius has never been

carefully worked out, though the insect has frequently been de¬

scribed as a not uncommon household pest. In 1818 Nitzsch (18)

published a description and figures of the digestive tract and re¬

productive organs of an insect which he called the book louse,

Psocus pulsatorius. In a number of ways my descriptions differ

from his, and it is probable that the insect which I have had is a

different species, although of the same genus, for the generic names

Psocus, Troctes, and Atropos have all been used in reference to the

genus to which the book louse belongs. Since the time of Nitzsch,

the only work which has been done on the book louse from an

anatomical point of view has been that on the mouthparts, and is

very little. However, considerable study has been made of the

mouthparts of a closely related family, the Psocidae. This has

been done by Burgess (6), Hagen (15), and others and has been

of interest largely because of the light it throws on the relation¬

ships of these to other insects.

The studies made here of Troctes are based on examinations of

hundreds of individuals in whole mounts, longitudinal, and cross

sections and in dissections made in glycerine or in balsam under

the dissecting microscope. The small size of the specimens, none of

them more than a millimeter and a half in length, has made neces¬

sary the working out of many details by comparison in a large

number of specimens. For the gross anatomy the micro-dissections

stained in alum carmine, or haemalum gave most information;

while the smaller details and histology could only be determined

from sections. These were stained with Heidenhain’s or Delafield’s

haematoxylin. A number of fixatives were used, but probably the

best results came from Carnoy’s I and II, and Kahle, Tower and

Petrunkewitsch. Very much assistance in the work was given by

Professor W. S. Marshall, under whose direction it was carried out.

196 Wisconsin Academy of Sciences, Arts, and Letters,

The specimens for use were easy to procure at certain times of

the year, for during the summer and autumn months a sheet of

paper left for a few days on one of the laboratory tables is sure to

cover a number, large and small, but in the winter, beginning with

November and continuing until May, the insects almost entirely

disappear. Their favorite habitat is a dry and undisturbed place,

and the tables of a laboratory, if dust-covered, suit them perfectly.

It is useless to search for them on a clean table, even though it has

been well sheltered by papers or books. Books infrequently used,

or herbarium shelves, or dried grains packed away in boxes re¬

mote from sunlight are favorable habitats for them. As has often

been noted in descriptions, the femur of the posterior pair of legs

of Troctes is greatly enlarged, a characteristic of jumping insects,

but observers have claimed this to be of no use to the animal. How¬

ever, in escaping from the collector, as they probably do in eluding

animals which prey upon them, they put this feature to good use

by making leaps backward before fleeing swiftly into some dark

crevice. Before they can hide, they are easily picked up with a

moist brush and dropped into hot water.

Mouthparts

The structure of the head and mouthparts of the Corrodentia has

been of more interest to entomologists, particularly systematists,

than any other feature of the anatomy of these insects, because of

the similarity shown by them to corresponding parts of the Mal-

lophaga. The oesophageal sclerite or ‘‘bonnet,” the so-called max¬

illary fork, and the chitinous structures of the labium known as the

lingual glands are similar, to a striking degree, in the two orders.

The first work which revealed to any extent the structure of the

mouthparts of a form similar to Troctes was that of Burgess (5)

in 1878 on the anatomy of the head in the Psocidae. In this he

figured in detail some portions of the head and mouthparts of

Psocus, with a few references to those of Troctes (called by him

Atropos). Since his time the studies which have been made on

these insects have been largely to determine the nature of the

“lingual glands” and maxillary fork. Doubt has been expressed

as to the glandular nature of the labial structures by Bertkau (4),

Cummings (8) and Enderlein (10), and the true relationship of

the maxillary fork is unknown. Is it an independent organ, not

Noland — Anatomy of Troctes Divinatorins Muell.

197

maxillary in origin as Burgess concluded, or does it represent the

inner lobe or lacinia of the maxilla, as claimed by Enderlein f

The head of Troctes is flattened dorso-ventrally with a large pro¬

jecting clypeus filled with muscles which move the oesophageal

sclerite. The internal support of the head is given it by the ten¬

torium (fig. 1), which consists of a central plate and two pairs of

arms; the anterior ones (fig. 1, a.t.) arise from the head capsule

at the point of attachment of the mandibles, and the posterior pair

(fig. 1, p.t.) appear to arise from the head capsule posterior to the

muscles attaching the mandibles and maxillary fork to the posterior

wall of the head in the occipital region.

In Troctes the inner surface of the clypeus, rather than the

labrum as in Psocus, is furnished with tufts of hair (fig. 15, c.h.),

and the labrum extends downward from the clypeus to meet the

labium. The mandibles (fig. 2), heavy, chitinized structures, lie

beneath the labrum. The dorsal portion of their inner or mesad

surface is molar in nature for crushing (fig. 2, m.s.), while the

ventral edge is made up of three sharply pointed teeth (fig. 2,

v.p.), heavy enough to provide a satisfactory cutting surface.

The dorso-lateral angle of each mandible is adapted to serve as

the point of attachment to the lateral wall of the head capsule.

The middle part of this area is rounded into a knob (fig. 2, m.p.)

which fits into a corresponding socket in the wall of the capsule.

Ventral to this is another projection (fig. 2, a.ab.) from which a

small abductor muscle passes into the head just ventral to the base

of the antenna. The mesad dorsal surface of the mandible is in¬

dented slightly (fig. 2, a.ad.) to form the point of attachment for

a chitinous rod (fig. 1, c.m.) which is held in place by tendons.

This rod spreads out posteriorly into several large bundles of

adductor muscles (fig. m.m.) which fill a great part of the head

cavity and are inserted in the posterior wall of the head. The

maxillae and maxillary forks lie between the mandibles and labium.

Each maxilla proper is composed of a main stipes (fig. 5, s.) which

supports a four-jointed maxillary palp (fig. 5, m.p.) and a broad

galea (fig. 5, g.). Burgess (6) describes a cardo in Psocus, but in

Troctes this was not found. The inner edge of the galea is chitin¬

ized and has a few blunt teeth or ridges. This structure may be

swung on the stipes by means of two muscles extending to the

outer edge of the stipes (fig. 5, gme. and gmf.). The first muscle,

attached to a place on the stipes anterior to the second, serves as

the extensor, pulling the galea out laterally; the second acts as a

198 Wisconsin Academy of Sciences, Arts, and Letters,

flexor to draw it inward toward the median line. The maxillary-

palp is pubescent. Its flrst or basal segment is very short, and the

third is like it ; while the second and fourth are longer, each fully

three times the length of either of the other two. The largest

muscles of the palp (flg. 5, f.p.) extend to the inner edge of the

stipes, crossing it ventral to those of the galea. Thus their con¬

traction would serve to draw the palp in toward the median line;

a shorter muscle (fig. 5, ep.) attached near to those of the galea

swings it out. On the stipes, just at the base of the galea, is a

groove (fig. 5, fs.) through which the maxillary fork slides as if in

a slot, and in which it is held in place by a band of tissue which

arises from one wall of the slot and passes over the fork to attach

on the other side. This slot has not been referred to before ex¬

cept as Hagen (15) described the fork as sliding “in the outer

lobe as in a vagina ’ ’ ; yet this he did not figure. He spoke of the

tip of the fork as sliding through a chitinous ring of the tip of the

outer lobe, but this has not been seen in any of my preparations.

The fork itself in Troctes is a straight rod ending distally in three

sharp points of unequal length, the two outer longer than the

inner. These ends are unlike the truncated tips of Psocus. At its

base the fork is covered by strong muscles which in turn connect

with a muscle (fig. 5, r.f.), not a ligament, as claimed by Burgess.

This is probably the fork retractor, as it extends to the posterior

wall of the head and would thus serve to draw the fork within the

head by its contraction. The function of the forks is difficult to

conceive, unless they could be used in so delicate a process as that

of picking up mold spores and other small particles and drawing

them into the mouth. With the close attachment which the maxil¬

lary fork bears through the slot to the stipes of the maxilla, it be¬

comes possible to consider it the homologue of an inner lobe, since

there is no other sign of one in connection with the stipes. How¬

ever, even considering this, with the ultimate attachment of the

fork to the posterior wall of the head, the old hypothesis of

Burgess, that it is an independent organ, is still possible. In dis¬

section it is so easy to separate the fork from the maxilla that its

real attachment through the slot is difficult to ascertain. Until

observation is made of its origin in development this question will

remain a matter of doubt.

The labium is made up of a broad mentum (fig. 6, m.) which

bears a pair of one-jointed labial palps (fig. 6, l.p.) — short, bluntly

pointed, and covered with bristle-like hairs, — and two median, hair-

Noland — Anatomy of Troctes Divinatorius Muell.

199

covered fleshy lobes which form the ligula (fig. 6, v.L). The men-

tum is borne on a very short submentum (fig. 6, s.m.). So closely

attached to the dorsal side of the labium as really to form a part

of it, though designated by Burgess the tongue, is a thin plate

(fig. 6, d.p.) which extends undivided as far forward as the tips

of the lobes of the ligula, and in it and the labium itself the labial,

or ^‘lingual’’ glands (fig. 6, l.g.) are imbedded. The glandular

nature of these structures has been doubted, but their appearance

when examined in section is distinctly that of glandular cells.

Each cell has a nucleus large in proportion to the size of the cell

and containing a distinct nucleolus. It is difficult to explain the

fact that the outer walls of these glands are of heavy chitin, but

they are obviously neither reservoirs, nor merely chitinous sup¬

porting structures as has been supposed. Unlike the glands of

Psocus, those of Troctes are not supported by any sort of cap or

peduncle. They have no support except their position, imbedded

in the labial tissue. From them chitinous ducts extend a short dis¬

tance forward and then turn backward to unite with one-another

between the two glands. The common duct so formed leads back

between rows of gustatory cones (fig. 6, t.c.) or pegs, more rounded

than hairs, into the oesophageal sclerite between its two anterior

horns. This sclerite (figs. 4 and 6, o.s.) is a chitinous ‘^bonnet-

shaped’’ structure which extends two anterior arms, or horns, for¬

ward for support in the labium, and one backward onto a flat chit¬

inous plate which is held on each side by two smaller rods extend¬

ing out toward the sides of the head. The large masses of muscles

of the clypeus, already mentioned, which are attached to the dorsal

wall of the oesophageal sclerite, are said by Burgess (5) to function

in closing the oesophagus, but it seems improbable that their con¬

traction, situated as they are, could produce such an action.

The Digestive System

The mouth opens through the oesophageal sclerite into the long

slender oesophagus, or fore-intestine. This extends through the

thorax into the first or second abdominal segment where it joins

the mid-intestine. The anterior portion of the oesophagus is nar¬

row and has its origin at the oesophageal sclerite, turning in a

slightly dorsal direction before passing posteriorly. In the thorax

it enlarges into a crop-like structure (fig. 7, cr.) which has thin

walls histologically the same as those of the oesophagus. At the

200 Wisconsin Academy of Sciences, Arts, and Letters.

posterior end of the crop, lying close to this part of the digestive

tract are four structures, presumably salivary glands, though it

is impossible to find an outlet for them. The two of these lying in

closest proximity to the crop are large and nearly oval in outline

(fig. 7, l.g.) ; while the others are longer, more slender, and with

rounded ends posteriorly. Their position is lateral and ventral to

the larger ones. When studied in microscopic section, there appear

to be six structures instead of four, for the anterior portion of the

tubular pair is histologically distinct. The posterior part of each

tubular organ (figs. 7 and 8, g.r.) has a broad lumen and the cells

which make up its walls are probably syncitial, since no cell walls

show, though nuclei are scattered throughout it at intervals. The

structure of these leads to the conclusion that they are gland reser¬

voirs rather than glands. The anterior part (fig. 8, s.g.) has no

lumen, but is made up of cells which sometimes show droplets of

secretion in their cyptoplasm. The histological arrangement of

these cells is similar to that of the large oval bodies mentioned

above and the two pairs of structures are probably connected. The

oval bodies have no lumen and are composed of large glandular

cells, the nuclei of which are large and stain heavily, while the

cytoplasm stains irregularly. These structures are glands and are

connected with the reservoirs through their anterior ends, though

this connection is not discernible. Their appearance . suggests the

three pairs of glands described by Uzel (22) in the thoracic region

of the Thysanoptera. He does not show any ducts leading from

them.

Faure-Fremiet (13) describes in the Hydrocorises a pair of

labial glands which lie in the thorax, even extending into the ab¬

domen, and are composed of an anterior and posterior lobe uniting

anteriorly, and, where they come together, joining the excretory

duct of a gland reservoir. An arrangement similar to this is very

possible in Troctes, because of the relation the various parts bear

to each other.

The fore-intestine with its thin walls and wide channel, narrows

abruptly at its posterior end to form the cardiac constriction, or

valve (fig. 15, c.J.

The mid-intestine (fig. 7, m.) beginning with this valve shows in

section a sharp transition lo thick glandular walls, markedly folded.

The content of this part of the digestive tract reveals the food of

the book louse, for it is often made up of the spores of molds and

small particles of other organic matter. At times the contents of

Noland — Anatomy of Troctes Divinatorius Muell.

201

the digestive tract is made up largely of brown crystals. The diet

on the whole appears to be gained from the organic material in

particles of dust^ rather than from gnawing portions from dried

grain or specimens. Frequently the mid-intestine is found to be

two-thirds filled with large gregarines, the only parasites discov¬

ered in the digestive tract.

The hind-intestine is at its beginning much narrower than the

mid-intestine. The anterior portion, or ileum, is the longest part,

and is coiled once toward the dorsal wall of the abdomen. Imme¬

diately after coiling, the ileum enlarges into the short, broad rec¬

tum (fig. 7, rg.) which is fully as wide as the mid-intestine. Its

walls contain four swellings which, in section, appear to be glandu¬

lar. These rectal glands are composed of large cells with large

nuclei, and both cells and nuclei stain heavily. The rectum nar¬

rows abruptly before it opens to the outside in the ninth segment

of the abdomen.

At the beginning of the ileum four Malpighian tubules (fig. 7,

m.t.) open. These extend anteriorly half or two-thirds of the

length of the abdomen. Each tubule stretches anteriorly for about

half of its length and then doubles back on itself, coiling about,

with its blind end in the posterior part of the abdomen, near the

rectum.

The Female Reproductive System

The two ovaries of the female reproductive system of Troctes,

lying one on either side of the mid-intestine and between the intes¬

tine and the dorsal wall of the abdomen, are each made up of five

ovarian tubules (fig. 9, o.). Each tubule is of the type in which

nutritive cells alternate with egg cells. The young oocyte (fig. 11,

ov.) is found at the proximal end. Distal to it in the tubule are

three or more large nutritive or nurse cells (fig. 11, n.c.) in a sin¬

gle chamber which supply nutriment to the oocyte. The apex of

the ovarian tubule forms a terminal chamber (fig. 11, t.c.) contain¬

ing a mass of undifferentiated cells, and the tubule terminates dis-

tally in a long terminal filament which unites with those from the

other ovarian tubules to form a single filament (figs. 9 and 11, f.)

that serves to hold them in place in the abdomen. The proximal

ends of the tubules are long and slender and unite to form the two

short oviducts (fig. 9, od.) which join almost immediately in the

broad vagina (fig. 9, v.) that opens to the outside at the posterior

202 Wisconsin Academy of Sciences, Arts, and Letters.

ventral edge of the eighth segment. On the left side of the abdom¬

inal cavity lies the seminal receptacle (fig. 9, s.r.), a thin-walled

sac, narrowed at each end and containing the spermatozoa. These

can be readily seen through its walls in cleared whole mounts of

the insect and in sections, and the thinness of its walls precludes

the possibility of glandular structure. From the posterior end of

the seminal receptacle a slender, delicate duct (fig. 9, d.) leads to

the vagina, entering it midway between the point of union of the

oviducts and its opening to the outside. No accessory glands are

present in Troctes.

In older individuals than the one represented in the figure, the

eggs are found in later stages of development. However no insect

was found with more than one fully developed egg at a time. This

egg is so large before laying that it occupies fully one-third of the

abdominal cavity ; in insects mounted in glycerine an outer chorion

and yolk granules can be seen. In ovarian tubules on the other

side of the abdomen from the well-developed egg, other eggs show

considerable development, but none equal in size to the first.

Though careful search was made during the breeding season, no

eggs were ever found after they were laid.

The Male Reproductive System

The most conspicuous structures of the male reproductive sys¬

tem are not the testes, but the paired seminal vesicles (fig. 13, v.)

which lie a little to the left of the median line of the abdomen.

These are large, when fully matured extending nearly to the

thorax, and they lie so close to each other as often to give the ap¬

pearance of a single organ. An examination of transverse sec¬

tions, however, proves it is a paired organ, its parts joining to

form a single large duct at their proximal ends. This, the ejacula¬

tory duct (fig. 13, e.d.), leads to the outside of the body through

the copulatory apparatus (fig. 13, c.a.). The testes (fig. 13, t.) are

round or oval bodies lying near the lateral abdominal walls. From

a surface view they give the impression of being a mass of coiled

tubes, and when sectioned, they appear to be separated into areas,

representing coils, which contain the spermatozoa at different

stages of development (fig. 12, s.). The central portion of ea^h

testis is formed of spermatogonia, and the posterior portion, from

which the vas deferens (fig. 13, v.d.) arises, is filled with spermato¬

zoa. At each end of the testis is a darkly staining mass (fig. 12,

Noland — Anatomy of Troctes Divinatorius Muell.

203

d.b.) whose function was not determined. The vasa deferentia,

which are long, slender, transparent tubes, lead by a somewhat

winding course to the seminal vesicles, entering them laterally just

anterior to their union in the ejaculatory duct.

The copulatory apparatus through which the ejaculatory duct

passes is a complicated chitinous structure, the anterior end of

which makes an acute angle ; while the posterior part terminates in

a pair of hooks which curve in toward each other and close about

the end of the ejaculatory duct. When drawn well into the body,

as figured, the anterior tip of the structure reaches as far anteriorly

as the posterior ends of the seminal vesicles, but the apparatus can

be thrust out of the abdomen to nearly one-half of its total length.

This copulatory apparatus consists of three pairs of chitinous

plates (fig. 13, prs. 1, 2, 3). The inner pair (fig. 13, pr. 1), broad

and flat and slightly curved at the posterior ends, is in contact with

the ejaculatory duct and forms the foundation to which the other

two pairs are attached. The acute angle of the apparatus is

formed by the second pair (fig. 13, pr. 2) which is more heavily

chitinized and slender than the others and comes in contact with

pair one at the anterior ends of the latter, ending there. The third

pair (fig. 13, pr. 3) is heavy, like the first, and is attached to both

of the other pairs, uniting with the first pair at its posterior end

and forming there the outer portions of the terminal hooks (fig.

13, t.h.). The anterior parts of the third or outer pair unite with

the second pair at about their middle. Thus the third pair vir¬

tually forms both a sheath for the posterior parts of the other two,

and the posterior hooks or ‘ ‘forceps themselves.

The Nervous System

The central nervous system of Troctes is of a simple type, com¬

posed of the supraoesophageal ganglion or brain (fig. 15, s.g.), the

suboesophageal ganglion (fig. 15, sb.g.), and three thoracic ganglia

(fig. 15, g.l, 2, 3). The last two thoracic ganglia are fused into a

large mass which appears externally to be but one, though in sec¬

tion its compound nature is revealed. The length of this last

ganglion and a slight constriction at its posterior end makes it

appear as if a third ganglion, originally abdominal, may have

taken part in its formation. This ganglion sends out a nerve, or

group of nerves (fig. 15, n.) as a prolongation from its posterior

204 Wisconsin Academy of Sciences, Arts, and Letters.

end which soon disappears in the abdomen and represents the total

nerve supply of that part of the insect’s body.

The supraoesophageal ganglion is broad, its central portion ex¬

tending anteriorly in a projection which may be an ocellar lobe

(fig. 14, 0.1), though no ocelli are visible on the exterior of the

head. Two lobes on the right and left of the median one send

nerves to the mouthparts, and just lateral to these are the anten-

nary lobes (fig. 14, a.n.). The largest lobes are the optic, (fig. 14,

op.l.) reaching laterally almost to the eyes. The supraoesophageal

ganglion ends posteriorly in two prominent projections which in

sections seem largely cellular with little “ punktsubstanz ”, or

medullary material, extending into them (fig. 14, p.l.). The func¬

tion of these is not clear.

The supraoesophageal ganglion connects with the suboesophageal

ganglion by the circumoesphageal commissures (fig. 15, c.o.c.)

which pass from its anterior ventral surface around the oesophagus.

The Respiratory System

The tracheal system can be distinguished only in specimens

which have been a short time in glycerine. So treated the air is re¬

tained in the tracheal tubes and they appear black against a trans¬

parent background. Even when thus prepared only parts of the

system are evident; so that any reconstruction of the tracheae

must be a composite from many specimens.

The respiratory system of Troctes is of a simple type. In the

abdomen are six pairs of stigmata or spiracles, one pair in each of

the first six segments. From each a short trunk leads to a main

lateral longitudinal one (fig. 16, l.t.) which extends the length of

the abdomen and is continuous with one of the main trunks of the

thorax and head. Each of the stigmatal trunks in the abdomen

gives off typically a visceral branch (fig. 16, vs.b.) to the digestive

and reproductive organs, a ventral branch (fig. 16, v.b.) to the

ventral body wall, and a dorsal branch (fig. 16, d.b.) which sup¬

plies the dorsal region of the body with oxygen. In the posterior

region of the abdomen the longitudinal trunks break up into nu¬

merous smaller branches which supply the last three segments.

In the prothorax and in the head, the longitudinal trunks are con¬

nected to each other by transverse trunks (fig. 16, t.t. and h.t.), and

in the head the tracheae are numerous and small, a large number

of branches reaching into the mouthparts, antennae, and eyes.

Noland — Anatomy of Troctes Divinatorius Muell.

205

Parasites of Troctes

In so small an insect, it was rather surprising to find parasites

so common. As has already been mentioned, gregarines not infre¬

quently are found in the mid-intestine. Once a parasitic protozoan

was recognized in a longitudinal section of the testes.

The parasite most commonly found is a large nematode worm

located in the body cavity. The small males and long, coiled

females were found twisted about in the abdomen, even extending

into the thorax. Sometimes one female nematode and as many as

a half dozen of the small males were dissected out of a single

Troctes. The males are short, less than one twenty-fifth the length

of the females, whose long bodies are filled with eggs. All the

specimens of the insect which contained this parasite were gath¬

ered from one place — a box which had for several years been kept

filled with stalks of dried grain to make a breeding place. The

hosts of the worms were as active as any of the non-parasitized

individuals, but could usually be detected by their greater size.

Bibliography

1. Banks, Natkan. Catalogue of the Neuropteroid Insects (except Odon-

ata) of the United States. Am. Ent. Soc., Philadelphia, 1907

(Separate publication).

2. Banks, Nathan. Two new species of Troctes. Entom. News, 11: 559.

1900.

3. Berlese, A. Gli Insetti, 1: Pt. 1.

4. Bertkau, P. Tiber die Speicheldriisen der Psociden, Verb. Nat. Ver.

Bonn, 39: 130. 1882.

5. Burgess, Edward. The anatomy of the head and the structure of the

maxilla in the Psocidae. Pros. Bost. Ent. Soc. Nat. Hist. 19:

291. 1878.

6. - On the structure of the head of Atropos. Psyche, 2: 87.

1877-78.

7. Butler, E. A. Our household insects. London, 1893.

8. Cummings, Bruce F. On some points in the anatomy of the mouth-

parts of the Mallophaga. Proc. Zool. Soc. London, Part 1: 128.

1913.

9. Derham, W. A letter concerning an insect that is commonly called the

Death-watch. Phil. Trans. Eoy. Soc. London, 22: 832. 1701.

- A supplement to the account of the Pediculus Pulsatorius,

or Death-watch. Phil. Trans. Eoy. Soc. London, 24; 1586. 1704.

10. EnderZein, G-. tiber die Morphologie, Gruppierung, und Systematische

Stellung der Corrodentien. Zool. Anz. 26: 423. 1903.

11. - Morphologie, Systematik und Biologie der Atropiden und

Troctiden. Eesults of Swedish Zool. Exped. to Egypt and the

White Nile, 1901. Uppsala, Part I. 1904.

206 Wisconsin Academy of Sciences^ Arts^ and Letters,

12. Evans, A. M. On the structures of maxillulae in insects. Journ. Linn,

Soc. London, 34: 429. 1921.

13. Faure-Fremiet, E. Contribution a 1’ etude des glandes labiales des

Hydrocorises. Ann. Sc. Nat., 9 ser. 12: 217. 1910.

14. Hagen, H. A. Psocinorum et Embidinorum Synopsis Synonymica. Ver-

handl. Zool.-Bot. Gesell. Wien. 16: 201. 1866.

15. - Some Psocina of the United States. Psyche, 3: 196, 206,

219. 1881.

16. - Synopsis of Neuroptera of N. America. Smithsonian Misc.

Coll. Washington. 1861.

17. Jordan, Karl. Anatomie und Biologic der Physapoda . Hann. Miinden.

1888.

18. Nitzsch, 0. L. Ueber die Eingeweide der Biicherlaus (Psocus pulsa-

torius) und uber das Verfahren bei der Zergliederung sehr kleiner

Insekten. Germar’s Mag. Entom. 4; 276. 1821.

19. Rambur, M. P. Histoire naturelle des insectes neuropteres. Suites a

Buffon. Paris. 1842.

20. Scudder, Samuel H. On the structure of the head of Atropos. Psyche,

2: 49. 1877.

21. Snodgrass, R. E. A Revision of the mouth-parts of the Corrodentia

and the Mallophaga. Trans. Am. Ent. Soc. 31: 297. 1905.

22. TJzel, Jindrich. Monographic Radu ‘ ‘ Thysanoptera. ’ ’ Koniggratz.

1895.

23. Viallanes, H. Centres nerveux et les organes des sens des animaux

articules. Bib. de PEcole des Hautes Etudes, 33: 1887.

24. Westwood, J. B. Introduction to the modern classification of insects.

2: 17.

Explanation of Figures

Plate IV

All figures drawn with camera lucida.

Fig. 1. Outline of head and tentorium from the ventral side. X 160.

a.s. The first segment of the antenna, a.t. The anterior arm of the ten-

otrium. c.m. The chitinous supports of the mandibles, e. The eye. 1. The

labrum. m. The mandibles, m.f. The maxillary fork. m.m. Mandibular

adductor muscle, o.s. The oesophageal sclerite. p.t. The posterior arm of the

tentorium.

Fig. 2. The Mandibles. X 250.

a. The right mandible, b. The left mandible, a.ab. The point of attach¬

ment of the abductor muscle, a.ad. The point of attachment of the ad¬

ductor muscle, m.p. The pivotal knob which fits into the head capsule, m.s.

The molar surface of the mandible, v.p. The ventral points of the mandible.

Fig. 3. Enlarged view of the cells of the right labial gland. X 930.

n. The nucleus, nu. The nucleolus.

Noland — Anatomy of Troctes Divinatorius Muell.

207

Fig. 4. Lateral view of the oesophageal sclerite. X 250.

a.h. The anterior horn of the sclerite. c.d. The common duct of the labial

glands. 0. Oesophagus, o.s. The oesophageal sclerite. p.h. The posterior

horn of the sclerite. t.c. The taste cones.

Pig. 5. Dorsal view of the maxilla and maxillary fork. X 250.

e. p. The extensor muscle of the maxillary palp. f.m. The muscles of the

maxillary fork. f.p. The flexor muscle of the maxillary palp. f.s. The groove

of the maxillary fork. g. The galea, g.m.e. The extensor muscles of the

galea, g.m.f. The flexor muscle of the galea, h.w. The posterior wall of the

hea^. m.f. The maxillary fork. m.p. The maxillary palp. p.m. The muscles

of the second segment of the maxillary palp. r.f. The retractor muscle of

the maxillary fork. s. The stipes.

Fig. 6. The labium and oesophageal sclerite from a dorsal view. X 250.

a.h. The anterior horn of the oesophageal sclerite. d.g. The duet of the

gland, d.p. The dorsal plate of the labium which contains the glands, l.g.

The labial gland, l.m. The labial muscles, l.p. The labial palp. m. The

mentum. o.s. The oesophageal sclerite. p.h. The posterior horn of the

oesophageal sclerite. s.a. The supports which help to attach the sclerite in the

mouth, s.m. The submentum. t.c. The taste cones on either side of the com¬

mon duct. v.l. The ventral lobe or ligula of labium as seen through the trans¬

parent dorsal plate.

Plate V

Fig. 7. The dorsal view of the digestive system. X 160.

cr. The crop. i. The ileum, g.r. The reservoir of the salivary gland, l.g.

The salivary gland, m. The mid-intestine, m.t. One of the Malpighian

tubules. 0. The oesophagus, o.s. The oesophageal sclerite. r.g. The rectal

gland.

Fig. 8. Longitudinal section of one of the salivary glands and its reservoir.

X 160.

g. The salivary gland, g.r. The gland reservoir, s.g. The anterior end of

gland reservoir.

Fig. 9. A dorsal view of the female reproductive system of a young specimen.

" X 160

d.The duct of the seminal receptable. f. The terminal filament, o. An

ovarian tubule, od. An oviduct, s.r. The seminal receptacle, v. The vagina.

Fig. 10. A cross section of the salivary glands and reservoirs. X 250.

cr. The crop. g. The salivary gland, g.r. The gland reservoir.

Fig. 11. A single ovarian tubule. X 465.

f. The terminal filament, g. The nucleus, n. The nucleus of a nurse or

nutritive cell. n.c. Nutritive cell. o.d. The duct of the tubule, ov. The

oocyte, t.c. The terminal chamber.

208 Wisconsin Academy of Sciences^ Arts^ and Letters.

Plate VI

Fig. 12. Longitudinal section of testis. X 465.

d.b. The dark body. s. Spermatozoa, sg. Spermatogonia, sp. Spermatids^

Fig. 13. The male reproductive system from a ventral view. X 160.

c.a. The copulatory apparatus, e.d. The ejaculatory duct. Pr.l, Pr.2, Pr.3.

The first, second, and third pairs of chitinous structures of the copulatory

apparatus, t. The testes, t.h. The terminal hooks of the copulatory ap¬

paratus. vd. The vas deferens, v. The seminal vesicle.

Fig. 14. Composite drawing of the brain. Outline of head. X 250. .

a.l. The antennal lobe. a.n. The antennal nerve, e. The eye. lb. The

labrum. o.l. An ocellar lobe. op.l. The optic lobe. p.l. The posterior lobe.

Fig. 15. Longitudinal section of the central nervous system. X 135.

c.h. Hairs on the clypeus. e.o.c. The circum-oesophageal commissure, c.

The cardiac valve, g.l., g.2., and g.3. The first, second and third thoracic

ganglia, n. The abdominal nerve, o.s. The oesophageal sclerite. sb.g. The

suboesophageal ganglion, s.g. The supra-oesophageal ganglion or brain, t.

A portion of the tentorium.

Fig. 16. A composite diagram of the tracheal system from a ventral view.

X 160.

a.b. The antennal branch, d.b. A dorsal abdominal branch, e.b. The branch

to eye. h.t. The transverse connecting trunk of the head. l.t. The main

longitudinal trunk of the abdomen, t.s.t. A thoracic stigmatal trunk, t.t.

The transverse trunk of the thorax, v.b. A ventral branch, vs.b. A visceral

branch.

Noland — Anatomy of Troctes Divinatorius Mnell. 209

TRANS. WIS. ACAD., VOL SXI

PLATE IV

14

210 Wisconsin Academy of Sciences, Arts, and Letters.

TEANS. WIS. ACAD., VOL XXI PLATE V

Noland — Anatomy of Troctes Divinatorius Miiell.

211

TRANS. WIS. ACAD., VOL. XXI

PLATE VI

ARRHENURI FROM WASHINGTON AND ALASKA

Ruth Marshall

In the summer of 1922 the author made an extended trip to

Alaska, from Seattle to Kodiak Island. A number of localities in

Alaska and the Canadian Northwest were visited and water mites

were collected in many small bodies of water. The preliminary

work on the study of this material has been done; it is proposed

to publish in the near future a complete account of the species

found and their distribution. The present paper is an account of

the Arrhenuri of the collections.

Arrhenurus tacomaensis nov. spec.

PI. VII, fig. 1-5

While en route to Alaska, the author had an opportunity to

make small collections of water mites in two localities in the state

of Washington. The first place visited, July 7, 1922, was the inlet

of Union Bay, at Seattle, near the University of Washington. In

the shallow water, bordered by tall grasses and other plants, six¬

teen individuals were found, representing four different genera

(Limnesia, Neumania, Fiona and Arrhenurus). There was but

one Arrhenurus, a male; this proved to be A. krameri Koenike,

which has already been reported by the author from Oregon.

The second collecting ground was at Tacoma, September 4.

Near the car-line at 52nd Street is a small shallow pond filled

with yellow pond lilies in which seven individuals were found.

Five of these are undetermined species of Fiona and Unionicola.

The remaining two specimens are Arrhenurus males of a new

species which has been given a specific name indicative of the local¬

ity from which it was first reported.

A. tacomaensis belongs to the subgenus Arrhenurus and some¬

what resembles A. lautus described by Dr. Koenike from Alberta.

The body is stout; two sickle-shaped projections arise dorsally at

the place where the body passes abruptly into the appendix. This

214 Wisconsin Academy of Sciences , Arts, and Letters.

caudal appendix has pronounced lateral projections and a clearly

defined hyaline appendage. The petiole is conspicuous; the sides

roll over dorsally to enclose a central oblong structure arising

out of it.

The palpi are stout, with a few large bristles. The fourth joint

of the fourth leg has a well defined thumb-like projection. The

color of the new species is dull red with indistinct brown areas

where the internal organs show through the body wall. The legs

have a greenish tinge. The total length is 1.15 mm., the greatest

width, 0.9 mm.

Arrhenurus elongatus nov. spec.

PI. VII, fig. 6, 7 ; PL VIII, fig. 14, 15

It is a pleasure to be able to record the finding of a new species

of Arrhenurus from Alaska, the first account of any work on the

water mites from this territory that the author is aware of. Quite

appropriately this species comes from the old Russian town and

former capitol, Sitka. ^ It was found in the border of a small shal¬

low pond choked with yellow pond lilies and like plants, in a boggy

meadow near the town. With it were a few individuals of other

genera, the species not yet determined.

A. elongatus is a large mite belonging to the subgenus Megalura-

carus. It is 1.5 mm. long, with a width of 0.73 mm. It resembles

A. solifer Marshall and A. pseudoconicus Piersig, both American

forms, and A. conicus Piersig, a European species. In this group

of related species the appendix narrows conspicuously at the end.

The new species, however, is more slender and has a more elongated

appendix than in any of the nearly related species, a feature so

characteristic that it has suggested the specific name.

Only one individual, a male, was found. The body is widest

and most elevated in the anterior half; likewise, the appendix is

largest in the proximal half, though constricted where it joins the

body. The end of the appendix is bowed out and has several small

humps and hairs, as shown in the figures. The three groups of the

epimera are close together ; the first and second pairs of plates have

sharp projecting anterior corners, and the fourth epimera are

very wide. The genital area is small and does not project beyond

the body wall. The color of the animal is dull red.

Marshall — Arrhenuri From Washington and Alaska. 215

Arrhenurus acerfonnis nov. spec.

PL VIII, fig. 8, 9, 13

At Chitna, Alaska, about one hundred and thirty-five miles in¬

land on the Copper River & Northwestern Railway from the sea¬

port town of Cordova, a number of small ponds were examined.

Only one of these yielded any Arrhenuri; this was a pond in the

mountains, surrounded by the forest, and called Lost Lake. It is

a circular body of water, perhaps a half mile in circumference,

apparently shallow, with a border of yellow water lilies among

sunken logs at one place. This pond is said to be seven hundred

and fifty feet above the town, some two miles away. Only two

Arrhenuri were found, both females, but representing two species.

As this genus is usually very common in the quiet waters of low¬

lands and plains, much interest attaches to the finding of these

specimens at this elevation. It is hoped that the males may be dis¬

covered in future collections by the aid of the accompanying fig*^

ures, since the palpi are alike in the two sexes.

A. acerformis, female, is oval in form, 1.45 mm. long and 1.1 mm.

across at the widest part. The color is dull olive green. The

epimera of the first pair end in rather sharp points. The genital

area is some distance from the fourth pair of epimera ; in form, the

wing-like areas bear a fanciful resemblance to the twin seeds of

certain maples, and this has suggested the specific name.

Arrhenunis hirsutus nov. spec.

PI. YIII, fig. 10-12

This is a smaller species, found with the preceding species, and

represented also by but one individual, a female. It is pyriform

in shape, 1.2 mm. long and 0.96 mm. in the widest part. The gen¬

ital area is close to the last epimera, and the wing-like expansions

are short and inclined from the plates guarding the opening. The

palpi have several long stout hairs or bristles on the second joint,

a character which has suggested the name of the species. The color

is the usual dull olive green.

216

Wisconsin Academy of Sciences, Arts, and Letters.

Explanation of the Plates

PLATE VII

tacomaensis, dorsal view.

tacomaensis, appendix, ventral view.

tacomaensis, lateral view.

tacomaensis, fourth joint of fourth leg.

tacomaensis, right palpus.

elongatus, dorsal view.

elongatus, lateral view.

PLATE VIII

Fig. 8.

Fig. 9.

Fig. 10.

Fig. 11.

Fig. 12.

Fig. 13.

Fig. 14.

Fig. 15.

Arrhenurus

acerformis fern., ventral view.

acerformis fern., dorsal view.

hirsutus fern., dorsal view.

hirsutus fern., genital area.

hirsutus fern., left palpus.

acerformis fern., left palpus.

elongatus, left palpus.

elongatus, fourth joint of fourth leg.

Eockford College,

Eockford, Illinois.

Marshall — Arrhenuri From Washington and Alaska. 217

TRANS. WIS. ACAD., VOL. XXI PLATE VII

218 Wisconsin Academy of Sciences, Arts, and Letters.

TRANS. WIS. ACAD., VOL. SXI PLATE VIII

NEW AND CORRECTED NAMES OF CERTAIN

MILK BACTERIA*

W. D. Frost and Ruth Chase Noland

We are not aware that there is anywhere a list of the bacteria

that have been found in milk later than that of Conn, Esten and

Stocking (1906). Since then, many species have been recognized

and some new ones described as occurring in milk. Furthermore,

the Society of American Bacteriologists, through their Committee

on Characterization and Classification (1920), have so modified the

genera that many species will have to be renamed. Few bacteriol¬

ogists have ever bothered with synonyms so that there is really

very great confusion in names. This is perhaps especially true

with dairy bacteria because of the wide use of Conn’s list in which

he has paid little attention to synonomy and made general use of

the unscientific trinomial system of nomenclature.

The adoption by the Society of American Bacteriologists of rules

and a definite system of classification makes for progress and it

would seem that the time is now ripe for an attempt to list the

milk bacteria and assign to them the correct names.

Such a task is a long and difficult one and perhaps is only likely

to be done piece-meal, yet if the work is carefully done, it should

be welcomed and valuable.

We list below a number of species in which the usual names have

been changed or new ones supplied because of the application of

the priority rule and those that have been changed from one genus

to another:

New Names

Bacillus involutus (adametz)

B. No. 15 y ADAMETZ, 1889, Landw. Jahrb. 18: 247.

Bact. turgidnm Chester, 1901, p. 195.

Note: This is now a bacillus but the name iurgidus used by Chester is

invalid because Duclaux has previously used the name. See Mace, 1897, p.

900, also Migula, 1900, p. 586.

*Published by permission of the Director of the Wisconsin Agricultural Experiment

Station.

220 Wisconsin Academy of Sciences, Arts, and Letters.

Bacillus Septimus (plugge)

No. VII. PLUGGE, Zeitsch. f. Hygiene 17:294. 1894.

B. plicatus CHESTER^ 1901, p. 275.

B. lactis No. 7 (plugge), Swithinbank & Newman, 1903, p. 423.

Note: Plicatus is not tenable because this name was given to a bacillus

by Deetjen in 1890.

Bacillus sextus (plugge)

No. VI. PLtiGGE, Zeitsch. f. Hygiene 17: 294. 1894.

B. lactis No. 6 (plugge), Swithinbank & Newman, 1903,

p. 423.

Bacillus tertius (plugge)

No. III. PLtiGGE, Zeitsch. f. Hygiene 17: 272. 1894.

B. lactis No. 3 (plugge), Swithinbank & Newman, 1903, p. 422.

Bacterium quintus (plugge)

No. V. PLtiGGB, Zeitsch. f. Hygiene 17: 272. 1894.

B. lactis No. 5 (plugge), Swithinbank & Newman, 1903,

p. 423.

New Combinations

Bacterium circulans ( Jordan)

B. circulans Jordan, Keport Mass. S. Board of Health, 1890,

p. 831.'

B. circulans II. conn, 1899, p. 58.

B. lactis circulans I. & II. conn, esten & stocking, 1906,

p. 174.

Note: Franklands’ description of spores is probably an error.

Bacterium cochleatus (conn, esten & stocking)

B. lactis cochleatus conn, esten & stocking, 1906, p. 181.

Bacterium liodermos (plugge)

B. liodermos plugge. Die Mikroorganismen, 1886, (II. Aufl.),

p. 323.

No. X. PLtiGGE, Zeitsch, f. Hygiene 17 : 293. 1894.

Gummibacillus loeffler, Migula, 1900, p. 577.

B. cremoris Chester, 1901, p. 274.

B. lactis No. 10 (plugge), Swithinbank & Newman, 1903,

p. 423.

Frost & Noland — Names of Certain Milk Bacteria. 221

Bacterium robertii (conn, esten & stocking)

B. lactis Bohertii conn, esten & stocking, 1906, p. 182.

Note: Description suggests Bact. vermiculosum (Zimmerman) Chester,

1901, p. 157, or B. virmiculosus Zimmerman, Die Bak. Nutz. u. Trinkwasser,

1890, p. 40.

Bacterium tenuis (duclaux)

Ty. tenius duclaux, Le Lait, Paris, 1887.

Ty. scaber duclaux, Le Lait, Paris, 1887.

B. tenuis trevisan, 1889.

B. scaber (duclaux) migula, 1900, p. 586.

B. lactis tenuis conn, esten & stocking, 1906, p. 176.

Note: We follow Conn who seems to regard Ty. scalier and tenuis as

identical.

Bacterium visco-symbioticum (buchanan & hammer)

B. visco-symbioticum buchanan & hammer, 1915, p. 261.

Chromobacterium ianthinum (zopf)

Bacterium ianthinum zopf, 1885, Spaltzpilze, p. 68.

Bact. ianthinum (zopf). Com. S. A. B. 1920, p. 222.

B. violaceus mac£:, 1887, Ann. d’Hyg. publ. et de Med. leg.

XVII.

B. violaceus frankland, 1889, Zeitsch. f. Hygiene, p. 394.

B. violaceus-laurentius Jordan, 1890, State Bd. of Health,

Mass., p. 838.

B. janthinus zimmerman, 1890, Die Bakt. nnser Trink. und

Nutzwasser, I. Kuhe, p. 36.

Ps. ianthina (zopf) mig., 1900, Migula, p. 94.

Ps. ianthina (zopf) Chester, 1901, Chester, p. 317.

Bact. violaceum (j. schroter), 1901, Lehmann and Neumann,

p. 277.

B. lividus (PLtiGGE and proskauer), 1887, Zeitsch. f. Hygiene

2: 463.

B. membranaceus amethystinus eisenberg, 1891, p. 421.

B. membranaceus amethystinus mobilis germano, 1892, Cen-

tralb. f. Bakt. 12 :516.

Erythrobacillus aurantiacus (frankland)

B. aurantiacus frankland, 1894, p. 449.

Note: Gr. and P. Frankland described but did not name it in an earlier

communication, Uber einige typische Organism im Wasser und im Boden,

Zeitsehr. f. Hygiene, Bd. VI., 1889, p. 390, also Frankland, 1894, p. 449.

222 Wisconsin Academy of Sciences, Arts, and Letters.

Erythrobacillus diffusus (conn, esten & stocking)

B. lactis diffusus conn, esten & stocking, 1906, p. 181.

Erythrobacillus aureus (frankland)

B. aureus frankland, 1894, p. 448 (See Note below) .

B. aureus lactis II. conn, 1899, (No. 100), p. 39.

Bact. aureum (frankland) mig., 1900, p. 480.

Bact. lactis aureum II. conn, esten & stocking, 1906, p. 130.

Note: The morphology of the form described by Frankland does not

fully agree with that described by Conn, Esten & Stocking.

Lactobacillus flocculus (conn, esten & stocking)

Bact. Lactis flocculus conn, esten & stocking, 1906, p. 149.

Lactobacillus gorinii (conn, esten & stocking)

Bact. C. MULLER, Arch. Hyg. 67 :127.

Bact. lactis Gorinii conn, esten & stocking, 1906, p. 148.

Lactobacillus healii (buck an an & hammer)

Bact. Healii buchanan & hammer, 1915, p. 249.

Lactobacillus magnus (conn, esten & stocking)

Bact. lactis magnum conn, esten & stocking, 1906, p. 149.

Lactobacillus surgeri (dornic & daire)

Bacillus surgeri dornic & daire, 1907, Bull. mens, de Toffice

de reseignements agricoles, 6:146.

Bact. surgeri (dornic & daire) buchanan and hammer, 1915,

p. 254.

References

Buchanan and Hammer. 1915. Slimy and rop}'' milk. Eesearch Bull. Iowa

State Agr. Exp. Sta.

Chester. 1901. Manual of determinative bacteriology. MacMillan Co., N. Y.

Committee. 1920. Families and genera of the bacteria. Winslow, Broad-

hurst, Buchanan, Krumwiede, Rogers, and Smith. Jour. Bact. 5:191.

Conn. 1899. 12th Annual Report Storrs Agric. Exp. Station, Storrs, Conn.

Conn, Esten and Stocking. 1906. 18th Annual Report Storrs Agr. Exp. Sta.

Frankland. 1894. Micro-organisms in water. Longmans, Green & Co.,

N. Y.

Migula. 1900. System der Bakterien. Gustav Fischer, Jena.

Swithinbank and Newman. 1903. Bacteriology of milk. John Murray,

London.

Trevisan. 1889. Schizomycetaceae in Sylloge Fungorum by A. P. Saccardo.

Vol. III.

THE CHARACTERISTICS OF CERTAIN FECAL BACTERIA

AS SHOWN BY THE LITTLE PLATE METHOD.

Ola E. Johnston and William D. Frost

(From the Department of Agricultural Bacteriology, University of Wisconsin,

Madison, Wisconsin.)

In another paper we propose to discuss the ‘^Use of the Little

Plate Method for the Bacteriological Analysis of Feces.” Here

we describe and illustrate by means of photomicrographs the most

commonly occurring colonies in the feces of infants, guinea pigs,

puppies and rats.

Infants. The ages of the infants studied ranged from three

days to eleven months. They were all breast fed except one.

Little plates made from the feces of these infants always showed

the presence of many diplococci. These organisms varied in

diameter from 0.6-0. 8 microns, although occasionally smaller ones

were found even down to 0.3 microns. Frequently these units

grouped themselves in short chains. Some plates contained almost

pure cultures of diplococci.

Usually colonies of staphylococci occurred in about the same

numbers as the diplococci. Besides the irregular grouping, a few

in tetrads and chains were found. The individual cells were about

the same size as the streptococci or perhaps more frequently a

little smaller.

With one exception all of the infant feces showed the presence

of as many rods as diplococci or staphylococci. These bacilli were

either Gram positive spore-bearers or Gram negative non-spore

bearing rods. They were from 0.5 to 0.8 microns wide and from

1.0 to 3.5 microns long but most frequently about 1.5 microns.

Comparing these colonies with the colonies obtained by making

little plates of pure cultures we have identified the following:

B. acidophilus y B.alholactis, B.bifidus, B.cereuSy Staph.albus{°l),

and Strep.lacticus. See figs. 1, 2, 3, 6, 11, 12, 13, 15, 16 and 19.

Guinea pigs. The fecal flora of fourteen guinea pigs was

studied by this method.

224 Wisconsin Academy of Sciences, Arts, and Letters.

A moderate number of colonies of diplococci was always found.

The individual cells varied in diameter from 0.5 to 0.8 microns

but were usually about 0.6 microns.

A few streptococci and a few staphylococci were found also.

Nearly as many colonies of small rods was found as diplococci.

These colonies had dense centers and open edges where the individ¬

uals were grouped in pairs with a slight tendency to form short

chains. The cells varied in width from 0.5 to 0.7 microns with

a few as wide as 1.0 micron. In length they ranged from 1.0 to

3.5 microns, averaging about 2.0.

Several of the guinea pigs showed large rods which were ar¬

ranged in extremely long chains. These lay parallel or interlacing

and most generally radiating from the dense center of the colony.

About one-third of these animals showed the presence of smaller

rods which grew into shorter chains than the foregoing.

Spores were quite abundant.

The dejecta of several scurvy guinea pigs was examined. The

bacteria varied little here from those found in the normal animals.

Except that the types were fewer and the colonies larger and

more compact than usual.

By comparison with little plates of pure cultures the following

species were recognized: Staph.alhus{^), Strep.lacticus, B.coli,

and B.subtilis. See figs. 9, 10, and 17.

Puppies. The fecal bacteria of the three puppies studied con¬

sisted of many diplococcus colonies, a moderate number of strep¬

tococcus colonies and a few staphylococcus colonies. Many large

bacillus colonies and a few colonies of very short bacilli.

Rats. There were many colonies of diplococci, streptococci

and staphylococci. Also many large bacilli of which some were ar¬

ranged in spreading interlacing threads and others isolated or in

pairs.

Some of the colonies were Gram positive and spore bearing and

others Gram negative and non-sporulating.

The organisms found are believed to be: Staph.alhusi^l),

Strep.lacticus, B. acidophilus, B.hifidus, B.mesentericus. See figs.

4, 5, 7, 8, 14, 18, 20, and 21.

Photomicrographs of some of the most frequently encountered colonies

are given in the accompanying plate (No. IX).

Colonies from the stools of infants are reproduced in figures 1, 2, 3, 6, 11,

12, 13, 15, 16, and 19. Those from guinea pigs in 9, 10, and 17; and from

rats in 4, 5, 7, 8, 14, 18, 20, and 21.

TRANS. WIS. ACAD., VOL. XXI

%%W'20

ON THE NATUKE OF DISEASE RESISTANCE IN PLANTS

J. C. Walker

Introduction

Since the pioneer work of DeBary (26) and Ward (87) upon

the nature of parasitism, numerous contributions to our knowledge

of the subject have been made. On the whole, however, we are

still very much in the dark as to the nature of the complex rela¬

tions which exist between the parasite and the host. Duggar (28)

in 1911 remarked that, ‘‘in general, the physiology of penetration

is poorly understood and it will reward investigation. . . .

There is now at hand ample material for a thoroughgoing study

of some of the factors governing resistance and susceptibility. The

problem is doubtless extremely difficult, but it is believed by

some the work may yield results. ... At present there is no

information respecting the cause of this difference in behavior as

regards resistance. ’ ’ In connection with the writer ’s studies upon

the nature of disease resistance in the onion (86), a review of

previous work upon the general subject was found necessary. The

results of this survey of the literature are presented at this time.

Types of Disease Resistance

Orton (61), followed by Freeman (32) and Butler (16), calls

attention to the distinction between avoidance of disease, endur¬

ance of disease, and true resistance to disease. It is the last of

these three classes with which we are concerned in this paper.

We shall include in this class, cases where through some inherent

quality in its composition, the plant is capable of successfully

resisting, to a greater or less extent, the attack of a given parasite.

A survey of the studied cases of disease resistance in plants

brings out at once the fact that all degrees are to be found, varying

from complete immunity to a high degree of susceptibility. We

have, for instance, in certain varieties of the potato complete

immunity to the wart disease ( Synchytrium endohioticum (Schil.)

15

226 Wisconsin Academy of Sciences, Arts, and Letters.

Perc. (60). On the other hand, in the case of cabbage yellows

(45) (Fusarium conglutinans Woll.) the resistant Wisconsin

Hollander variety under extreme conditions (high soil tempera¬

ture) shows a high percentage of plants slightly affected with the

disease. However, such plants resist the parasite so successfully

that they continue their growth, and with the return of somewhat

cooler conditions all symptoms of the disease disappear and the

plants mature normally. In contrast to this, plants of the common

susceptible Hollander variety of cabbage may entirely succumb

to the disease during the extreme conditions of high soil tempera¬

ture. We thus have in the case of Wisconsin Hollander not a

completely immune variety by any means, but unquestionably one

which may be said to have a high degree of disease resistance.

Moreover, as pointed out by Tisdale (78), this is a case of resis¬

tance acquired during the growth of the plant, since he has shown

that very young seedlings of the Wisconsin Hollander variety are

as susceptible as those of the common Hollander variety.

Kelation of Environment to Kesistance

The effect of environment upon resistance has been given only

occasional attention by investigators. Vavilov (82) points out

several cases where certain races of plants were equally resistant

to given parasites when tried out under a variety of environments

and in various parts of the world. However, we are not justified

in concluding that in other cases environment does not have its

effects.

Biffen (7) and Spinks (70) point out a distinct influence of

various fertilizers upon the resistance of wheat to yellow rust

(Puccinia glumarum (Schm.) Erikss. & Henn.). Stakman and

Aamodt (73), on the other hand, found that in the case of wheat

rust (Puccinia graminis tritici Erikss. & Henn.) “the amount of

rust was not changed directly by any fertilizer or combination of

fertilizers, although date of maturity, degree of lodging, crinkling,

shrivelling of seed, percentage of yellow-berry, and yield were

affected profoundly.^’ Tisdale (79) states that a strain of flax,

resistant to wilt (Fusarium Uni Boll.) under normal field environ¬

ment, showed a marked increase in susceptibility when grown at

high greenhouse temperatures. The value of results recorded on

the behavior of so-called resistant varieties in localities other than

those in which they originated is often limited by the fact that there

Walker — Nature of Disease Resistance* in Plants.

227

often exist biologic strains of the parasite possessing distinct

infective properties. This is especially true with the rusts where

the existence of distinct biologic strains of Puccinia graminis tritici

Erikss. & Henn. (74, 76), Puccinia graminis avenae Erikss. &

Henn. (75), Puccinia graminis secalis Erikss. & Henn. (47), and

Puccinia triticina Erikss. (51) has already been demonstrated.

Kanred, a wheat resistant to Puccinia graminis tritici in Kansas,

is not highly resistant in South Dakota nor in Minnesota. The

dilference in behavior is not attributed to any material difference

in the host plant under the three environments, but to a difference

in infective properties of the biologic strains of the parasite in the

respective regions (54). In the case of the bean anthraenose

organism ( Collet otrichum lindemuthianum (Sacc. & Magn.) B.

& C.) two biologic strains have been differentiated which behave

quite differently in their infective properties on individual varie¬

ties of bean (5). Much more study is necessary on the effect of

environing conditions upon parasitism and the expression of re¬

sistance over a wide range of cases.

Genetic Behavior of Resistance

The hereditary nature of resistance has been shown to be a

common phenomenon ; its genetic behavior, however, from the

limited results at hand, does not appear to follow any single

genetic law. Chief limiting factors in the advancement of our

knowledge of this phase of the subject have been the necessity of

assuming an arbitrary division between resistance and susceptibil¬

ity in analyzing the behavior of any group of plants, the present

imperfect understanding of the effect of environmental factors

upon the expression of resistant characters, and the lack of recog¬

nition, until recently, of the possible existence in many cases

of several distinct biologic strains of the parasite. Biffen (7)

found resistance in the case of yellow rust of wheat (Puccinia

glumarum) to be recessive and determined by a single factor.

Nilsson-Ehle (57), on the other hand, would explain the genetic

behavior of resistance to yellow rust in his crosses of wheat on

the basis of multiple factors. It is entirely possible that the

variance in results in these two cases is due to the existence of

different biologic strains in the two localities or to differences in

environment. Recent work with black stem rust of wheat (Puc¬

cinia graminis tritici) (1, 55) and of oat (Puccinia graminis

228 Wisconsin Academy of Sciences, Arts, and Letters.

avenae) (34) indicates that resistance to a given biologic strain is

determined by a single dominant factor. Crosses between varieties

of bean resistant and susceptible to anthracnose (Colletotrichum

lindemuthianum ) show that there is only a single factor difference

between the two when a single strain of the parasite is considered

(14, 49). Where two strains are involved a two factor difference

is indicated (50). In both cases resistance is dominant over

susceptibility. With two other diseases of bean, mosaic and root

rot (Fusarium martii A. & W. var. phaseoli Burk.), susceptibility

is at least partially dominant and resistance must be explained

on the basis of more than one factor (50). Vavilov (81) in study¬

ing resistance of wheat to mildew {Erysiphe graminis DC.) found

that from crosses between the immune Persian wheat (Triticum

vulgar e var. fuliginosum Al.) and common susceptible bread wheat

varieties, P^ hybrids were secured which were immune to the disease.

Klaphaak and Bartlett (46) find that immunity to powdery

mildew {Erysiphe polygoni DC.) in certain species of Oenothera is

determined by a single factor, which is dominant. In the case

of flax wilt (Fusarium Uni) Tisdale (79) finds the behavior of re¬

sistance more readily explained on the basis of multiple factors.

Gaines (33) reports there are different types of resistance to bunt

(Tilletia tritici (Beij) Wint.) in different varieties of wheat and

that in any particular case the resistant quality is composed of

multiple factors. Johnson (40) finds that inheritance of resistance

in tobacco to Thielavia hasicola (B. & Br.) Zopf is best explained

by a multiple-factor hypothesis.

Classes of True Resistance

The causes of true resistance in plants have been grouped by

Vavilov (82) into two classes: (1) mechanical or passive im¬

munity and (2) physiological or active immunity. The first class

includes those cases in which resistance depends upon certain

mechanical differences in structure or habit of growth of the

plant. The second class includes those cases in which resistance

is due to inherent physiological qualities of the host cells which are

sufficiently antagonistic to check the parasite. While this classi¬

fication is convenient, it is obviously unwise at our present state

of meagre knowledge of the subject to attempt to apply it too

rigidly.

Walker — Nature of Disease Resistance in Plants.

229

Cases of Eesistance Due to External Structure of Host

In a few instances disease resistance has been attributed to purely

external differences in mechanical structure of the host plant.

Hairiness of leaves and open habit of growth in certain varieties

of potato are cited by Stuart (77) and by Appel (3) as facilitating

more rapid evaporation of rain drops and hence as reducing lia¬

bility to attack by Phytophthora infestans (Mont.) DeBary.

Darnell-Smith (25) found a correlation between resistance to

bunt (Tilletia tritici) in certain varieties of wheat and lack of

hair on the terminal ends of kernels. Appel (3) noted that certain

varieties of raspberries which were covered by a thick, blue, waxy

layer remained free from attack by Coniothyrium, while other

varieties were severely attacked. He suggests that the waxy layer

may possibly influence infection by preventing penetration, or

causing drops of water to run off. Freeman (32) noted higher

resistance to stem rust (Puccinia graminis Pers.) in barley grown

on alkali soils where the amount of bloom’’ was increased.

The number and structure of stomata have been regarded in

some cases as factors in disease resistance. In a study of numer¬

ous varieties of wheat in relation to rust (Puccinia graminis), Cobb

(17) notes that in general the stomata were smaller and more

numerous in the resistant than in the susceptible forms, and to

these and certain other anatomical differences he attributes the

cause of resistance. Eriksson and Henning (30), the first to test

this theory, did not uphold it, and the more accurate studies of

Ward (88) upon brome rust (Puccinia dispersa Erikss.) failed to

confirm it'. Norton (59) points out a correlation between small

stomata and resistance to rust (Puccinia asparagi DC.) in the case

of the resistant Martha Washington variety of asparagus, but the

actual relation of this factor to penetration and infection was not

studied. Allen (2) reports that only a very small percentage of the

germ tubes of Puccinia graminis tritici enter the stomata of the

highly resistant variety of wheat, Kanred, and points out that

the stomatal slit of this variety is smaller and more slender than

that of the very susceptible variety, Baart. Appel (3) cites a case

in which resistance in “some Kemontant carnations is due to the

form of the stomata, which makes it impossible for the hyphae

to penetrate them.” Pool and McKay (63) point out that the

immature leaves of beet are nearly immune to infection by Cer-

cospora heticola Sacc. because the stomata are too small to permit

230 Wisconsin Academy of Sciences, Arts, and Letters.

penetration by the fungns hyphae. Valleau (80) suggests that

resistance to Sclerotinia cinerea (Bon.) Wor. in certain types of

plums may be due to the fact that the stomata early become plugged

with small parenchymatous cells which impede penetration.

The possible exclusion or retardation of parasites by means of

unusual thickness of cuticle or by outer layers of corky cells has

been suggested in a number of cases. Cobb (17) suggests that the

thicker cuticle which is common in rust resistant varieties of wheat

may contribute to resistance by preventing the maturation of the

rust sori. Later work, such as that of Ward (88), indicates that

this is not the important factor in rust resistance. An interesting

case of correlation between thickness of cuticle in the tomato fruit

and its resistance to Macrosporium tomato Cooke has recently

been described by Rosenbaum and Sando (66). Young immature

fruits are highly susceptible, while old fruits are resistant to attack.

This difference does not appear to be due to changes within the

host, since infection may readily be obtained in old fruits by

first injuring the skin. There are no natural openings in the skin

and the cuticle becomes thicker as the fruit matures. Moreover,

the outer layer is shown to become gradually more resistant with

age to mechanical puncture with a needle, and the authors suggest

that resistance in the older fruits may be due to the ability of the

thicker cuticle to resist puncture by the mycelium of the para¬

site. In discussing the pink disease of rubber (Corticium sal-

monicolor B. & Br.) in India, Butler (16) states that the attack

is most marked on the shady side of the tree where the thinner

cork offers less resistance to invasion. Appel (3) points out the

importance of corky layers as a factor in resistance to certain

parasites. In the ease of the sweet potato, Weimer and Harter

(91) have shown that wound cork layers may retard infection and

that suberization of cell walls at the surface of the wound is often

sufficient to prevent the entrance of microorganisms.

Cases of Resistance Due to Internal Character of the Host

In the foregoing cases, external differences in plant structure

have been considered the determining factors in disease resistance.

There is a much larger number of cases in which resistance is

seemingly due to internal causes which have been sought' either

in the cell membranes or cell contents. The pioneer work of

DeBary (26) upon Sclerotinia demonstrated the secretion of a

Walker — Nature of Disease Resistance in Plants.

231

substance by the parasite which killed the host cells by diffusing

in advance of the hyphae. As to the nature of this action he was

not sure, but he considered the cell wall solution enzymic in

character, and suggested the possible effects of a soluble oxalate.

Ward (87) in his study of the Botrytis disease of lily likewise

demonstrated the production of a wall-dissolving enzyme at the

tips of the hyphae. Nordhausen (58) working with Botrytis

cinerea Pers. also described its enzymic activity but in addition

a toxic action of the fungus due possibly to oxalic acid. Smith (68)

took the extreme position of attributing all of the action of Botrytis

upon the host to oxalic acid. Jones (43), working with the soft

rot bacillus (B. carotovorus Jones), ascribed the action of the para*

site to a secreted enzyme, pectinase, and gave little importance

to oxalic acid. Brown (9, 10, 11) improved greatly on methods

previously used. He distinguished a macerating and a lethal effect

of Botrytis cinerea upon the host. The former precedes the latter

and is due to a cytolytic enzyme produced largely at the actively

growing tips of the fungus hyphae. The nature of the lethal

principle was not determined. Brown points out that there are

chemical differences between cell walls and that a more thorough

study of hemicellulose and pectins is needed. The significance at¬

tributed by earlier writers to oxalic acid or a soluble oxalate is

definitely refuted.

In these early investigations the first stages of penetration were

not made clear. Biisgen (15) did not consider that the parasite

penetrated the cuticle by mechanical means alone. Miyoshi (56),

however, showed that Botrytis cinerea was capable of penetrating

paper, collodion, and other substances by mechanical pressure.

More recent researches with the same fungus by Blackman and

Welsford (8) and Brown (10) go to show that it does not secrete

a cutin-dissolving enzyme but penetrates the cuticle by mechanical

means alone. Further instances of apparent mechanical penetra¬

tion have since been demonstrated by Dey (27) in the case of bean

anthracnose, by Waterhouse (90) in the case of Puccinia gra-

minis on barberry and by Curtis (24) in the case of invasion of

potato by the zoospores of the wart organism.

Importance of Cell Membranes

Hawkins and Harvey (39) give evidence to support their theory

that in certain varieties of potato tubers, resistance to invasion

by Pythium deharyanum Hesse is due to resistance of the cell

232 Wisconsin Academy of Sciences ^ ArtSy and Letters,

walls to mechanical puncture by the fungus hyphae. Tisdale (79)

describes the formation of a layer of suberized cell walls in the

cortex of resistant flax plants which checks further advance of the

wilt fungus, Fusarium Uni. He points out, however, that the

actual resistant quality may possibly be contained in certain prop¬

erties of the protoplasm of the normal cortical cells which retard

the progress of the parasite during the suberization of the under¬

lying cells, the latter being a natural host reaction. Valleau (80)

considers it very possible that the slow development of Sclerotinia

cinerea in resistant plums is due to a slight difference in the com¬

position of the middle lamella which makes the latter less easily

soluble in the secretions of the fungus. Further work by Willaman

(93, 94) shows a higher crude fiber content in resistant than in

susceptible plums.

The Nature of Rust Resistance

The nature of resistance to rusts has been studied by a number

of investigators. The work of Cobb (17), stressing the importance

of the anatomical features of the host, has already been cited.

After a careful and comprehensive study of the question, Ward

(88, 89) failed to find any correlation between external host fea¬

tures and infection. He showed (89) that in the case of wheat

susceptible to Puccinia glumarum the hyphae of the latter “are

typically stout, branched, and contain hundreds of nuclei. They

also form numerous haustoria, and the attacked cells show no

evident signs of injury to the chlorophyll-corpuscles or nuclei

until a late stage of growth.’’ In the case of highly resistant

wheat, however, the substomatal chamber is invaded normally, but

when the hyphae come to invade the cells they “show evident

signs of degeneration in all respects and we conclude, from com¬

parison with experimentally starved hyphae, that they are under¬

going death-changes owing to one of two events, viz., they are either

starving for want of food supplies or they are being poisoned.”

The same investigator (88) described earlier a similar condition

existing between Puccinia dispersa and susceptible and highly

resistant varieties of bromes. He (89) sums up his conclusions

on the subject as follows :

In other words, infection, and resistance to infection depend on the

power of the fungus-protoplasm to overcome the resistance of the cells of

the host by means of enzymes or toxins; and, reciprocally, on that of the

Walker — Nature of Disease Resistance in Plants. 233:

protoplasm of the cells of the host to form anti-bodies which destroy such

enzymes or toxins, or to excrete chemotactic substances which repel or at¬

tract the fungus-protoplasm.

Other work on the relation of rust fungi to their hosts has been

carried on by Gibson (36) with Puccinia (TJredo) chrysanthemi

Roze. on chrysanthemum, Marryat (52) with Puccinia glumarum

on wheat, by Stakman (71, 72) with Puccinia graminis on wheat,

by Reed and Crabill (65) with Gymnosporangium juniperi-virgin-

ianae Schw. on apple. In all cases the parasites were found to

enter normally the substomatal chambers of both susceptible and

highly resistant varieties of their respective hosts, but the differ¬

ence in resistance lay in the reaction between the fungus hyphae

and the mesophyll cells about the substomatal chamber. Giddings

(37) describes a gradual development of resistance to rust (Gym¬

nosporangium juniperi-virginianae) in the leaves of York Imperial,

a susceptible variety of apple. He believes that this acquired im¬

munity is due primarily to food factors and secondarily to dif¬

ferences in the composition of the cell wall.

The recent contribution by Miss Allen (2) advances considerably

our knowledge of the nature of rust resistance. She has made

a careful eytological study of infection of Baart, a very susceptible

variety of wheat, and Kanred, a highly resistant variety, to a certain

biological strain of Puccinia graminis tritici. We quote from her

summary as follows:

The germination of the spores and the formation of the appressoria on

the stomata take place in the same way in the susceptible and immune hosts.

In Baart the fungus enters freely and grows rapidly. In Kanred, under

greenhouse conditions, only a few of the fungi pass through the stomata;

the rest remain outside until they shrivel and die.

In a congenial host, numerous haustoria are formed. A slender-growing

hypha strikes a host cell, swells at the tip, its pair of nuclei divide, and

a septum is formed, marking off a short terminal cell. This haustorium

mother cell is closely appressed to the host cell, forms a fine pore through

its wall and the host wall, and its contents, including both the nuclei,

which have decreased in size, and the cytoplasm, now pass in, forming the

haustorium. The osmotic membrane of the host appears to be invaginated

by the haustorium, but apparently is still intact.

In Kanred the process is similar until a small haustorium is formed,

which, either by its presence, or, as is more likely, by secreting some

substance in the host cell, sets up chemical reactions within that cell,

causing its collapse and death. The further diffusion of toxic substances

into healthy host tissues is cheeked by the formation of thickened contact

walls. One or more of the substances formed in the host cell diffuse into

234 Wisconsin Academy of Sciences, Arts, and Letters.

the haustorium, killing it, and causing collapse of the mother cell and the

death and plasmolysis of the hypha back of it for some distance. If this

reaction is rapid, the haustorium is destroyed while still very small; if

more sluggish, a full-grown haustorium may be formed and some nourish¬

ment for further growth be extracted by the fungus.

She thus points out that the resistant variety Kanred possesses

three means of defense against this strain of rust, namely, the

stomata which exclude most of the fungi, the heavy contact walls

adjoining the attacked cells which are interpreted as preventing

the diffusion of toxic substances to uninvaded cells, and ‘‘a true

immunity.’’ In connection with the last the starvation theory

discussed by Ward (89) and his students (36, 52, 70) does not

seem tenable since the fungus hyphae appear to grow well in

the intercellular spaces as long as the host cell is not penetrated.

Upon invasion of the latter, there is evidence that the host cell

undergoes chemical changes due to the invasion by the parasite

and that, moreover, some substance diffuses into the fungus haustor¬

ium resulting in its death. That there is irregularity in the

balance of forces of interaction between parasite and host is

shown by the fact that in some cases the one collapses first while

in other cases, the other dies first.

Preliminary studies of the reaction of Kanred to another bio¬

logic strain of Puccinia graminis tritici to which it is known to be

less ' resistant, indicate that a higher percentage of appressoria

enter the stomata. It will be interesting to learn the results of

further study on the interactions between the host and this second

strain of the parasite.

Chemotropism as a Factor in Resistance

Massee (53) would explain the susceptibility or resistance of a.

plant by the presence or absence of a positively chemotropic sub¬

stance in the host cells. The evidence is not convincing, however.

Numerous cases have been recorded where fungi penetrated plants

very resistant to their attack. Gibson (36), for instance, found

that the germ tubes of a number of rust fungi readily entered a

wide range of plants other than their respective hosts, but no

further development of the fungus occurred. Wiltshire (95)

found that the pear scab fungus (Venturia pirina) would invade

apple fruit and that the apple scab fungus (Venturia inaequalis)

would invade the pear, but each parasite was capable of producing

the disease only upon its respective host. Tisdale (79) showed

Walker — Nature of Disease Resistance in Plants.

235

that under certain conditions the cabbage yellows fungus (Fusar-

ium conglntinans) would invade the root hairs of flax, but was

incapable of progressing farther and producing a wilt similar to

that caused by Fusarium Uni. In a comparative study of two

closely related organisms causing the leaf spots of alfalfa and red

clover, respectively, Jones (41) showed that when sown under

proper conditions upon clover leaves, the ascospores of Pseudo-

peziza medicaginis (Lib.) Sacc. germinated and the hyphae pene¬

trated the epidermal cells, but the progress of the fungus was

checked at this point; likewise when ascospores of the red clover

organism, Pseudopeziza trifolii (Bernh.) FcL, were sown upon

alfalfa leaves, the germ tube penetrated but did not advance beyond

the epidermal cell. Salmon (67) noted that the haustoria of the

wheat mildew (Erysiphe graminis) penetrated the epidermal cells

of the barley leaf, but that they eventually shriveled and died with¬

out producing the characteristic disease symptoms. Explanation

of the cases just mentioned on the basis of the absence of positively

chemotropie substances in the host cell is unsatisfactory, since

several other possible explanations might be offered, such as the

presence of inhibitive substances in the host cell, or the absence

in the parasite of proper enzymes or toxins to bring about the

chemical or physical changes in the host cell necessary to provide

food for the growth of the invader.

Osmotic Pressure as a Factor in Resistance

From his earlier experimental studies upon phanerogamic para¬

sites, McDougal (48) concluded that an osmotic pressure of the

parasite higher than that of the host was essential. In the light

of more recent researches he believes that osmotic pressure may be

of minor importance in the establishment of the haustorium, and

that absorption by imbibition and the force of expansion of the

invading protoplast are important factors at this early stage. In

this connection the work of Hawkins (38) is significant. Using

two potato-rotting fungi, and one strawberry-decaying fungus,

he found that they would all grow on solutions of glucose, sucrose,

potassium nitrate or calcium nitrate at diffusion tensions much

higher than the total diffusion tensions of the dissolved substances

in the juices of their respective host plants. Vavilov (82), after

examining many species and varieties, was unable to establish

any correlation between osmotic pressure and resistance.

236 Wisconsin Academy of Sciences, Arts, and Letters.

Cell Sap Acidity as a Factor in Resistance

A correlation between resistance and a higher acidity of the cell

sap of the host tissues has been pointed out by several investigators.

Averna-Sacca (4) reports a higher acidity in grapes resistant to

Oidium and Peronospora, and Comes (18, 19) reports a similar

correlation for a variety of wheat (Rieti) resistant to rust. On

the other hand, Vavilov (82) found no connection between the

acidity of cell sap of many varieties of oats, wheat, and roses

and their resistance to rusts and mildew. No such correlation was

found in the potato by Jones and co-workers (44) in the case of late

blight (PhytopJithora infestans), by Hawkins and Harvey (39) in

the case of leak (Pythium debaryanum), nor by Weiss and

Harvey (92) in the case of black wart (Sychitrium endobio-

ticum). Gardner and Kendrick (35) in their work upon a tomato

fruit spot caused by Bacterium exitiosum G. & K. find a corre¬

lation between the hydrogen ion concentration of the plant tissues

and their resistance to infection. The organism did not grow in

culture media more acid than a Ph value of 5. Examination of

various tomato plant parts yielded Ph values as follows : seedlings

and leaves, 6.3 to 6.5; green fruits, 5 to 5.4; ripening and mature

fruits, about 4.6. Seedlings, leaves, and green fruits were very

susceptible to the disease, but inoculations of ripe fruits were

usually unsuccessful. Aside from the last instance, we have

as yet very little convincing evidence that cell sap acidity is an

important factor in disease resistance.

Tannin as a Factor in Resistance

The relation of tannin to the growth of a number of fungi was

studied by Cook and Taubenhaus (21). They found that in

general when tannin was added in increasing amounts to a favor¬

able medium, germination was inhibited and finally spores were

killed. Considerable variation in the reaction of different fungi

was noted, and in general, parasitic fungi were less resistant to

the toxicity of tannin than saprophytic fungi. Cook and coworkers

(20) later claimed that tannin as such does not exist in the host

cell, except in small amounts. There does exist a polyatomic

phenol, which, upon injury to the cell, results in the formation

of a tannin or tannin-like substance. They point out that condi¬

tions for such a reaction prevail in normal immature pomaceous

fruits which are injured by the invading hyphae. A germicidal

Walker — Nature of Disease Resistance in Plants.

237

fluid containing the soluble tannin is thus formed; on the basis

of the germicidal action of this fluid, the resistance of such fruits

to parasitic attack is explained. Cook and Taubenhaus (22) in a

later paper point out that many fruits lose their power of resistance

very soon after removal from the plant. This loss of resistance is

proportionate to the reduced activity of the enzyme. They extended

their studies to the vegetable acids and found the toxicity of the

latter to vary with the organisms 'used. Tannic acid was the most

toxic, but the character and the true importance of this substance

within the living plant remains to be determined by future investi¬

gations. Valleau (80) found no correlation between tannin content

of plums and their resistance to Sclerotinia cinerea. The investi¬

gations upon tannin and organic acids in relation to resistance,

while suggestive, are still inconclusive.

Anthocyans and Flavones in Relation to Resistance

The pigments of the anthocyan class have been suggested in

several instances as substances contributing toward disease resist¬

ance, but in most cases the evidence is only observational and

is not supported by experimental investigation. Sorauer (69)

noted that red potatoes are in general more resistant than white.

Jones (42) says that many potato experts, American and Euro¬

pean, regard red, rough-skinned varieties of potatoes as less

liable to rot than thin, white-skinned potatoes. He points out,

however, that there is abundant evidence of high disease resistance

coupled with a thin white skin. Voges (83) considered red color

in apple fruits a protection against scab but the large amount of

data to be found in literature does not bear out this statement.

Comes (19) attributed certain cases of resistance to the occurrence

of anthocyan but his evidence is not convincing. In this con¬

nection, it is interesting to note the work of Cook and Wilson (23)

who studied the effect of ^‘commercial” tannin and extracts from

chestnut bark upon the growth of the chestnut blight organism,

Endothia parasitica (Murr.) And. & And. Results with “com¬

mercial” tannin were of the same general nature as those secured

with other fungi by Cook and Taubenhaus (21, 22) noted above.

Since “commercial” tannin was variable in composition and since

it contained a certain percentage of impurities, extracts from

chestnut bark were also used. Through the cooperation of Kerr,

three extracts were made: “1-X”, described as the water soluble

238 Wisconsin Academy of Sciences, Arts, and Letters.

tannin, insoluble in alcohol; “2-X”, described as similar in its

reactions to except that it is soluble both in water and in

alcohol; and “3-X”, described as the ‘‘coloring matter of the

bark^’ . . . which “is estimated as tannin in bark analysis,”

but “its real nature is unknown.” Extracts “1-X” and “2-X”

were less toxic than “commercial” tannin while “3-X”, the

coloring matter, was very toxic. They point out further a state¬

ment from Kerr to the effect that “chestnut trees of northern

growth, say on a line north of the southern boundary of Pennsyl¬

vania, contain very materially less coloring matter than the growth

south of it, and, as we all know, the wood in the latitude referred

to seems to have been more susceptible than that further south.”

The importance of the coloring matter as an inhibitive substance is

thus suggested in this instance. Promme and Wingard (31) in a

study of varietal susceptibility of beans to rust (Uromyces appen-

diculatus (Pers.) Lev.) found that all varieties with solid red or

red mottled seed were resistant, while those with white seed as

a class were more susceptible than those of any other color. In the

case of onion smudge ( Collet otrichum circinans (Berk.) Vogl.), a

bulb rot. Walker (84, 85, 86) has shown a very strict correlation

between scale pigments and resistance. The white varieties are

uniformly susceptible while the red and yellow varieties are only

slightly attacked. The resistant principle can be secured by

making a cold water extract of the dry, outer colored scale. This so¬

lution is highly toxic to the spores and mycelium of the fungus. A

similar extract from white scales promotes normal germination

and growth. When the dry outer scale is removed even colored

bulbs are readily infected. The interpretation of this is found

in the fact that the toxic substance is readily dissolved from the

dead outer scales into the soil water thus deactivating the fungus

before invasion. In the succulent scale the epidermal cells which

are the ones containing the pigment show reduction of this pig¬

ment while the mycelium is still in the outer cell wall and has not

actually invaded the cell lumen. This is due either to autolytic

processes within the cell or to a diffusible substance secreted by

the fungus. Resistance by the host, however, is accomplished by

the dry outer scales which serve as a barrier to the fungus through

the action of their soluble toxic substance.

Walker — Nature of Disease Resistance in Plants.

239

Other Cell Contents in Eelation to Resistance

Jones, Giddings, and Lutman (44) considered that resistance

of certain varieties of potato to the late blight fungus (Phytoph-

thora infestans) is due to something within the tissues of the leaf

and the tuber, rather than to difference in epidermal structures.

Their studies on the relation of the potato cell sap to the fungus

indicate that acidity of the former has little to do with the disease

resistant quality. Attempts to study the relation of the extracted

juice from susceptible and from resistant varieties to the growth

of the fungus yielded largely negative results. In conclusion they

say “this (disease resistant) product may, therefore, be assumed

to be either a compound, modified or destroyed by cooking and

weakened or removed by filtration through porcelain, or else it

may be so intimately associated with the living protoplasm as to.

be inseparable from it by the processes employed.’’

Wiltshire (95) studied the relation of apple and pear scab

fungi to plants susceptible and plants highly resistant to their

attack. Since these fungi penetrated the cuticle of both susceptible

and resistant plants he concluded that immunity does not depend

upon freedom from attack. A study of the germination and growth

of spores of the apple scab fungus (Venturia inaequalis) in ex¬

pressed juice of the host showed in one experiment that germina¬

tion was inhibited in the sap of both susceptible and resistant

varieties, while in the diluted sap no difference from germination

in water was observed. In another experiment the germ tubes

of the conidia appeared to grow better in the sap of susceptible

than of resistant varieties. These experiments indicate that some

chemical substance which had an inhibitive effect upon the fungus

was present in the expressed cell sap, but Wiltshire apparently

did not carry the investigation far enough to definitely establish

the fact or to throw any light upon the nature of the inhibitive

substance.

The fungicidal effect of volatile substances within the plant cell

has received some attention. Pasteur (62) early noted the retard¬

ing effect of onion juice upon yeasts. Bernard (6) noted the

fungicidal effect of volatile substances in the tissue of an orchid

(Loroglossum) upon the mycorhizal fungus from other closely

related species. Walker (85, 86) noted that the volatile onion

oil has a retarding effect upon germination and growth of the

onion smudge fungus ( Collet otrichum circinans). It was shown.

240 Wisconsin Academy of Sciences, Arts, and Letters.

however, that although this may be a factor in limiting the para¬

sitism of the latter fungus, it was not evidently a factor contribut¬

ing to the dilference in resistance between colored and white varie¬

ties. Brown (12, 13) has recently pointed out the effect on

germinating of fungus spores of substance diffusing from the

host cell into the infection drop; and in this work it was further

shown that volatile oils from various plants may have in some

cases detrimental and in other cases stimulative effects upon the

spores.

Summary

For purposes of convenience let us outline the main evidence to

date as to the nature of disease resistance, remembering that since

much of the work is fragmentary and not uniformly reliable this,

is merely a suggestive and tentative summary based wholly on

statements in the original texts.

Orton (61) classifies the various known types of resistance into

three classes: (1) avoidance of disease, (2) endurance of disease,

and (3) true resistance to disease. We include in the discussion

the latter class, in which we consider cases where through some

inherent quality in its composition the plant is capable of success¬

fully resisting the attack of a given parasite. In consideration

of the subject the importance of environment as affecting the ex¬

pression of resistant qualities is emphasized. In the few instances

where the hereditary nature of resistance has been studied it is

determined in some cases by a single factor, in other cases by

multiple factors. Vavilov’s (82) separation of the causes of true

resistance into two groups, mechanical or passive and physiolog¬

ical or active is useful. Numerous instances where mechanical

structures of the host plant are responsible for resistance have

been noted. These include hairiness of leaves and open habit of

growth in the case of potatoes resistant to blight, lack of hair on

terminal end of kernels in case of wheat resistant to bunt, external

waxy layers in case of raspberries resistant to Coniothyrium and of

barley resistant to stem rust. Differences in stomatal structure or

size have been cited in several cases as responsible for the resistant

quality. The exclusion of parasites by means of thick cuticle or

the formation of corky layers is quite well established in a few

eases.

Physiological or internal factors as causes of resistance have

been noted in a variety of instances. These internal differences are

Walker — Nature of Disease Resistance in Plants.

241

found in the cell membranes or in the cell contents. In the first

class are included (1) the internal cell walls as a factor in limiting

the potato-rotting fungus (Pythium deharyanum) , (2) the forma¬

tion of a layer of suberized cell walls in the cortex of flax plants

resistant to wilt, and (3) evidence of differences in cell wall

composition in plums resistant or susceptible to attack by Sclero-

tinia cinerea.

In the second case — that of internal differences in the cell con¬

tents — ^we have first the examples in the various cases of rust re¬

sistance. While the physiology of the process is not clear we know

that invasion of the cells of the resistant plant occurs but that

progress is stopped either by lack of adjustment on the part of the

fungus or by the presence of toxic substances in the host protoplasm.

A theory of chemotropism to explain resistance has been offered

by Massee (53) but it has never been well substantiated. The

correlation between cell sap acidity and resistance was brought

out by certain earlier workers; more recent investigations, how¬

ever, with other diseases has failed to show such a relation.

Tannins have been shown to be more or less toxic to fungi and there

are suggestions that in some cases this may contribute to resist¬

ance. Correlation between the anthocyan or flavone pigments and

resistance has been pointed out in a number of instances. In only

one case, that of onion, is the evidence sufficient to justify the sup¬

position that the coloring matter is the resistant principle, and

even there further investigation is necessary to ascertain definitely

whether it is the coloring matter or some other closely associated

substance.

The presence within the host tissues of a volatile substance

more or less toxic to parasites of the same tissue has been shown

in the case of onion. Curiously enough this substance has not been

correlated with resistance although it may play a role in limiting

the parasitism of the smudge fungus.

The above summary is sufficient to show that the true nature of

resistance has been exceedingly difficult to determine and, more¬

over, it is quite obvious that almost every case of resistance very

probably presents a specific problem. In certain cases, no doubt,

external factors, such as size of stomata, hairiness of leaf surface,

or presence of bloom are factors contributing to resistance. In

a larger number of cases, however, the underlying causes will be

found in the interrelations between the parasite and the host cell

membranes or the host cell contents. We can conceive of several

242 Wisconsin Academy of Sciences, Arts, and Letters.

factors entering into this relationship. Resistance may be due

to a substance (or substances) in the host tissue which is either

toxic to the parasite or is capable of neutralizing or destroying

the enzymes or toxins produced by the parasite and hence may

preclude or check invasion; it may be due to a lack of proper

nutrients in the host tissue for the development of the invader;

or it may be due to a lack of ability on the part of the parasite,

first, to penetrate the host plant or, secondly, so to adjust itself

to the chemical and physical complex of its new substrate as to

make food materials, even though present, available for absorption

and assimilation. It is entirely possible that one or all of these

factors may enter into a given case of disease resistance. It should

also be recognized at the outset that a clear conception of the

nature of the parasitism occurring in any given case is essential to a

clear understanding of resistance. Indeed, the two processes are

merely phases of a single phenomenon which involves the inter¬

relation of two living organisms. Moreover, it should be made

clear that, as Duggar (28) has pointed out, ‘‘there are all grades

of parasitism and there must be a variety of effects induced in the

host, including changes essentially autolytic.’’ He compares, for

instance, the effect of Cyst opus candidus (Pers.) Lev. upon the

host tissues with that of closely related fungi, as Phytophthora in-

festans or Pythium deharyanum; and again, the rusts which have

“little tendency to kill immediately the tissues which they invade”

with the large group of parasites which have no contact with the

living protoplasm but kill the cells in advance by means of

secreted diffusible substances. The greatest advance in our knowl¬

edge of the nature of disease resistance will therefore come from a

thoroughgoing, persistent study of individual cases, and especially

of those which seem to offer the most accessible approach.

Literature Cited

(1) . Aamodt, Olaf S. The imheritance of resistance to several biologic

forms of Puccinia graminis tritici in a cross between Kanred and

Marquis wheats. (Abstract.) Phytopathology 12: 32. 1922.

(2) . Allen, Buth. F. A cytological study of infection of Baart and Kan¬

red wheats by Puccinia graminis tritici. Jour. Agr. Eesearch 23:

131-152. 1923.

(3) . Appel, Otto. Disease resistance in plants. Science n. s. 41: 773-782.

1915.

(4) . Averna-Sacca, Bosario. acidita dei succhi delle piante in rapporto

alia resistenza contro gli attach! dei parassiti. Staz. Sper. Agr.

Ital. 43: 185-209. 1910.

Walker — Nature of Disease Resistance in Plants.

243

(5) . Barrus, M. F. Varietal susceptibility of beans to strains of Colleto-

trichum lindemuthianum (Saec. & Magn.) B. & C. Phytopathol¬

ogy 8: 589-614. 1918.

(6) . Bernard, Noel. Sur la fonction fungicide des bulbes d’Ophrydees.

Ann. Sci. Nat. Bot. 9® serie 14: 221-234. 1911.

(7) . Biffen, R. H. Studies in the inheritance of disease resistance. Jour.

Agr. Sci. 2: 109-128; 4: 421-429. 1907-1912.

(8) . Blackman, V. H. and Welsford, E. J. Studies in the physiology of

parasitism. II. Infection by Botrytis cinerea. Ann. Bot. 30:

389-398. 1916.

(9) . Brown, William. Studies in the physiology of parasitism. I. The

action of Botrytis cinerea. Ann. Bot. 29: 313-348. 1915.

(10) . Studies in the physiology of parasitism. III. On the rela¬

tion between the ^infection drop’ and the underlying host tissue.

Ann. Bot. 30: 399-406. 1916.

(11) . - Studies in the physiology of parasitism. IV. On the dis¬

tribution of cytase in cultures of Botrytis cinerea. Ann. Bot. 31:

489-498. 1917.

(12) . Studies in the physiology of parasitism. VIII. On the exos¬

mosis of nutrient substances from the host tissue into the in¬

fection drop. Ann. Bot. 36: 101-119. 1922.

(13) . - Studies in the physiology of parasitism. IX. The effect on

the germination of fungal spores of volatile substances arising

from plant tissues. Ann. Bot. 36: 285-300. 1922.

(14) . Burkholder, W. H. The production of an anthracnose resistant white

marrow bean. Phytopathology 8: 353-359. 1918.

(15) . Biisgen, M. Ueber einige Eigenschaften den keimlinge parasitischer

Pilze. Bot. Ztg. 51: 53-72. 1893.

(16) . Butler, E. J. Immunity and disease in plants. Agr. Jour. India

(Spec. Indian Sci. Cong. No.) 1918: 10-28. 1918.

(17) . Cobb, N. A. Contributions to an economic knowledge of the Austral¬

ian rusts (Uredineae). Agr. Gaz. N. S. Wales 1: 185-214, 1890;

3: 44-68; 181-212. 1892. Literature cited 3: 190-200.

(18) . Comes, Orazio. Della resistenza dei frumenti alle ruggini. Stato

attuale della quistione e provvedimenti. Atti. R. Istituto d’ln-

corraggiamento Napoli. Ser. 6, 64: 419-441. 1913. (Rev. in

Bui. Agr. Int. and PL Dis. 4: 1117-1119. 1913.)

(19) . - Della resistenza dei frumenti alle ruggini ed in generale delle

piante alle loro cause nemichi. Ann. d. R. Scuola Sup. di Agri-

coltura. Ser. 2, 12: 419-473. 1914.

(20) . Cook, M. T., Bassett, H. P., Thompson, F., and Taubenhaus, J. J.

Protective enzymes. Science n. s. 33: 624-629. 1911.

(21) . - and Taubenhaus, J. J. The relation of parasitic fungi

to the contents of the cells of the host plants (I. The toxicity

of tannin). Del. Agr. Exp. Sta. Bui. 91. 77 p. 1911.

(22) . - and Taubenhause, J. J. The relation of parasitic fungi

to the contents of the cells of the host plants (II. The toxicity

of vegetable acids and the oxidizing enzyme.) Del. Agr. Exp.

Sta. Bui. 97: 53 p. 1912.

244

Wisconsin Academy of Sciences y Arts, and Letters.

(23) . - and Wilson, G. W. The influence of the tannin content of

the host plant on Endothia parasitica and related species. Bot.

Gaz. 60: 346-361. 1915.

(24) . Curtis, K. M. The life-history and cytology of Synchytrium endohiot-

icum (Schilb.) Perc., the cause of wart disease in potato. Phil.

Trans. Eoy. Soc. London. Ser. B, 210: 409-478. 1921.

(25) . Damell-Smitli, G. P. Some observations on bunt and fungicides.

Agr. Gaz. N. S. Wales 21: 751-756. 1910.

(26) . DeBary, A. Ueber einige Sclerotinien und Selerotienkrankheiten.

Bot. Ztg. 44: 377-387; 393-404; 409-426; 433-441; 449-461; 465-

474. 1886.

(27) . Dey, P. K. Studies in the physiology of parasitism. V. Infection

by Collet otriclium lindemutJiianum. Ann. Bot. 33: 305-312. 1919.

(28) . Duggar, B. M. Physiological plant pathology. Phytopathology 1:

71-78. 1911.

(29) . Durrell, L. W., and Parker, J. H. The comparative resistance of

varieties of oats to crown and stem rusts. Iowa Agr. Exp. Sta.

Ees. Bui. 62: 25-56d. 1920.

(30) . Eriksson, J., and Henning, E. Die Getreideroste. 464 p. Stockholm,

1896. Literature cited, p. 355-363.

(31) . Fromme, F. D., and Wingard, S. A. Varietal susceptibility of beans

to rust. Jour. Agr. Eesearch 21: 385-404. 1921.

(32) . Freeman, E. M. Eesistance and immunity in plant diseases. Phyto¬

pathology 1: 109-115. 1911.

(33) . Gaines, E. F. The inheritance of resistance to bunt or stinking smut

of wheat. Jour. Amer. Soc. Agron. 12: 124-131. 1920.

(34) . Garber, E. J. A preliminary note on the inheritance of rust re¬

sistance in oats. Jour. Amer. Soc. Agron. 13: 41—43. 1921.

(35) . Gardner, M. W., and Kendrick, J. B. Bacterial spot of tomato.

Jour. Agr. Eesearch 21: 123-156. 1921.

(36) . Gibson, C. M. Notes on infection experiments with various Uredi-

neae. New Phytologist 3: 184-191. 1904.

(37) . Giddings, N. J. Infection and immunity in apple rust. West Va.

Agr. Exp. Sta. Bui. 170: 71 p. 1918.

(38) . Hawkins, L. A. Growth of parasitic fungi in concentrated solu¬

tions. Jour. Agr. Eesearch 7: 255-260. 1916.

(39) . - , and Harvey, R. B. Physiological study of the parasitism of

Pythium deharyanum Hesse on the potato tuber. Jour. Agr. Ee¬

search 18: 275-297. 1919.

(40) . Johnson, James. Inheritance of disease resistance to Thielavia

basicola. (Abstract.) Phytopathology 11: 49. 1921.

(41) . Jones, F. R. The leaf-spot diseases of alfalfa and red clover caused

by the fungi Pseudopeziza medicaginis and Pseudopeziza trifolii,

respectively. U. S. Dept. Agr. Bui. 759: 38 p. 1919.

(42) . Jones, L. R. Disease resistance of potatoes. TJ. S. Dept. Agr., Bur.

PI. Ind. Bui. 87: 39 p. 1905.

Walker — Nature of Disease Resistance in Plants,

245

(43) . - Pectinase, the cytolytic enzyme produced by Bacillus carotovorus

and certain other soft-rot organisms. N. Y. (Geneva) Agr. Exp.

Sta. Tech. Bui. 11: 289-368. 1909; also Vt. Agr. Exp. Sta. Bui.

147: 289-360. 1909.

(44) . - , Giddings, N. J., and Lutman, B. F. Investigations of the

potato fungus Phytophthora infestans. U. S. Dept, of Agr. Bur.

PI. Ind. Bui. 245: 100 p. 1912.

(45) . - - , Walker, J. C., and Tisdale, W. B. Fusarium resistant cab¬

bage. Wis. Agr. Exp. Sta. Ees. Bui. 48. 34 p. 1920.

(46) . Klapliaak, P. J., and Bartlett, H. H. A preliminary notice of geneti-

cal studies of resistance to mildew in Oenothera. Amer. Jour.

Bot. 9: 446-458. 1922.

(47) . Levine, M. N., and Stakman, E. C. Biologic specialization of

Puccinia graminis secalis. (Abstract.) Phytopathology 13: 35.

1923.

(48) . MacDougal, D. T. The beginnings and physical basis of parasitism.

Plant World 20: 238-244. 1917.

(49) . McBostie, G. P. Inheritance of anthracnose resistance as indicated

by a cross between a resistant and a susceptible bean. Phyto¬

pathology 9: 141-148. 1919.

(50) . - Inheritance of disease resistance in the common bean. Jour.

Amer. Soc. Agron. 13: 15-32. 1921.

(51) . Mains, E. B., and Jackson, H. S. Strains of the leaf-rust of wheat,

Puccinia triticina, in the United States. (Abstract.) Phytopa¬

thology 13: 36. 1923,

(52) . Marryat, Dorothea C. E. Notes on the infection and histology of

two wheats immune to the attacks of Puccinia glumarum,

yellow rust. Jour. Agr. Sci. 2: 129-138. 1907.

(53) . Massee, George. On the origin of parasitism in fungi. Phil. Trans.

Roy. Soc. London. Ser. B, 197: 7-24. 1905.

(54) . Melchers, Leo E., and Parker, John H. Rust resistance in winter-

wheat varieties. U. S. Dept. Agr. Bui. 1046: 32 p. 1922.

(55) . - - - Inheritance of disease re¬

sistance to black stem rust in crosses between varieties of com¬

mon wheat {Triticum vulgare). (Abstract.) Phytopathology

12: 31-32. 1922.

(56) . Miyoshi, M. Die Durchbohrung von Membranen durch Pilzfaden.

Jahrb. Wiss. Bot. (Pringsheim) 28: 269-289. 1895.

(57) . Nilsson-Ehle, H. Resistenz gegen Gelbrost beim Weisen. Kreuz-

ungsuntersuchungen an Hafer und Weisen. II. Lund’s Univ.

Arsskr. n. f. afd. 2, 7: 57-82. 1911.

(58) . Nordhausen, M. Beitrage zur Biologie parasitarer Pilze. Jahrb.

Wiss. Bot. (Pringsheim) 33: 1-46. 1898.

(59) . Norton, J. B. Methods used in breeding asparagus for rust resist¬

ance. U. S. Dept. Agr. Bui. PI. Ind. Bui. 263: 60 p. 1913.

(60) . Orton, C. R., and Weiss, Freeman. The reaction of first generation

hybrid potatoes to the wart disease. Phytopathology 11: 306-

310. 1921.

246

Wisconsin Academy of Sciences , Arts, and Letters.

(61) . Orton, W. A. The development of farm crops resistant to disease.

U. S. Dept. Agr. Yearbook 1908: 453-464. 1909.

(62) , Pasteur, L. Memoire sur la fermentation appelee lactique. Ann. d.

Chemie et d. Physique. Ser, 3, 52: 404-418. 1858.

(63) . Pool, V. W., and McKay, M. B. Eelation of stomatal movement to

infection by Cercospora heticola. Jour Agr. Eesearch 5: 1011-

1038. 1916.

(64) . Beed, Geo. M. Varietal resistance and susceptibility of oats to

powdery mildew, crown rust, and smuts. Mo. Agr. Exp. Sta.

Ees. Bui. 37: 41 p. 1920.

(65) . Reed, H. S., and Crabill, C. H. The cedar rust disease of apples

caused by Gymnosporagium juniperi-virginianae Schw. Va. Agr.

Exp. Sta. Tech. Bui. 9: 106 p. 1915. Literature cited, p. 40-43.

(66) . Rosenbaum, J. and Sando, C. E. Correlation between size of the

fruit and the resistance of the tomato skin to puncture and its

relation to infection with Macrosporium tomato Cooke. Amer. Jour.

Bot. 7: 78-82. 1920.

(67) . Salmon, E. S. On the stages of development reached by certain

biologic forms of Erysiphe in cases of non-infection. New

Phytologist 4: 217-222. 1905.

(68) . Smith, R. E. The parasitism of Botrytis cinerea. Bot. Gaz. 33: 421-

436. 1902.

(69) . Sorauer, P. TJeber die pradisposition der Pflanzen fiir parasitare

Krankheiten. Arb. Deut. Landw. Gesell. Jahresber. Sonder-

aussch. Pflanzenschutz 12: 193-210. 1903. Literature cited,

p. 198.

(70) . Spinks, G. T. Factors affecting susceptibility to disease in plants.

Jour. Agr. Sci. 5: 231-247. 1913.

(71) . Stakman, E. C. A study in cereal rusts. Physiological races. Minn.

Agr. Exp, Sta. Bui. 138: 56 p. 1914.

(72) . - Relation between Puccinia graminis and plants highly resistant

to its attack. Jour. Agr. Eesearch 4: 193-199. 1915,

(73) . - , and Aamodt, Olaf S. The effect of fertilizers on the develop¬

ment of stem rust of wheat. Phytopathology 12: 31. 1922.

(74) . - , and Levine, M. N. The determination of biologic forms of

Puccinia graminis on Triticum spp. Minn. Agr. Exp. Sta. Tech. Bui.

8: 10 p. 1922.

(75) . - , and Bailey, D. L. Biologic specialization of

Puccinia graminis avenae. (Abstract.) Phytopathology 13: 35.

1923.

(76) . - , and Leach, J. G. New biologic forms of

Puccinia graminis. Jour. Agr. Eesearch 16: 103-105. 1919.

(77) . Stuart, William. Disease resistance of potatoes. Vt. Agr. Exp. Sta.

Bui. 122: 105-136. 1906.

(78) . Tisdale, W. B. The influence of soil temperature and soil moisture

upon the Fusarium disease in cabbage seedlings. Jour. Agr.

Research. 24: 55-86. 1923.

(79) . Tisdale, W. H. Flaxwilt: a study of the nature and inheritance of

wilt resistance. Jour. Agr. Research 11: 573-606. 1917.

Walker — Nature of Disease Resistance in Plants.

247

(80) . Valleau, W. D. Varietal resistance of plums to brown-rot. Jour.

Agr. Eesearch 5: 365-396. 1915.

(81) . Vavilov, N. I. Breeding for resistance to rust and mildew in wheat.

Jour. Gen. 4: 49-65. 1914.

(82) . Vavilov, N. I. Immunity of plants to infectious diseases. Proc.

of the Petrovski Agricultural Academy 1918: 1-239. 1919.

(With a resume in English.)

(83) . Voges, Ernst. Die Bekampfung des Fusicladium. Ztschr. Pflanzen-

krank. 20: 385-393. 1910.

(84) . Walker, J. C. Notes on the resistance of onions to anthracnose.

(Abstract.) Phytopathology 8: 70-71. 1918.

(85.) - Onion smudge. Jour. Agr. Research 20: 685-722. 1921.

(86) . - Disease resistance to onion smudge. Jour. Agr. Research.

24: 1019-1040. 1923.

(87) . Ward, H. Marshall. A lily-disease. Ann. Bot. 2: 319-382. 1888.

(88) . - On the relations between host and parasite in the bromes and

their brown rust, Puccinia dispersa (Erikss). Ann Bot. 16: 233—

315. 1902.

(89) . - Recent researches on the parasitism of fungi. Ann. Bot. 19:

1-54. 1905.

(90) . Waterhouse, W. L. Studies in the physiology of parasitism. VIII.

Infection of Berberis vulgaris by sporidia of Puccinia graminis.

Ann. Bot. 35: 557-564. 1921.

(91) . Weimer, J. L., and Harter, L. L. Wound-cork formation in the

sweet potato. Jour. Agr. Research 21: 637-647. 1921.

(92) . Weiss, F., and Harvey, R. B. Catalase, hydrogen-ion concentration,

and growth in the potato wart disease. Jour. Agr. Research 21:

589-592, 1921.

(93) . Willaman, J. J., and Sandstrom, W. M. Biochemistry of plant dis¬

eases III. Effect of Sclerotinia cinerea on plums. Bot. Gaz.

73: 287-307. 1922.

(94) . - , and Davison, F. R. Biochemistry of plant diseases IV,

Proximate analysis of plums rotted by Sclerotinia cinerea. Bot.

Gaz. 74: 104-109. 1922.

(95) . Wiltshire, S. P. Infection and immunity studies on the apple and

pear scab fungi (Venturia inaequalis and V. pirina). Ann. Appl.

Biol. 1: 335-350. 1915.

J

SOME FERNS OF SOUTHWESTERN WISCONSIN

Sister M. Ellen

An attempt to discover and to identify the pteridophytic flora

in the vicinity of Sinsinawa Mound (Grant County) was both in¬

teresting and attractive. The types of habitat in this locality are

so varied within a radius of three or four miles that a fern lover

might hope to find almost any or all of the native, temperate forms.

These various plant habitats may be classified as follows: deep

moist woods, dry open woods, wet rocky bluffs, tall limestone cliffs,

and waste lands.

It was somewhat disappointing but none the less interesting to

find not a single plant of Polypodium vulgare, the common rock

fern, which is so generally distributed throughout the state. . The

Woodsias, rock-loving Aspleniums, Aspidium marginale, the Lyco¬

podiums, and other common forms were also hunted in vain.

In the following list the nomenclature follows Gray^s Manual,

seventh edition.

POLYPODIACEAB

Adiantum pedatum L.

Common.

Aspidium spinulosum (0. F. Muller) Sw.

Occasional plants in deep woods.

Asplenium Filix-femina (L) Bernh.

Common everywhere.

Camptosorus rhizopJiyllus (L.) Link.

Three places on rocks; (a) on Sinsinawa Mound; (b) about

three and one-half miles west of the Mound; and (c) about four

miles southwest of the Mound.

Cyst'opteris bulbifera (L.) Bernh.

Very abundant on rocks.

Cystopteris fragilis (L.) Bernh.

Common everywhere.

250 Wisconsin Academy of Sciences, Arts, and Letters,

Cryptogramma Stelleri (Gmel.) Prantl.

Abundant on rocks.

Pellaea atropurpurea (L.) Link.

Very abundant on rocks.

Onoclea sensihilis L.

Abundant in moist woods.

Onoclea Struthiopteris (L.) Hoffm.

In wet fields and transplanted on the campus of St. Clara College.

Pteris aquilina L.

Abundant in open woods.

OSMUNDACEAE

Osmunda Claytoniana L.

Very abundant in moist woods.

Ophioglossaceae

Botrychium virginianum (L.) Sw.

Common in deep woods.

Equisetaceae

Equisetum arvense L.

Common everywhere on roadsides and in margins of fields.

Equisetum hyemale L. var. rohustum (A. Br.) A. A. Eaton.

On rocks.

Department of Botany,

University of Wisconsin

NOTES ON PARASITIC FUNGI IN WISCONSIN— IX

J. J. Davis

The greater part of the field work in 1920 was done along the

lower Wisconsin and the Mississippi river, the Chippewa and its

tributaries, the Yellow, the Fisher, and the Jump rivers. The sea¬

son was an unfavorable one because of low precipitation, which

however allowed an unusual proportion of working days.

Plasmopara acalyphae Wilson previously known only from the

type station at Madison was found in July 1920 at Caryville in

the Chippewa valley. As usual the development of conidiophores

was scanty. [Collected since at Lone Rock, Arena and Oconto but

always scanty.]

Plasmopara ohducens Schroet. in summer is sometimes confined

to small angular leaf areas which become brown, thus causing spot¬

ting of the affected leaves. The summer development of this mil¬

dew is more common northward.

What is presumed to be mycelium and conidia of Sphaerotheca

humuli (DC.) Burr, is sometimes abundant on leaves of Bubus

allegheniensis at Madison during the early part of the season, dis¬

appearing in summer without development of perithecia. It has

been noticed that the thickened leaves of the plants bearing Caeoma

are especially liable to this infection.

In a collection on twigs and young leaves of Physocarpus opuli-

folius from Fish Creek referred to Sphaerotheca humuli (DC.)

Burr, the perithecia bear long tapering appendages like those of

Podosphaera leucotricha (E. & B.) Salmon in addition to short

rhizoid basal ones. These long appendages are not apical, however,

but basal or equatorial, and the spores are not crowded in the

ascus. Nevertheless the Sphaerotheca would appear to be the

‘^next of kin^’ to the apple mildew. The mildew on Physocarpus

appears to over winter in the twigs or buds.

Phyllachora oryzopsidis Theiss. & Syd. has been collected on

Oryzopsis asperifolia at Mosinee. The collection was made in

252 Wisconsin Academy of Sciences, Arts, and Letters.

September and the spores are smaller than indicated in the de¬

scription.

For the parasite recorded in ^ ‘ Notes V, p. 696, under the name

Lophodermium linear e Pk. (Ehytisma linear e Pk., Hypoderma

lineare Pk.) von Hoehnel has proposed a new genus, Bifusella (Ann.

My col. 15:318-19.)

In ‘‘Notes” IV pp. 683-4 reference was made to a group of

variable Sphaerioidaceae on Atriplex and Chenopodium members

of which have been referred to Phyllosticta, Ascochyta, Diplodina,

Septogloeum, Stagonospora, Phleospora and Septoria and to the

reference of the group as a whole to Stagonospora atriplicis West,

by Lind and to Septoria chenopodii West, by Grove. In a discus¬

sion of some members of this group by J. B. Ellis the conclusion

was reached that “This variability would seem to strengthen the

supposition that all the forms here enumerated may be referred to

Septoria (Phyllosticta) atriplicis Desm. ” {Journ. My col. 4:117-18

[1888].) A parasite collected on Chenopodium album at Cary-

ville July 21, 1921, appears to be an extreme variant of this group

because of the long and slender sporules. The following notes

were made : Spots suborbicular, immarginate, light yellow to yel¬

lowish green, averaging about % cm. in diameter ; pycnidia

amphigenous, scattered evenly over the spots; sporules discharged

in colorless cirri, hyaline, straight or curved, 1-3 septate,

24-67x2%-3%/>t. The sporules often appear continuous in a water

mount. Apparently they are catenulate, their length depending

upon the number of abstrictions of the primary sporule.

In a collection of Septoria aquilegiae Penz. & Sacc. from Durand

(July 12, 1920) the sporules are mostly about twice the typical

length but without noticeable increase in diameter.

Septoria ampelopsidis Ellis appears to be a better developed

state of the parasite recorded in the provisional list under the

name Septogloeum ampelopsidis (E. & E.) Sacc. This form with

definite pycnidia and long and slender sporules has been collected

on Psedera at Madison, Wausau and Durand. I have labeled it

Septoria ampelopsidis (E. & E.) Ellis.

In a collection of Septoria on Cicuta maculata made at Weyer¬

haeuser, September 9, 1918, the pycnidia are borne on small (1 mm.

or less) subcircular white arid spots having a dark reddish brown

border; the globose pycnidia are 80-85/x in diameter, the sporules

Davis — Notes on Parasitic Fungi in Wisconsin — IX, 253

30-45xl-l%/-t. It was labeled provisionally Septoria umhellif era-

rum Kalchb.

In “Notes’’ V it was stated that while in Oloeosporium caryae

Ell. & Dearn. as found on Cary a ovata the acervuli are hypophyl-

lous in the collections on C. cordiformis they are epiphyllous. In

a collection on the latter host made at Caryville in 1920 they are

amphigenous.

In a collection of Marssomna fraxini Ell. & Davis from the

Mississippi river bottoms at Glen Haven the sporules range from

30-50x2%-4/x. No matter what the length of the sporule but a

single septum appears.

R. E. Stone states that the ascogenous state of Marssonina poten-

tillae (Desm.) Magn. is Mollisia earliana (E. & E.) Sacc.

In the parasite recorded as Cylindrosporium clematidis in the

provisional list the sporuligerous stroma, under favorable condi¬

tions, forms a hollow sphere and the spore body is therefore a

pycnidium.

In “Notes” IV, p. 687 Septocylindrium caricinum Sacc. was

recorded as occurring in Wisconsin. Examination of the specimen

on Carex grisea from Blue Mounds however reveals only tufts of

hyphae indistinguishable from the conidiophores of Cercospora

caricina Ell. & Dearn. which are sometimes nearly or quite hyaline.

The Septocylindrium record therefore should be stricken out.

Cercosporella apocyni E. & K. and Cylindrosporium apocyni

E. & E. sometimes occur on the same spots. May it be that they

are conidial forms of one species?

Specimens of the parasite on Populus tremuloides referred to

Cladosporium subsessile Ell. & Barth, have been collected in which

the tufts are scattered over unaltered portions of the leaves.

W. B. Tisdale has shown the connection between Heterosporium

gracile Sacc. and the ascigerous stage Didymellina iridis (Desm.)

Hoehn. {Phytopathology 10:153-4.)

Cercospora arctostaphyli Davis {Trans. Wis. Acad. 18:268)

seems to have been founded upon a misapprehension. There is no

specimen in the University of Wisconsin herbarium and the char¬

acters ascribed are those of Cercospora gaultheriae E. & E. It

should be stricken out.

254 Wisconsin Academy of Sciences, Arts, and Letters.

In “Notes’’ V p. 694 reference was made to a small collection on

Ambrosia trifida from Maiden Rock which was referred to Enty-

loma polysporum (Pk.) Pari. This form with definite, orbicular,

yellow, thickened, concavo-convex spots is however merely a late

stage of E. compositarum Pari, although it differs strikingly in

appearance from the earlier conidiophorous state. The spore walls

do not become thick. Pield observation is said to indicate that

Entyloma calendulae (Oud.) D By. as it occurs in Europe is com¬

posed of races adapted to various genera of Compositae and H. & P.

Sydow have proposed eight binomials for the designation of these

races {Ann. Mycol. 16:244). The same kind of evidence indicates

that an equal number of races of Entyloma compositarum Pari,

exist in Wisconsin but I see nothing to be gained by using bi¬

nomials for them as it would tend to obscure their very close kin¬

ship. It would require a large amount of experimentation to dem¬

onstrate that under no circumstances is there passage from one

host genus to another. There seems to be a very close adaptation

of parasite to host in the Ustilaginales in general and with that in

mind the citation of the host gives the information in this case that

would be conveyed by a binomial without obscuring the conception

of phylogenetic unity. It seems quite possible that no violence

would be done in considering E. calendulae (Oud.) D By. and E.

compositarum Pari, as two geographical series of races of a single

species.

In Notes II p. 106 it was stated that Uromyces graminicola Burr,

had been collected at Madison on the railroad right of way. It

was not again observed until 1920 when it was found at Durand

in the western part of the state.

Por the rust on Koeleria cristata having its aecial stage on Liatris

I am using the designation Puccinia koeleriae-liatridis instead of

P. liatridis Bethel. P. koeleriae Arth. with aecia on Berberis is con¬

sidered to be distinct.

Puccinia pustulata (Curt.) Arth. with aecia on Comandra is

treated as a race of P. andropogonis Schw. in North American

Flora.

Puccinia simillima Arth. is united with P. magnusiana Koern. in

North American Flora.

In the provisional list Rubus idaeus aculeatissimus, B. occiden-

talis and B. triflorus were given as hosts of Pucciniastrum arcticum

Davis — Notes on Parasitic Fungi in Wisconsin — IX. 255

americanum Farl. The collections on Ruhus triflorus however

should be referred to the type while I find no specimens in the

herbarium which I can determine as being on B. occidentalis. A

specimen on this host collected by B. 0. Dodge in Wisconsin at

Algoma was reported by Arthur as P. arcticum but' Dr. Arthur in¬

forms me that this was an error and that the specimen is of the

americanum type which is not rare in Wisconsin on the red rasp¬

berry and that the host is B. strigosus. Arthur has proposed the

raising of the variety americanum Pari, to specific rank (Bull. Torr.

Bot. Club 47 : 468) but certain forms on Bubus idaeus aculeatissimus

having globose peridia seem closely allied to the type.

aculeatissimus.

Fig. 4. Section through a uredo sorus of Puceiniastrum on Bubus triflorus.

Drawn by E. M. Gilbert with the aid of camera lucida. Pig-. 4 was drawn

from a small sorus and the disparity in size is somewhat exaggerated.

For the Abies-Salix rust recorded in ‘‘Notes” II & III under the

binomial Melampsora arctica Rostr. Arthur proposes M. ameri-

cana {Bull. Torr. Bot. Club 47;465--68.) Besides the localities

which he gives the Caeoma has been collected in Wisconsin at

Mountain and at Kelly lake in Oconto county and at Solon Springs.

All of the localities are in either the northeastern or the north¬

western corner of the state. Various collections of uredinia and

telia on species of Salix have been referred to this species with

doubt as they do not seem to be readily separable by their morpho¬

logical characters from those of Melampsora bigelowii Thuem.

Additional Hosts*"

Not previously recorded as bearing the parasites mentioned in

Wisconsin.

Synchytrium aureum Schroet. On Hydrocotyle americana Cary-

ville. Many leaves bore the parasite at this restricted station but

the sori were much scattered ; only one or very few on a leaf.

256 Wisconsin Academy of Sciences, Arts, and Letters.

Peronospora leptosperma DBy. was collected at Jump River

on a host that was referred to Artemisia serrata.

Peronospora ruhi Rabh. was found in June, 1920, at Blue

Mounds on Bubus allegheniensis growing along the railroad. The

mildew was apparently confined to plants parasitized by Caeoma.

Premia lactucae Regel. On Lactuca Scariola integrata, Madison.

Basidiophora entospora Roze & Cornu. On Aster TradescantL

Madison. Rather abundant on this host in the autumn of 1920. •

Gnomonia ulmea (Schw.) Thuem. Immature material was col¬

lected on Ulmus racemosa at Nekoosa in September.

In September, 1919, Phyllachora on Melica striata was collected

at Mosinee. The material is immature but the characters of the

ascomata are not those of Phyllachora melicae Speg.

^ Phyllosticta phomiformis Sacc. On Quercus bicolor. La Crosse.

Globose, dark brown, opaque cells 7-10/x diameter are also found

in this collection.

Phyllosticta grossulariae Sacc. On Ribes oxyacanthoides. Dur¬

and. Sporules smoky tinted.

Septoria nocti florae Ell. & Kell. On leaves and calyes of Silene

nivea. Durand. In the collection referred to this species the

sporules are 30-45x1 %-2V2/>«', continuous, but the material is not

mature.

Septoria gei Rob. & Desm. On Geum canadense. Glen Haven.

The pycnidia are imperfect in these specimens.

Septoria scrophulariae Pk. On Scrophularia leporella. Glen

Haven.

Septoria stachydis Rob. & Desm. On St achy s palustris. Coon

Valley.

Septoria commonsii Efll. & Evht. On Cirsium discolor. Durand.

Phleospora ulmi {Pt.) Wdllv. On Ulmus fulva. Fountain City.

In ‘ ‘ Notes ’ ’ VII mention was made of the fact that Gloeosporium

salicis West, had been found in Wisconsin on foreign species of

Salix only. In 1919 a tree of Salix alba vitellina bearing the Gloeo-

Davis — Notes on Parasitic Fungi in Wisconsin — IX. 257

sporium was observed at Fish Creek and under it a shrub of Salix

lucida also bearing the Gloeosporium.

Gloeosporium fraxineum Pk. On Fraxinus pennsylvanica.

Mosinee.

Gloeosporium aridum Ell. & Hoi. On Fraxinus nigra. Cary-

ville.

Collet otrichum graminicolum (Ces.) Wilson. On Melica striata,

Mosinee. A Colie totrichum which was abundant on leaves of Car ex

ehurnea at Pish Creek I do not distinguish from this species.

Marssonina rhahdospora (E. & E.) Magn. On Populus tremu-

loides. Millston, Cornell and Holcombe. My observation of this

parasite does not lead me to look with favor on the suggestion of

von Hoehnel that it is a form of Septoria populi Desm.

The parasite recorded under the name Marssonina potentillae

tormentillae Trail in ‘‘Notes’’ III p. 259 was collected on Bubus

canadensis at Tomahawk but immature or imperfectly developed

as usual. Collections of a similar character were made on a host

referred to Bubus hispidus at Millston in 1912.

Cylindrosporium vermiforme Davis. On Alnus crispa. Minoc-

qua. In the collection on this host, which was made in August, the

sporules are slender, being but about Sjn thick.

Bamularia lysimachiae Thuem. On Lysimachia terrestris. Vilas

County and Holcombe. Conidiophores up to 40/it long.

Bamularia virgaureae Thuem. On Solidago nemoralis. Cary-

ville.

Bamularia asteris (Phil. & Plowr.) Bubak. On Aster lateri-

florus. Nekoosa. Aster Tradescanti. Caryville and Madison.

Cercosporella pyrina Ell. & Evht. On Pyrus ioensis, Nekoosa.

A specimen on Erigeron annuus collected at Madison in Septem¬

ber and referred to Cercosporella cana Sacc. bears conidiophores

50— 100x3— 6 ju.

Cladosporium subsessile Ell. & Barth. On Populus grandi-

dentata. Muscoda.

Cladosporium triostei Pk. On Triosteum aurantiacum. Glen

Haven. In the material that I have seen the patches of conidio-

17

258 Wisconsin Academy of Sciences, Arts, and Letters.

phores may precede the spotting, the conidia are devoid of apical

papillae and the narrow base is truncate.

Cercospora caricina Ell. & Dearn. On culms, leaves and bracts

of Cy perns Schweinitzii. Caryville.

Cercospora oxybaphi Ell. & Hals. On Oxyhaphus hirsutus.

]\Ieridean.

Cercospora velutina Ell. & Kell. On leaves and stipules of Bap¬

tism bracteata. Caryville.

The Cercospora that was tentatively referred to C. flag ellif era

Atk. in “Notes” VIII has been collected at Glen Haven on what

was taken to be Lespedeza frutescens.

Cercospora viciae Ell. & Hoi. On Lathy rus palustris. Cary¬

ville.

From the examination of a collection on Solidago serotina made

at Durand August 18, 1920, the following notes were made : Spots

angular, blackish brown becoming alutaceous and finally white and

arid especially above, 1-2 mm. in diameter; conidiophores hyp-

ophyllous, single, in pairs or in small fascicles, erect, simple, terete

and continuous or torulose and septate, fuligenous, 33-147x4-6/x;

conidia obclavate, straight or but little curved, subhyaline,

67-200x4-6/x. Although the spots are smaller and more definite

and the conidiophores and conidia of a fuligenous cast rather than

brown I have referred this provisionally to Cercospora stomatica

Ell. & Davis as perhaps a modified form on a different species of

host.

Of a collection on the same species of host made at La Crosse

August 24, 1920, it was noted that the spots are definite, angular,

brown, mostly l-2mm. in diameter ; the conidiophores having

more or less of a fuligenous coloration, 33-63x4-6/x, the conidia

hyaline, upwardly attenuate, up to llO^a or more in length by 3-4/a

thick. I was at first inclined to consider this to be a form of Bamu-

laria virgaiireae Thuem. modified by the dry weather but it can

hardly be distinct from the collection referred to above.

On some of the leaves of the Durand material are small hypo-

phyllous patches of Ramularia having straight, hyaline, subulate

Davis — Notes on Parasitic Fungi in Wisconsin — IX, 259

conidiophores, 15-25x2/>i and catenulate, hyaline conidia which are

straight, cylindrical with acute ends, 10-30xl%-2^. I have

labeled this Bamularia virgaureae Thuem. Macroscopically the

frost like patches look like those of Cercosporella.

Schizonella melanogramma (DC.) Schroet. On Car ex longiros-

tris, Madison. (J. R. Heddle.)

Entyloma compositarurn Farl. On Ambrosia psilostachya. Tom¬

ahawk. This host was given in Trelease’s Preliminary List and

should have been included in the provisional list.

Of a Uredo collected at Cadott, July 23, 1920, on Car ex intume-

scens Dr. Arthur reports, “appears to me to belong to TJromyces

minutus Diet, rather than to Puccinia minutissima. I judge this

from the fact that the pores are almost entirely equatorial and also

because the host is more closely of the character of the Uromyces

hosts than of the Puccinia forms. It is just possible that these

two rusts are correlated but as to that we have made insufficient

study to say with certainty.”

Uromyces hedysari-paniculati (Schw.) Farl. Uredinia and telia

on Desmodium canadense. Westby.

A single spot of Aecidium fraxini Schw. (Puccinia periderm-

iospora (B. & T.) Arth.) on a leaf of Praxinus nigra was collected

at Caryville,

Puccinia graminis Pers. Telia on Agropyron tenerum. Mosinee.

In “Notes” II Puccinia gigantispora Bubak was recorded from

Glen Haven on a host that was not determined as between Anemone

virginiana and A. cylindrica. A collection of aecia and telia from

the same locality made in 1920 is on A. virginiana.

Dr. Arthur is of the opinion that the aecia and telia in these

collections are not genetically related but are aecia of Puccinia

agropyri E. & E. and telia of Puccinia anemones-virginianae

Schw.

Puccinia absinthii DC. Uredinia on Artemisia ludoviciana.

Prairie du Chien and Caryville.

Sclerotium deciduum. Davis occurred at Caryville in 1920 on

Aster paniculatus or an allied species.

260 Wisconsin Academy of Sciences, Arts, and Letters.

Additional Species

Not previously recorded as occurring in Wisconsin.

Plasmopara cuhensis (B. & C.) Humphrey. On Cucumis sativa

(cult.). Madison. The first collection of this parasite in Wiscon¬

sin, to my knowledge, was made in 1920. It is hoped that it was

merely casual.

Eocronartium muscicola (Fr.) Fitzp. (Typhula muscicola

(Pers.) Fr.) has been collected a number of times in Wisconsin

on unidentified mosses. Most of the collections were made by

Cheney in the northern part of the state.

Cordyceps clavulata (Schw.) Ell. & Evht. On scale insects

(Lecanium) on Fraxinus nigra and Ilex verticellata. Hannibal.

Yery abundant on the black ash in 1920; seen on but two plants of

Ilex. Immature specimens on Lecanium corni collected by J. G.

Sanders in 1913 are probably of this species. Apparently parasitic

on and destructive to the Lecanium.

Phyllosticta phaseolina Sacc. On Strophostyles helvola. Cass-

ville. The flattened pycnidia with dark and much thickened upper

wall and pale basal wall hr ve a leptostromatoid appearance.

Phyllosticta fraxinicola Curr. On Fraxinus pennsylvanica.

Durand. Spots along the midvein, angular, pale margined; spor-

ules 3-6x2%-3jt>i, smoky hyaline.

On looking over a collection of Cercospora dioscoreae E. & M. one

of the leaves was found to bear Phyllosticta dioscoreae Cke. Many

of the sporules are two celled.

Marssonina thomasiana (Sacc.) Magn. On Evonymus atropur-

pureus. Glen Haven. In the Wisconsin collection the leaves are

largely unspotted but pale spots appear late and red bordered

ones are numerous. The subcuticular acervuli are mostly epi-

phyllous, quite variable in size and soon naked. The sporules

are 17-23x7-14/x, the lower cell narrower. In the description of

the variety or “subspecies’’ fautreyana the width of the sporules

is given as 4/x, but in Yestergrens Micromycetes rariores selecti

1240, said to have been submitted to Saccardo, it approximates

that of the sporules in the Wisconsin collection.

Davis — Notes on Parasitic Fungi in Wisconsin — IX. 261

A parasite that is probably conspecific with Septogloeum con¬

volvuli Ell. & Evht. was collected on Convolvulus spithamaeus

at Tomahawk August 22, 1919. The small, circular, pale brown

spots are clustered on dead or dying leaf areas. The hyaline,

straight, cylindrical sporules are 24-36x3-3%ju, becoming triseptate.

Ramularia tenuis n. sp.

Spots yellowish, becoming black with age, subcircular to angular,

immarginate, 2-lOmm. in diameter ; conidiophores epiphyllous,

densely fasciculate from a prominent stromatoid base, hyaline,

subulate to cylindrical, simple, continuous, usually straight, apex

sometimes oblique, 10-20x2-3%ja, conidia hyaline, cantenulate,

usually straight, fusoid to cylindrical, acute, continuous to tri¬

septate, 7-37xl-3/x. On leaves of Solidago latifolia. Holcombe,

Wisconsin. August 9, 1920. This may prove to be a form of

Cercosprella reticulata Pk.

Botrytis epichloes Ell. & Dearn. On Epichloe typhina growing

on Glyceria nervata. Hannibal.

Fusicladium ejfusum Wint. On Cary a cordiformis at Muscoda

on both sides of the Wisconsin river.

In these collections are spots 2-4mm. in diameter which are

light brown above but much darker and less regular in outline

below and also dark dead areas following the veins as well as

indefinite marginal areas; the conidiophores are hypophyllous,

septate, usually curved or undulate, 40-140x3ju, in places swollen

to 5/x thick ; the conidia are obovate to fusoid or amygdaliform, sub¬

truncate at base, continuous, 13-23x6-7/x. The host tissue affected

becomes friable. Fusicladium caryigenum Ell. & Langlois is per¬

haps the same parasite.

Fusicladium cerasi (Rabh.) Sacc. (Cladosporium carpophilum

Thuem.) On fruit of Prunus americana. Caryville and Durand.

This was abundant on wild plums in August', 1920. Venturia

cerasi Aderh. is said to be the ascigerous stage (Ann. My col. 16:

81). Dr. G. W. Keitt informs me that the conidial form has been

observed on cherries and also on leaves in Door county.

Cercospora moricola Cke. On Morus rubra. Millville. Conidia

40-75x3-3%/a, becoming 3-6 septate; spots immarginate, indefi¬

nite below.

262 Wisconsin Academy of Sciences, Arts, and Letters.

Cercospora euonymi Ellis. On Evonymus atropurpureus. Dur¬

and.

In this collection the conidiophores are hyaline, 35-50x3/ji the

hyaline conidia cylindrical to obclavate-cylindrical, 30--50x3-4i/2/x.

Despite the Mucedinous character and the smaller conidiophores

and conidia it is unquestionably the parasite described under this

name by Ellis and issued in Fungi Columhiani, 2211. The white

spots with the broad dark purple border are conspicuous.

Cercospora teucrii Ell. & Kell. On Teucrium canadense. Cary-

ville.

Ustilago hypodytes (Schlecht.) Fr. On Stipa spartea. Meri-

dean.

Entyloma linariae Schroet. var., gratiolae n. var. Causing spots

which are orbicular, yellowish white, 1mm. or less in diameter

or larger and indefinite; spores broadly elliptical to globose with

epispore varying from nearly smooth to verrucose, 13-18/i, long. In

leaves of Gratiola virginiana. Cadott, Wisconsin, July 22, 1920.

The epispore in Entyloma linariae Schroet. is variously described

by authors as being smooth, irregularly angularly thickened or

as having a wavy outline because of low projections. In the col¬

lection referred to here some of the spores are studded with pale

verrucosities which are perhaps the remains of the gelatinized

hyphae. The form on this host is probably physiologically distinct

from the var. veronicae Wint. (E. veronicae Lagh.) as well as from

the type.

Heebarium of the University of Wisconsin, March, 1921

Notes on Distribution and Abundance

It is purposed to supplement the list of parasitic fungi occur¬

ring in Wisconsin with some notes on the distribution and fre¬

quency of the species in the state based upon the specimens pre¬

served in the Davis and the University of Wisconsin herbaria

and field observation. In attempting this one meets the fact that

infection is dependent on variables and hence a parasitic flora is

far from uniform. This seems to be especially true of those forms

which infect by means of zoospores.

Synchytrium cellulare Davis. Where first found, at Devils

Lake, (1913) the station, the bottom of a kettle hole, was very

Davis — Notes on Parasitic Fungi in Wisconsin — IX, 263

restricted and the parasite disappeared therefrom in 3 or 4 years.

It was not seen again until found at Babcock (1919) again very

restricted in a low spot in the river bottoms. In 1920 it was found

at Caryville, this time in more abundance, in an oxbow of Coon

creek where it flows through the bottom lands along the Chippewa

river and again in small quantity at Durand on the bank of the

Eau Galle river near its mouth. From the situations in which

this parasite has been observed it would appear that infection of

the host is favored if the water in which the zoospores develop is

impounded, thus preventing the carrying away of the infecting

agents.

Synchytrium scirpi Davis. The collections of this form have

been made only in the vicinity of Racine. It was first observed in

1904 southwest of the city and later northwest also, and appeared

in the same places in successive years. It has been found upon the

single species of host Scirpus atrovirens. It is possible that it is a

form of the following species.

Synchytrium aureum Schroet. To this species are referred col¬

lections on a considerable number of hosts of widely different

affinities and additions are made to the list yearly. When young

and especially on succulent hosts the sori, as seen through a hand

lens, are golden yellow, the color due apparently to the content of

yellow oil. When older this disappears and the sori, in section

are whitish. I have not seen an authentic specimen of Synchytrium

glohosum Schroet. but judging from the description some of the

Wisconsin collections could be placed in that species. My observa¬

tions lead me to suspect, however, that they represent a single

species in different stages and on hosts of different character.

The distribution is quite general as is indicated by the following

list of collections:

Clintonia borealis, Athelstane, July 24, 1915.

Geum virginianum, Berryville, July 3, 7 and 14, 1892; August

30, 1892; June 10, 1894; October 9, 1894. Some of these are

labelled ^^Geum alburn/^ (G. canadense.)

Geum strictum, Two Rivers, July 17, 1918.

Bubus triflorus, Turtle Lake, September 3, 1914; Solon Springs,

August 6 and 14, 1915; Two Rivers, July 17, 1918; Bruce,

September 4, 1918; Mosinee, September 1, 1919.

264 Wisconsin Academy of Sciences, Arts, and Letters.

Buhus hispidus, Millston, August 26, 27, 28, 1912; August 17, 19,

1915; July 21, 22, 1916; Athelstane, August 26, 1913; Wild

Eose, July 9, 1918.

Buhus villosus ( ? ) , Millston, September 26, 1912.

Viola pallens, Solon Springs, June 17 and 23, 1914; Athelstane,

July 27, 1915.

Viola pubescens, Barryville, July 3 and 9, 1892; Somers, June 22,

1902 ; Wild Eose, July 3, 1918.

Viola conspersa, Solon Springs, August 14, 1915.

Viola sp. indet., Solon Springs, August 7, 1915.

Hydrocotyle americana, Caryville, August 16, 1920.

Lysimachia terrestris, Millston, September 26 and 27, 1912,

Halenia deflexa, Solon Springs, August 7, 1915.

Lycopus uniflorus, Millston, September 26, 1912; Babcock, Sep¬

tember 11, 1919.

Pedicularis canadensis, Eacine, June 29, 1907.

Budheckia laciniata, Bruce, September 6 and 7, 1918.

Petasites palmatus, Prentice, August 18 and 19, 1918.

Prenanthes alba, Kenosha, July 4, 1907.

Synchytrium asari Arth. & Hoi. This maintained itself from

1897 to 1902 at a station in southeastern Wisconsin. In 1908 it

was observed in small quantity near Mellen in the northwestern

part of the state. I have not seen it since.

Synchytrium pulvereum Davis. Found only in Eusk county in

the north central part of the state. It was abundant at one station

subject to overflow from a creek. [This has since been collected

at White Lake and Sauk City.]

Synchytrium anemones (DC.) Wor. In Wisconsin, as elsewhere,

this well marked species is of wide distribution and varying

abundance. As is apt to be the case with common species it is not

as well represented in the herbaria as are some that occur less

frequently,

Synchytrium decipiens Farl. is the most frequent, abundant,

and conspicuous species of the group. By some mycologists only

Davis — Notes on Parasitic Fungi in Wisconsin — IX. 265

the first of these species is referred to Synchytrium, the last one

to Woroninella and all of the others to Pycnochytrium. If one may

borrow from the usage in the rusts and represent repeating spores

by II and resting spores by III they could be presented thus :

II, III Synchytrium.

Ill Pycnochytrium

II Woroninella

Physoderma menyanthis (DBy). Collected but once (1902)

near Found lake in the northern part of the state. But little was

seen.

Physoderma vagans Schroet. But a single scanty collection

has been made of this. It was found on bottom lands of the Wolf'

river. [Collected since at Spring Green.]

Cladochytrium maculare (Wallr.) has been collected only in

the southern portion of the state (Kenosha, Racine, and Dane

counties) where it is not frequent but sometimes rather abundant

and able to maintain itself in one station for several succesive

years.

Urophlyctis major Schroet. Observed only in the northeastern

part of the state in Kewaunee, Oconto and Marinette counties. It

is infrequent.

Urophlyctis pluriannulata (B.&C.) Farl. Not frequent but

sometimes locally abundant on Sanicula in both southern and

northern Wisconsin. On Zizia aurea it has been seen in but one

station. This was in 1907 and no trace of it was found on this

host at the station in subsequent years.

Albugo bliti (Biv.) Kuntze. Frequent and abundant on Amar-

anthus retroflexus and hybridus, often with development of abun¬

dant and conspicuous oospores ; infrequent on other species and on

Acnida.

Albugo portulacae (DC.) Kuntze. Not infrequent and some¬

times abundant.

Albugo Candida (Pers.) 0. Kuntze. Frequent and often abun¬

dant on various Cruciferae.

Albugo tragopogonis (DC.) S. F. Gray. Frequent, although on

some of the hosts it is but rarely seen. It is more frequent prob-

266 Wisconsin Academy of Sciences, Arts, and Letters.

ably on Cirsium muticum than on any other of the hosts. It is

sometimes very abundant on cultivated Tragopogon.

Phytophthora thalictri Wilson & Davis. This occurs throughout

the state and is collected every year. There are no collections on

Thalictrum dioicum. As T. revolutum occurs in but the south¬

eastern corner of the state T. dasycarpum is the usual host,

Phytophthora infestans (Mont.) DBy. In Wisconsin, as else¬

where, there is wide fluctuation in the frequency and abundance

of this parasite.

Premia lactucae Eegel. Frequent and abundant, especially on

Lactuca spicata. It has not been collected often on cultivated

lettuce in the open but is often abundant under glass.

Plasmopara humuli Miyabe & Takahashi. In southeastern and

southwestern Wisconsin. A scanty development has been seen

also in central Wisconsin. It appears to be indigenous. It was

collected at Caryville in the Chippewa valley in July, 1920, with

oospores.

Plasmopara pygmaea (Unger) Schroet. Frequent and abundant

on Hepatica and Anemone quinquefolia. The var. fusca (Pk.)

Davis has been collected in southern Wisconsin and on Hepatica

only. This form is peculiar in not developing conidia.

Plasmopara ribicola Schroet. Although some of the hosts occur

throughout the state this has been observed in the northern part

of the state only, the southernmost localities being in Shawano

county. Within its range it is not infrequent and fairly abundant

on the hosts of northern range Bibes prostratum and B. triste.

Plasmopara geranii (Pk.) Berk & DeToni. Widely distributed

on Geranium maculatum.

Plasmopara obducens Schroet. Frequent throughout the state

in spring, less so in summer when it is more often found north¬

ward.

Plasmopara acalyphae Wilson. This has been found only at a

station near Madison and in very small quantity at Caryville. It

is inconspicuous and as a rule very few conidiophores are de¬

veloped. [Since collected at Arena and at Lone Rock and a trace

at Oconto.]

Davis — Notes on Parasitic Fungi in Wisconsin — IX. 267

Plasmopara viticola (B.&C.) Berl. & DeToni. A common and

abundant species.

Plasmopara australis (Speg.) Swingle. Frequent and abundant

in the southwestern quadrant of the state.

Plasmopara viburni Pk. The only Wisconsin collections were

made at two localities in Marinette county in August, 1913.

Plasmopara cephalophora Davis. While this has been recognized

but r'ecently it has been collected on the banks of the Wolf, the

Wisconsin, and the Chippewa rivers and is probably not rare.

Plasmopara halstedii (Farl.) Berl. & DeToni. Frequent and

abundant. It is perhaps a congery of host-linked races. It usually

occurs upon a single species of host at any particular station.

Peronospora schleideni Unger. This is reported to have occurred

sporadically and sparingly about Madison.

Peronospora urticae (Lib.) DBy. This was recorded by Trelease

in the Preliminary List of Parasitic Fungi of Wisconsin (1884) as

having been collected at Kirkland (now Devils Lake) and La

Crosse. Dr. E. A. Burt kindly examined the Trelease hebarium

at the Missouri Botanical Garden and found a specimen from

“Kirkland.” No collections have been made since in Wisconsin.

Peronospora polygoni Thuem. This has been collected in the

southwestern part of the state along the Mississippi and lower

Wisconsin and Chippewa rivers and in Kenosha county in south¬

eastern Wisconsin.

Peronospora effusa (Grev.) Eabh. Frequent and abundant

especially on Chenopodium album.

Peronospora obovata Bon. A single collection made in 1911.

The host is rare in Wisconsin.

Peronospora silenes Wils. Collected at Necedah and Adams

only. Local.

Peronospora alsinearum Casp. Observed at Racine and Madison

only. Local and not permanently established.

Peronospora ficariae TuL Rather common throughout the state.

Peronospora corydalis DBy. Rare and local. It has been col¬

lected in the southern part of the state only.

268 Wisconsin Academy of Sciences, Arts, and Letters.

Peronospora parasitica (Pers.) Tul. Frequent and abundant

on various species of Cruciferae. It is probably a group of host-

linked races.

Peronospora potentillae DBy. Not infrequent on Agrimonia,

Geum and Potentilla and perhaps includes three races which are

host adapted.

Peronospora rubi Rabh. Infrequent. Most of the collections

were made in northern Wisconsin but in 1920 it was found along

the railroad at Blue Mounds where leaves infected by Caeoma

seemed to be more susceptible. One of the specimens from north¬

western Wisconsin is on leaves bearing Caeoma.

Peronospora trifoliorum DBy. This is rather frequent in alfalfa

fields but not destructive in Wisconsin. There is but a single

specimen each on Lupinus and Astragalus in the herbarium, both

from western Wisconsin.

Peronospora viciae (Berk.) DBy. This occurs rather frequently

in the pea fields but does not do much damage to the crop.

Peronospora viciae americana Davis. Local on Yicia americana

mostly in the western part of the state.

Peronospora chamaesycis Wils. This is probably more frequent

than the small number of specimens would indicate as it is not

conspicuous.

Peronospora floerkeae Kell. The collection made at St. Croix

Falls by Holway in 1904 is the only one that has been made in

Wisconsin.

Peronospora arthuri Farl. Not infrequent on Oenothera biennis

to which is seems to be confined. It usually shows evidence of

systemic infection of the host.

Peronospora alta Fckl. This is a frequent and abundant species.

Peronospora calotheca DBy. Not frequent but sometimes locally

abundant. All of the specimens are from the southern half of

the state.

Peronospora leptosperma DBy. This is a rare species in Wis¬

consin, being represented only from the following collections:

Racine, 1897 ; Berryville, 1900 ; Shiocton, 1917.

Davis — Notes on Parasitic Fungi in Wisconsin — IX. 269

Basidiophora entospora Roze & Cornu. This is probably more

frequent than the specimens in the herbaria would indicate. It

was especially abundant about Madison in 1920.

Sclerospora graminicola (Sacc.) Schroet. This again is doubt¬

less more frequent than the few Wisconsin specimens would in¬

dicate.

Protomyces andinus Lagh. This was frequent and abundant on

Bidens before 1911, but the only collections since were made along

the Mississippi river. On Ambrosia it has continued to be frequent

and sometimes abundant.

Herbarium of the University of Wisconsin,

Madison, Wisconsin, April, 1921

NOTES ON PARASITIC FUNGI IN WISCONSIN— X

J. J. Davis

The season of 1921 was characterized by high temperature and

low humidity and was consequently unfavorable for the develop¬

ment of most fungous parasites. The low stage of water in the

Wisconsin River, however, gave opportunity to explore bottom

lands that are ordinarily difficult of access.

After one’s conceptions of generalities have changed, there is a

lag in the application of such changes to particulars. When the

writer began the collection of data on the parasitic fungus flora

of Wisconsin there was in his mind a conception of such a flora as

something fixed, static; given time and application it could be

fully set forth. The fact of quantitative variation was quickly

brought home to him and was expressed in his first contribution.

More slowly the conception of vegetation as dynamic and mutable

has come to be applied to a special group in a local flora. The

records are then seen as datum points having a time as well as a

space value. This conception increases rather than lessens their

value but shows no finality as a goal. It does not however at all

lessen the importance of completeness in the record.

The downy mildews as they occur in Wisconsin are interesting

from an evolutionary viewpoint. Typically they bear two kinds of

spores, summer dispersion conidia and winter resting oospores, and

cause local infections. In many species, however, general infec¬

tion takes place and allows overwintering as mycelium. Such spe¬

cies show a tendency toward suppression of oospores and abun¬

dance of conidia. In Plasmopara pygmaea (Ung.) Schroet. each

of these directions of change seems to have been followed. On

Hepatica and Anemone quinquefolia and A. canadensis races pro¬

ducing local infections and both kinds of spores ; on Anemone quin¬

quefolia a race with general infections and suppression of oospores ;

on Hepatica acutiloha local infections, abundance of oospores and

no conidia. The latter is what Peck described under the name

Protomyces fuscus but which I have designated Plasmopara pyg-

272 Wisconsin Academy of Sciences, Arts, and Letters.

maea var. fusca (Pk.). I take it that the state with local infec¬

tions and both spore forms is the older and that the others have

been derived therefrom.

Synchytrium scirpi Davis on Scirpus atrovirens previously

known only from the vicinity of Racine was collected at Oconto in

September, 1921, mostly in an abortive state, presumably because

of the hot, dry summer. In examining sections of the leaves what

appeared to be a Sphaerulina was observed and the following notes

made: Perithecia sparse, innate, black, globose, 85-lOOjtJi in diam¬

eter; asci sessile, fusoid-cylindrical, 60-70xl6-18/x; spores long

fusoid, slightly curved, hyaline, obtuse, 5-septate, 35-40x5-6/a;

paraphyses none; The inconspicuous perithecia are too few to

warrant taking the material as a type. [In 1922 Synchytrium scirpi

was collected on the same species of host opposite Sauk City. There

is also a specimen from Little Suamico.]

Sept or ia rubi West, was found by Roark to have an ascigerous

stage for which he proposed the name Mycosphaerella rubi (Phyto¬

path. 11 :329, [1921]. Although the Septoria is abundant through¬

out the state he was able to find the Mycosphaerella only in Door

county which lies between Green bay and lake Michigan.

Piggotia vaccinii Davis (“Notes” IX, p. 436, fig. 3) is doubtless

conspecific with Leptothyrium conspicuum Dearn. & House (A. Y.

State Museum, Report of the Botanist for 1919, p. 37, [1922] and is

antedated thereby.

A parasite of Gentiana Andrewsii was collected in Wisconsin

thirty odd years ago and sent to Ellis who identified it as Depazea

gentianaecola Pr., but referred it to Phyllosticta (see N. A. Phyl-

lostictas No. 176) and it was so recorded in the supplementary list.

Baeumler in 1889 recorded as Leptothyrium, a fungus on Gentiana

which he supposed to be the same as the one treated by De Can¬

dolle and Pries and used the specific name proposed by De Candolle.

In compiling the provisional list there was no time to investigate

such matters but simply to follow the Tom Johnson rule, “decide

at once and be right half the time. ’ ’ On the principle that things

that were equal to the same thing were equal to each other I used

Baeumler ’s binomial. Dr. Brenckle sent me a specimen from North

Dakota and I wrote him that it was the fungus recorded in the

provisional list under the name Leptothyrium gentianaecolum

(DC.) Baeuml. and he issued it in Fungi Dakotenses under that

Davis — Notes on Parasitic Fungi in Wisconsin — X. 273

name. This fall I took the matter up and find that it can not be

referred to Leptothyrium but that it agrees with Allescher’s de¬

scription of Asteroma gentianae Fckl. (Bahh. Krypt. fl. Pilze:

7: 464). I have seen no specimens of Sphaeria gentianaecola DC.,

Xyloma gentianaecola DC., Depazea gentianaecola (DC.) Fr. and

do not know their relationship to our parasite. Neither have I

seen a specimen of Fuckel’s Asteroma gentianae. As matters

stand now, I am inclined to use Fuckel’s name provisionally, as

the fungus has the characters of Asteroma. [In 1922 this was

collected on Gentiana puherula at Arena. Sometimes the proximal

portion of the pycnidial wall is thin but distinct while the distal

portion is thick and black while other pycnidia have a thick, black

wall throughout.]

Fusidium pteridis Kalchbr. was recorded in Trelease’s Prelimi¬

nary List of parasitic Fungi of Wisconsin and was copied there¬

from into the provisional list. In ‘‘Notes” V, p. 701, a parasite of

Pteris was recorded under the name Gloeosporium leptospermum

Pk. which is the same fungus. Gloeosporium pteridis Hark, as

represented by Griffith’s West American Fungi 324 and 324a and

Jackson’s No. 1688 is the same species. The synonymy is

Fusidium pteridis Kalchbr.

Gloeosporium pteridis Hark.

Gloeosporium leptospermum Pk.

Gloeosporium pteridis (Kalchbr.) Kabat & Bubak.

It is said to be a sporuligerous state of Cryptomyces pteridis (Eeb.)

Rehm. (Gloeosporium ohtegens Syd. Ann. Mycol. 2:172), C. F.

Baker, Pacific Slope Fungi 3757 can hardly be other than a pro¬

fusely developed state of the same parasite. Another parasite of

Pteris was recorded in the provisional list under the name Mars-

sonina necans (E. & E.) Magn. The sporules of this fungus are

developed in a pycnidium and it is referable to Ascochyta and does

not differ from Ascochyta pteridis Bres. as represented in Krieger ’s

Fungi Saxonici 989. The synonymy is :

Gloeosporium necans Ell. & Evht. Journ. Mycol. 4: 104, (Octo¬

ber, 1888).

Marsonia necans (E. & E.) Sacc.

Marssonina necans (E. & E.) Magn.

274 Wisconsin Academy of Sciences^ Arts, and Letters.

Ascochyta pteridis Bres. Hedwigia, 1894.

Ascochyta necans (E. & E.) n. comb.

In specimens of Bamularia desmodii Cke. on Dcsmodium illino-

ense the penicillate conidiophoral fasciculi sometimes exceed lOOfi

in length.

In examining a collection of Cerosporella cana Sacc. on Erigeron

annuus it was observed that the conidiophores ranged up to 100/^

in length.

Specimens on leaves of Crataegus collected in 1890 were re¬

ferred to Mr. J. B. Ellis for determination who reported as follows :

‘‘9039. Phleospora oxyacanthae (Kze. & Schm.) I think it must

be, but your specimens are much better than any I have in my

European collections. ’ ’ At the close of the letter he wrote : “ I have

had to examine the things in great haste but I think you will find

them correct.^’ In the Supplementary List the fungus was re¬

corded under this name and carried from there to the Provisional

List where by a slip the specific name was given as “crataegi”.

This was corrected in “Notes” III p. 254 and some remarks on the

character of the parasite added which indicated that it could not

be a Phleospora. To the notes there given, I would add that in

some specimens the conidia have divided at' some of the septa re¬

sulting in shorter conidia which sometimes becomes thicker,

15-45x3-7/x. This is quite different from Phleospora oxyacanthae

(Kze. & Schm.) Wallr., but appears to be Cercosporella mirahilis

Pk., for a specimen of which I am indebted to Dr. House which,

however, lacks the differentiated conidiophores of Cercosporella.

The first Wisconsin collection of Cercospora on Smilax was made

at Racine. It was a form with small spots and preponderance of

the dark border and was referred to Cercospora mississippiensis

Tracy & Earle. Subsequent collections lead me to believe that

there is but one species on Smilax in Wisconsin and that it is

C. smilacis Thuem., as described and figured by Peck {33d Report,

p. 29, figs. 1-3). In different collections the spots vary from 1 to

8 mm. in diameter, the border varies in width and intensity of

color, the conidiophores are longer or shorter (30-83x4|u) and the

conidia are variable in size (30-115x4-5/x) and depth of color.

They may be attenuate or subcylindrical and obtuse and, with the

conidiophores, vary in septation. Fungi Columhiani 2208 labeled

Cercospora mississippiensis Tracy & Earle I have referred to C.

Davis — Notes on Parasitic Fungi in Wisconsin — X. 275

smilacis Thuem. Saccardo however thought the parasite described

and figured by Peck to be distinct from Cercospora smilacis Thuem.

and called it C. smilacina Sacc. (Michelia 2: 364). I have not seen

European specimens but Peck accepted von Thuemen’s descrip¬

tion as applying to his species {33d Report, explanation of plate 2,

footnote).

A collection of Cercospora davisii E. & E. on Melilotus alha made

at Madison in June 1921 (Bensaude, McFarland, & Davis) bears

conidiophores up to 140/x in length. Evidence accumulates that

length of conidiophores and conidia in this and similar genera as a

specific character is to be used with caution. A Cercospora occur¬

ring on dark areas on branches of Melilotus alha at Grays Mills is

referred to this species. The conidia seen were only about 3jtx thick.

Cercospora epigaeina Davis (Trans. Wis. Acad. 16: 758) is evi¬

dently not distinct from C. epigaeae Ell. & Dearn. which is the

older name.

Examination of a collection of Cercospora saniculae Davis from

Blue Mounds shows that when not crowded the conidiophores are

not always straight, that they sometimes occur on the upper leaf

surface and that the longest ones may attain a length of 60/x.

Cercospora platyspora Ell. & Holw., is doubtfully distinct from

Cercospora sn E. & E. and from Fusicladium depressum (B. & Br.)

Sacc. Specimens on Angelica were issued in Fungi Columhiani

1924 under the name Didymaria platyospora (Ell. & Holw.), but

F. Col. 4230 on Taenidia integerrima is labeled Fusicladium de¬

pressum (B. & Br.) Sacc.

In a collection of Cercospora stomatica Ell. & Davis made at

Woodman, July 4, 1921, the conidia are narrow (about 3/x) and of

nearly uniform diameter throughout. A result perhaps of the hot,

dry season.

Doassansia ranunculina Davis which had not been seen in Wis¬

consin for upwards of 20 years was collected at Shiocton in Sep¬

tember, 1921. Although the host was abundant there was but very

scanty development of the parasite.

Puccinia zygadeni Trel. is merged into P. atropuncta Pk. & Cl.

in North American Flora.

276 Wisconsin Academy of Sciences, Arts, and Letters.

Additional Hosts

Not previously recorded as bearing the fungi mentioned in Wis¬

consin.

It is customary in collecting in Wisconsin to find each year sori

of Synchytrium on an additional host at a single station and in

small quantity and they have usually been referred to Synchytrium

aureum Schroet. Such a collection was made July 25, 1921, on

leaves of Acalypha virginica in the Wisconsin Kiver bottom lands

at Lone Rock. In this collection the sori are first pale yellow be¬

coming castaneous with age. The galls are hypophyllous, discrete,

hemispherical, but little larger than the sori which are globose to

elliptical, 125-17 5 fi in diameter with wall about 5/>t thick.

[Collected in 1922 at Arena and Prairie du Sac.]

Peronospora ficariae Tul. On Eanunculus recurvatus. Ridge¬

way.

Peronospora calotheca DBy. This was collected in small quan¬

tity on Galium asprellum at Madison in November, 1920.

Basidiophora entospora Roze & Cornu. On Aster lateriflorus.

Muscoda.

Erysiphe cichoracearum DC. On Vernonia fasciculata and

Helenium autumnale. Muscoda.

Phyllactinia corylea (Pers.) Karst. On Betula nigra. Richland

County opposite Muscoda.

Phyllachora graminis (Pers.) Fkl. On Elymus hrachystachys.

Richland County opposite Muscoda.

A Phyllachora forming black patches with effused ascomata on

leaves of Panicum virgatum has been collected at Muscoda. Pro¬

visionally it is labeled P/i. graminis panici (Schw.) Shear although

it differs widely in macroscopic appearance from specimens on

other species of Panicum.

Phyllachora amhrosiae (B. & C.) Sacc. {Physalospora amhrosiae

E. & E.). On Amhrosia psilostachya. Muscoda.

Pseudopeziza singularia Pk. On Ranunculus septentrionalis.

Blue River and Iowa County opposite Lone Rock.

Davis — Notes on Parasitic Fungi in Wisconsin — X. 277

Phyllosticta apocyni Trel. On Apocynum androsaemifolium,

Ridgeway.

Ascochyta pisi Lib. On Vida angustifolia segetalis, Barneveld

and Ridgeway.

Ascochyta lophanthi lycopina Davis. This has been collected on

Lycopus virginicus and it is quite possible that other collections are

on this species of host.

Darluca filum (Biv.) Cast. On Puccinia asteris growing on

Aster Tradescanti, Blue River. Two telial hosts were given in

^ ‘ Notes VI, p. 707.

Septoria annua Ell. & Evht. On Poa annua. Black Earth.

(McFarland & Davis.)

Septoria caricinella Sacc. & Roum. A specimen on Car ex chordo-

rrhiza from Lost Lake, Vilas County (July 4, 1901) is referred to

this species.

Septoria polaris Karst. Specimens on Ranunculus septentrion-

alis from Richland County opposite Muscoda are provisionally re¬

ferred to this species. The sporules are 20-30xl-l%ju-.

Septoria oenotherae West. On Oenothera rhombipetala. Mus¬

coda.

Septoria sii Rob. & Desm. On Cicuta hulhifera. Oconto.

Septoria solidaginicola Pk. On Solidago patula. Cecil. Aster

lateriflorus. Woodman. Of the latter collection it was noted ‘ ‘ not

abundant on this host and some of the spots atypical”.

Septoria atropurpurea Pk. On Aster paniculatus. Lone Rock.

The strongly curved sporules range up to more than 100xl%-2/x.

Phleospora ulmi (Fr.) Wallr. On TJlmus fidva. Richland

County opposite Boscobel. Sporules mostly about 30x7ja.

A Marssonina on leaves of Salix lucida collected at Shawano I

do not distinguish from forms on Populus and have labeled it

Marssonina populi (Lib.) Magn. The aeervuli are amphigenous

but more abundant and better developed below where they have a

resinous appearance. The curved sporules are 13-17x3-4/ji. A

collection on Populus halsamifera from Little Suamico has hypo-

phyllous subcuticular aeervuli with sporules ll-17x3%-5|a.

278 Wisconsin Academy of Sciences, Arts, and Letters.

Marssonina pote^itillae (Desm.) Magn. On Fragaria virginiana.

Crandon and Little Suamico. Of the latter collection it was noted

— Immaculate, spornles 17-24x3%-6/x. On Potentilla anserina.

Oconto.

Septocylindrium concomitans (Ell. & Holw.) Hals. On Bidens

vulgata puberula. Wauzeka.

Bamularia uredinis (Voss) Sacc. On Salix amygdaloides.

Oconto.

Bamularia fraxinea Davis. On Fraxinus americana ( ? ) Gays

Mills and Blue River. This parasite has been seen only in river

bottom lands.

Bamularia effusa Pk. On Gaylussacia haccata. Ridgeway. Ap¬

parently causing defoliation.

Bamularia dispar Davis. A collection on Eupatorium urticae-

folium from Crandon is referred to this species.

In a specimen on Solidago serotina from Gays Mills the conidio-

phores are mostly subulate, 10-20x3ja, the conidia fusiform to cylin¬

drical, 7-36x2/x. This was referred to Bamularia virgaureae

Thuem.

Scolecotrichum graminis Fkl. On Dactylis glomerata. Madi¬

son. (Bensaude, McFarland & Davis.)

Cercospora diffusa Ell. & Evht. On Physalis heterophylla.

Oconto.

Cercospora antipus Ell. & Holw. On Lonicera Sullivantii. Wer-

ley. The brown, tufted conidiophores are 40-70x3jut.

Ustilago striaeformis, (West.) Niessl. On Agrostis alha. Madi¬

son. (W. H. & J. J. Davis) Collected also on Poa pratensis at

Madison by W. H. Davis.

Urocystis agropyri (Preuss) Schroet. On Agrostis alba. Madi¬

son. (W. H. Davis)

Entyloma ranunculi (Bon.) Schroet. Conidiophorous material

on Thalictrum dasycarpum collected at Oconto September 8, 1921

Davis — Notes on Parasitic Fungi in Wisconsin — X, 279

I can not distinguish from this species either in the field or in the

herbarium. Typical material of E. thalictri Schroet. was collected

in the same locality.

TJromyces appendiculatus (Pers.) Lk. Uredinia on Strophostyles

helvola. Kichland County opposite Blue River.

Puccinia graminis Pers. Uredinia on Poa annua. Black Earth.

(McFarland & Davis). Spores 17-24xl3~17ju.

Puccinia peridermiospora (E. & Tr.) Arth. On Fraxinus penn-

sylvanica and var. lanceolata. These are probably the most sus¬

ceptible hosts of the Aecidium in Wisconsin.

Puccinia polygoni-amphihii Pers. Uredinia on Polygonum Per-

sicaria. Oconto.

Pucciniastrum myrtilli (Schum.) Arth. Uredinia on Gaylus-

sacia baccata, Oconto.

Aecidium dicentrae Trel. the type locality of which is in Wis¬

consin has been shown by Mains to be the aecial stage of a

Melampsoraceous rust on Laportea canadensis (Am. Journ. Bot.

8: 445) which has been collected in Wisconsin at Hannibal, Jump

River, Holcombe and Blue Mounds. For this Dr. Mains proposes

the binomial Cerotelium dicentrae (Trel.) Mains & Anderson, a

name which does not accord with the usage followed in this series

of notes. [See Saccardo: De Diagnostica et Nomenclatura Myco-

logica; admonita quaedam (Annates Mycologici 2:197.) English

translation in Journal of Mycology 10:111-2.] I am therefore

using Cerotelium urticastri Mains (loc. cit. 451).

Additional Species

Not previously recorded as occurring in Wisconsin.

In August 1892 a Synchytrium on Ranunculus recurvatus was

collected in Kenosha County and the following description written

and filed with the specimen. As it was not seen again, however,

some doubt arose as to its being distinct and it was recorded under

S. aureum Schroet. in the provisional list which is doubtless an

aggregate species as treated in Wisconsin. In 1921 it was found

on Ranunculus septentrionalis at the base of the Wisconsin River

bluff in Iowa County opposite Spring Green readily recognizable

as being the same as the original collection. I therefore publish

the description.

280 Wisconsin Academy of Sciences, Arts, and Letters.

Synchytrium cinnamomeum n. sp. Galls cinnamon brown, hemi¬

spherical to obtusely conical, scattered or aggregated, frequently

confluent, 125-150/x in diameter; resting sori solitary, globose to

elliptical to pyriform, 42-66/x in the longer diameter; wall brown,

contents brown black, granular. On petioles and leaves of Ranun¬

culus recurvatus, Somers, Wisconsin, August 13, 1892 (type).

Ranunculus septentrionalis, Iowa County opposite Spring Green,

July 20, 1921. Readily recognized by the brown color which sug¬

gests rust.

Synch3rtrmm nigrescens n. sp. Sori hypophyllous, scattered,

subepidermal, at flrst pale yellow with abundant oil content, be¬

coming black with content in part black and granular, spherical to

ovoid 80-180/x; wall thin, homogeneous, chitinous, black by re¬

flected, fuscous by transmitted light, outer surface smooth or

minutely tuberculate. But slight prominences are produced, the

sori often extending through to the upper epidermis without caus¬

ing hypertrophy. On Aster laterifiorus on bottom lands of the

Wisconsin river at Spring Green, Lone Rock and Blue River.

Plasmopara illinoensis (Farl.) n. comb. On Parietaria penn-

sylvanica. This was described by Farlow {Bot. Gaz. 8: 332 [1883] )

from collections made by Seymour at Quincy and Camp Point on

the Mississippi river in southern Illinois. No further collections

seem to have been made. Guy West Wilson gave a new description

and referred it to his proposed genus Rhysotheca {Bull. Torr. Bot.

Cluh 34: 407 [1907]). In 1921 it was collected at Blue Mounds,

Ridgeway, Fennimore, Werley and Woodman. A collection from

Blue Mounds made July 9 contains globose oospores 23-SOix in

diameter with wall thick filling the rather thin-walled oogonia.

Assuming that southern Wisconsin is the northern limit of this

species its development was probably favored by the abnormally

hot season of 1921.

Phacidium taxi Fr. On Taxus canadensis. Crandon. The ex-

ciple is thick and black and ruptures irregularly in the center.

The asci are clavate-cylindrical, about 50x6ja. No mature spores

were seen.

Claviceps nigricans Tul. Sclerotia on Eleocharis palustris.

Sturgeon Bay (J. E. Sanders), Madison (J. R. Heddle).

Phyllosticta congest a Heald & Wolf {Mycologia 3: 8). On

Prunus pennsylvanica. Devils Lake (C. E. Owens).

Davis- — Notes on Parasitic Fungi in Wisconsin— X, 28 J.

Phyllosticta pyrolae Ell. & Evht. Collected in small quantity

and not quite mature on leaves of Pyrola elliptica at Blue Mounds.

Phyllosticta steironematis Beam. & House. On 8teironema cilia-

tum. Lone Eock.

Phyllosticta verbascicola Ell. & Kell. On Verhascum Thapsus.

Barneveld.

Phoma alliicola Saec. & Eoum. A collection on Allium canadense

from Madison is referred to this species. The ostiole is often gap¬

ing, the sporules 4--6x2~3fi.

Macrophoma arens n. sp. Pycnidia black, scattered, subepider-

mal, globose, 130”150ft in diameter ; sporules narrow ovoid, becom¬

ing subfuligenous, 27xl0/x at maturity. On more or less of the

distal portion of leaves of Koeleria cristata which become sere and

involute. Boscobel, Wisconsin, July 5, 1921.

Asteromella astericola n. sp. Pycnidia epiphyllous on indefinite

purple areas in compact orbicular groups, black, globose, 100-1 65ju,

in diameter ; wall parenchymatous of dark firm cells 6-10/x in diam¬

eter; sporules sessile (?), hyaline, terete to fusoid-cylindrical,

mostly straight, 20x30x3-4/x. On Aster lateriflorus, Blue River,

Wisconsin, August 3 and 4, 1921. It may be that this is a better

developed form of Asteromella asteris Pk. {Report for 1912, p. 38).

Stagonospora albescens n. sp. Spots orbicular, sordid white,

%-2 mm. in diameter surrounded by a broad indeterminate reddish

brown border; pycnidia few, innate, dark brown, globose, thick

walled, about 150/i, in diameter; sporules hyaline, fusoid-cylindri¬

cal, straight or sometimes curved, 5-7 septate, 45-67xl0-13ju. On

living leaves of Carex tribuloides. Muscoda, Wisconsin, October

1920. Macroscopically this resembles Septoria caricinella Sacc.

sufficiently to have been mistaken for it in the field. The contents

of the cells or cytoplasmic divisions are homogeneous.

May 13 and 19, 1921, spots were observed on leaves of a few

plants of Melilotus alba at a station near Madison. The following

notes have been made from these collections : Spots definite, circular

to elliptical to irregular, argillaceous with a paler center, 1-10 mm.

in diameter; pycnidia in the paler area, hypophyllous, exception¬

ally epiphyllous also, brown, globose to lenticular, ostiole circular

with a dark margin about 30/x across, 135-165/a in diameter;

282 Wisconsin Academy of Sciences , ArtSy and Letters,

sporules cylindrical, hyaline, usually straight, sometimes slightly

curved or bent, 1-3 septate, about 20 (‘‘13-23”, “ 10-27 ”)x3-3 %/a.

I have labeled it Stagonospora meliloti (Lasch) Petr.

Three collections on leaves of Acer Negundo were made in 1921

that were referred to Septoria negundinis Ell. & Evht. They are

evidently members of the acericolous group referred to in “Notes”

I, pp. 81-2. In the collection from Werley the round arid spots

are but 2 to 3 times the diameter of the usually solitary pycnidia

and the curved sporules 32-40x1 %-2%/a indistinctly 3-septate.

This is much like Septoria saccharina Ell. & Evht. The collection

from Barneveld is similar with slightly greater range in spore

length and septation not apparent. In the collection from Madison

(Bensaude, McFarland & Davis) the spots are pale argillaceous

with a narrow raised darker margin, amphigenous, circular to

angular in outline, 1-3 mm. in diameter, often confluent ; pycnidia

one to few on the spot, hypophyllous, subepidermal, globose to

lenticular, up to 180/a in diameter ; sporules hyaHne becoming curved

and 3-septate, 25-40x1%-2%/a. A collection made at Galesville

in 1914 and recorded in “Notes” III, p. 264, as Septoria acerella

Sacc. but belonging with these is similar, the sporules being

23-33x1%-2%/a becoming curved and triseptate. In this collec¬

tion indeflnite leaf areas upon which the small spots are numerous

become dead and brown. In examining this epiphyllous subcuticu¬

lar acervuli were seen bearing oblong hyaline sporules 10-13x4/a.

These collections are evidently Septoria acerella Sacc. as treated

by Ellis in Septorias of North America No. 160 {Journ. Mycol.

3: 79) but subsequently described by Ellis & Everhart as a new

species for which the name Septoria negundinis was proposed in

the Proceedings of the Academy of Natural Science, Phila. for

1893, p. 165.

Of a Septoria on Caeoma-infected leaves of Buhus allegheniensis

collected at Madison June 2, 1921, the following notes were made:

Spots epiphyllous, circular to angular, subolivaceous, immarginate,

1-5 mm. in diameter, often confluent; sporules hyaline, usually

curved, continuous, 30-50x1-2/a. While it is possible that this is

a form of Septoria ruhi West, modified by the character of the sub¬

stratum it has been kept separate under the name Septoria comitata

n. sp. ad interim.

Septoria lycopi Pass. Collected on Ly copus uniflorus on the

river bottoms opposite Muscoda in October 1920. This was a dry

Davis — Notes on Parasitic Fungi in Wisconsin — X. 283

season and the sporules are not well developed, being but about

SOxl^jLt.

Cylindrosporium caryogenum Ell. & Evht. On Carya cordi-

formis, Werley and Woodman. This bears some resemblance to

Microstroma in the field.

I

Of a collection on Aster sagittifolius obtained at Woodman July

4, 1921, and referred to Septoria solidaginicola Pk. the following

notes have been made : Spots angular, limited by the veinlets, be¬

coming confluent, reddish brown to pale brown, without halo,

2-7 mm. long, 1-2 mm. wide; pycnidia epiphyllous-innate, globose

to ovoid, often with a black ring around the ostiole which is some¬

times conical, 60-90/a in diameter; sporules 30-36x1-1 1/^/a. Sep¬

toria angularis Tharp to which, judging from the description, this

bears resemblance was described as having pycnidia 75-80x100-200/a

and sporules 35-50x3/a {Texas Parasitic Fungi, Mycologia 9: 121).

The name is antedated by Septoria angularis Dearn. & Barth.

{Mycologia, 8: 103) on Solidago latifolia (Ontario, Dearness) in

which the spots are said to be limited “when the pycnidia are well

developed by a narrow, raised, sharply defined border’’. The col¬

lection on Aster sagittifolius here referred to is quite similar to

this as represented in Fungi Columhiani 4875. There seems war¬

rant for the suspicion expressed by the authors that this may be

Septoria fumosa Pk.

Phleospora anemones Ell. & Kell. On Anemone virginiana. Iowa

County opposite Lone Rock. This forms a well-developed pycni-

dium and might be referred to Septoria without doing violence.

Cylindrosporium guttatum Wint. What is perhaps this species

was collected on Hypoxis hirsuta at Blue Mounds bearing lunate

sporules but 18-24x2/ji. From the examination it was thought that

the short sporules might have been formed by division as in the

acervuli ( ? ) were found longer straight ones.

Cylindrosporium toxicodendri (Curt.) Ell. & Evht. On Bhus

toxicodendron. Barneveld and Lone Rock. Septoria irregularis

Pk. as represented by a specimen collected by Peck at Bolton Land¬

ing, N. Y., is the same fungus.

Septogloeum querceum n. sp. Spots or areas indefinite, becom¬

ing mottled brown; acervuli hypophyllous, subcuticular, sporules

sessile, hyaline, falcate, 7-9 septate, 35-50x5-7/a. On languishing

284 Wisconsin Academy of Sciences, Arts, and Letters,

leaves of Quercns bicolor. Blue River, Wisconsin, August 2, 1921.

Exceptionally straight conidia occur while some might perhaps be

called rostrate.

Fig. 1. Vertical section of acervulus of Septogloeum querceum n. sp. on

leaf of Quercns bicolor with sporules in various stages of development. Drawn

by E. M. Gilbert with the aid of camera lucida.

[This was collected again in 1922 and was grown on nutrient

agar by Miss Helen Johann and in addition to the conidia there

was development of pycnidia with spermatioid contents. Later a

collection was made at Blue River that no longer bore conidia on

the lower surface but with pycnidia on the upper surface with

spermatioid, imperfectly developed, contents. Some of these leaves

were kept in a moist chamber by Miss Johann and developed hya¬

line, globose, delicate walled sporules 3-4/a in diameter. When

germinating in water these sporules developed first a bud and then

bud and sporule developed each a germ tube. Often the bud was

nearly and sometimes quite as large as the sporule. Later (Sept.

2) a collection was made at Arena with similar sporules in the

pycnidia and acervuli on the lower leaf surface. For the purpose of

filing I have designated this pycnidial state Phyllosficta quercea

n. sp. ad interim.

In circular groups or later on orbicular brown spots or irregular

areas; pycnidia epiphyllous, immersed in the palisade layer, dark

brown, globose to elliptical, the longer axis parallel with the pali¬

sade cells, 60-100/a in diameter; sporules hyaline, globose, delicate

walled, 3-5/a in diameter. On leaves of Quercns bicolor. Arena and

Blue River, Wisconsin.

From observation of the cultures Miss Johann is of the opinion

that the Septogloeum and the Phyllosticta represent distinct organ¬

isms, the mycelium of the former growing very slowly, that of the

latter much more rapidly. By placing leaves in a moist chamber

she brought about the development of sporules in which one or

Davis — Notes on Parasitic Fungi in Wisconsin — X. 285

two of the cells is divided by a vertical septum. She is quite sure

that these are a further development of the sporules represented

in fig. 1.]

Fig. 2. Selected sporules from leaf of Quercus Mcolor kept in a moist cham¬

ber. Drawn by Helen Johann with the aid of camera lucida.

Bamularia tanaceti Lind. On Tanacetum vulgare. Fennimore.

Epiphyllous tufts sometimes occur.

Cercosporella celtidis (Ell. & Kell.) n. comb. {Bamularia

celtidis E. & K. Journ. My col. 1: 75). On Celtis occidentalis.

Bank of Wisconsin river opposite Boscobel. Well characterized

by the short conidiophores. The slender, filiform conidia up to 75ju,

in length are more nearly of the Cerosporella type. To their de¬

scription the authors appended the statement ‘ ‘ approaches Cercos-

pora’k

Of a collection on ash leaves in 1921 the following notes were

made: Spots epiphyllous, orbicular, sordid white with a reddish

brown to black border, 1-2 mm. in diameter ; conidiophores fuscous,

single or in small fascicles, more or less irregularly undulate and

finely denticulate toward the apex, simple, septate, 40-100x3/x;

conidia brown, uniseptate, about 13x4/x. On leaves of Fraxinus

pennsylvanica. Blue River. This has been provisionally labeled

Cladosporium simplex Schw. of which I have not seen an authentic

specimen. It may be that the Cladosporium is not the cause of

the spots.

Cladosporium astericola Davis. ' On leaves of Solidago serotina.

Lone Rock.

Cercospora moUugiiiis n. sp. Showing first small pallid spots

but the infected leaves becoming sere and yellow before the appear¬

ance of the fasciculi which blacken the areas upon which they ap¬

pear; conidiophores amphigenous, fasciculate, fuligenous, suberect

or sometimes curved, undulate or geniculate, simple, usually con-

286 Wisconsin Academy of Sciences , Arts, and Letters.

tinuous, 25-65x3-4/4 ; conidia subhyaline, slender, tapering, straight

or slightly curved, 50-100x3/4.

On leaves of Mollugo verticillata. Lone Kock, July 22, 1921.

Cercospora verhenae-strictae Pk. On Verbena stricta. Lone

Eock. The conidia appear before tissue death has occurred and

therefore before spotting has appeared. There is but a trace of

color in the conidiophores.

Of a specimen on Lepachys pinnata from Pennimore the follow¬

ing notes have been made. Leaves mottled above with indefinite

slightly paler areas; conidiophores amphigenous, solitary or in

small fascicles, brown, straight, curved or undulate, sometimes sep¬

tate, often subnodulose and geniculate distally, 50-100x3-4/4 ;

conidia hyaline, straight, obclavate-cylindrical, becoming septate,

50-100x3 %-5/4. Pending opportunity to examine more material

I have placed this with Cercospora ratibidae Ell. & Barth.

TJstilago sphaerogena Burrill was collected on Echinochloa crus-

galli near MillviUe in 1913 but has not been recorded in the

‘‘ Notes It was collected again on the same species of host at

Madison in 1921 by a class in mycology.

University op Wisconsin Herbarium,

Madison, Wisconsin, April, 1922.

NOTES ON PARASITIC FUNGI IN WISCONSIN— XI

J, J. Davis

The Synchytrium that has been reported as occurring in Wis¬

consin on Lycopus and referred to Synchytrium aureum Schroet.

has been found to develop summer sori and consequently cannot be

a form of that species. As the summer sori develop an empty basal

cell I am now referring it to Synchytrium cellulare Davis. The

galls of the summer sori are simpler than in the type of that

species on Boehmeria cylindrica. The summer sporangia are

globose to polyhedral, 18-24//, in diameter. The resting sori some¬

times develop in prominent multicellular galls. I am now labeling

this var. lycopodis n. var. In addition to the ordinary host, Lyco¬

pus unifloruSj one collection has been made on L. americanus. Ves-

tergren’s Micromycetes rariores selecti 1673 is the form referred

to. It may be that forms on other hosts in Wisconsin that have

been referred to S, aureum will be found to form summer sori when

collections have been made at the proper time.

Fig. 1. Left, two conidipliores of Pldsmopara Tcellermani bearing conidia.

Right, apical portion of a conidiophore with conidia more highly magnified.

Drawn by E. M. Gilbert with the aid of camera lucida.

The record of Iva xanthifolia as a host of Albugo tragopogonis

(D.C.) S. F. Gray in the provisional list was an error, the fungus

on that host being Plasmopara kellermani (Ell. & Hals.) Swingle.

288 Wisconsin Academy of Sciences, Arts, and Letters.

It has been collected at Spooner, Dresser Junction and Alma. This

differs from Basidiophora in the intrafoliar origin of the conidio-

phores and the absence of distinct basidia. From Albugo it differs

in the conidia being borne in a cluster on the apex of the conidio-

phore instead of being catenulate.

Circular black spots 3-4 mm. in diameter sometimes occur on

leaves of Yitis vulpina and Vitis Mcolor in Wisconsin. They ap¬

pear like a young stage of Rhytisma but have been seen only on

living leaves in summer.

The parasite of TJlmus americana and TJ. racemosa recorded in

“Notes” VI, p. 11, under the name Phyllosticta ulmicola Sacc. is

referred to Ph. melaleuca E. & E. in North American Flora 6 : 67.

In the provisional list a parasite of Prunus virginiana was re¬

corded under the name Phyllosticta destruens Desm. regarding

which a note was published in “Notes” I, p. 79. For this I am

now using the binomial Phyllosticta virginiana (Ell. & Hals.)

Seaver (A. A. Flora 6^ : 70). This is not a typical Phyllosticta in¬

asmuch as the sporules are formed by successive basipetal divisions

of filaments the proximal, as yet undivided, portions of which I

take to be the “long sporophores” mentioned in the description of

the similar Ph. innumerahilis Pk. in N. A, Flora 6^ : 52. The micro-

conidia, of some species at least, of Cylindrosporium are produced

in the same way. The form on Amelanchier has been collected in

Wisconsin on hosts referred to A. ohlongifolia and A. spicata.

The Septoria which occurs on Cacalia at riplici folia in Wisconsin

produces white arid spots with a more or less broad dark purple

border like those of S. nahali B. & C. Specimens from Missouri

and Kansas are similar. The single specimen on Cacalia reniformis

has brown spots with a narrow darker border. This developed in a

moist deeply shaded station while those on C. at riplici folia developed

in the open where they were exposed to the direct rays of the sun.

The parasite of Spiraea described by Trelease under the name

Ascochyta salicifoliae, referred to Septoria by Berlese & Voglino

and to Cylindrosporium in “Notes” IV, p. 673 is referred to

Phleospora by Petrak ( Ann. My col. 20 : 210-11 ) . I quite agree

with those who see the genus Phleospora as a mixture of species

referable to other genera and hence one that should be dropped as

was done by Diedicke in Kryptogamenflora der Mark Brandenburg

and by Migula in Thome’s Flora von Deutschland.

Davis — Notes on Parasitic Fungi in Wisconsin — XI. 289

For the species recorded in the provisional list as Ovularia

ohliqua (Cke.) Oud. the name 0. monosporia (West.) Sacc. should

be used because of priority (Sylloge Fungorum 22: 1296).

A collection on Mentha arvensis canadensis made at Arena in

September and referred to Bamularia variata Davis bears conidia

but about IfjL thick.

When well developed the conidia of Cercospora ampelopsidis Pk.

are obclavate 60-80x5-6/^.

The conidial tufts of Cercospora galii Ell. & Hoi. are usually

red until death of the host-tissue takes place. Fasciculi of this

color are more often seen on the small leaved species of host.

There are a number of Compositae that bear Cercospora of a

brown color in both Europe and America. In “Notes” VIII, p.

430 forms on Kudbeckia and Prenanthes were included in Cer¬

cospora tahacina Ell. & Evht. Until more is known of their rela¬

tionship to each other it is perhaps better to keep the forms on the

different host genera distinct. I am therefore labeling the speci¬

mens on Prenanthes Cercospora hrunnea Pk., although it may be

that this is not distinct from C. prenanthis Ell. & Kell, as the

brown color is not always conspicuous. C. rudbeckiae Pk. seems to

be a synonym of 0. tabacina Ell. & Ev. which was published in

1888 not 1886 as given in the Sylloge Fungorum. I append notes

of a specimen on Prenanthes alba collected in Iowa County July 1,

1922. Conidiophores in dense fascicles arising from substomatal,

black, stromatoid tubercles, fuligenous, simple, flexuose, sometimes

geniculate, continuous or sparingly septate, 50x80x4/x; conidia cyl¬

indrical to subclavate, nearly hyaline, straight or curved,

20x60x3-5/x. Macroscopically the angular areas are tobacco brown.

The form on Ambrosia issued in Fungi Columbiani 2117 under the

name Cercospora racemosa E. & M. appears to be a member of

this group which seems closely related to C. ferruginea Fckl. oc¬

curring on Compositae in Europe.

Although Puccinia cryptotaeniae Pk. was recorded in the fourth

supplementary list I find that it was not included in the Provi¬

sional list. It has been collected at Kacine, Ridgeway, and at

the Dells of the Wisconsin River in Adams County.

If one reads the descriptions of Septoria besseyi Pk. on Fraxinus

pennsylvanica lanceolata and Marssonina fraxini (Ell. & Davis)

on Fraxinus nigra they appear to be quite distinct in their micro-

290 Wisconsin Academy of Sciences^ Arts, and Letters.

scopic characters while similar macroscopically. In the former the

sporules are borne in pycnidia, are obtuse, continuous 40-55x about

4:11. In the latter they are borne in acervuli, are acute, 1-septate,

17-33x2-3/a. Examination of specimens, however, shows that the

matter is not as simple as it appears. To illustrate I give some

notes of various specimens.

On Fraxinus pennsylvanica lanceolata.

7-22-21. Pycnidia distally imperfect (hemispherical) ; sporules

subacute to obtuse, becoming 1-septate, 27x50x3ju.

7-21-22. Pycnidia more or less imperfect; sporules continuous,

40— 60x3/x.

7-7-20. Distal portion of pycnidia imperfect ; sporules obtuse or

acute at one end, continuous, 30-50x2%-4/a.

On Fraxinus nigra.

7-29-20. Pycnidia more or less imperfect; sporules acute, con¬

tinuous, 27-33x3jLt.

7- 1-18. Pycnidia distally imperfect'; sporules acute, continuous,

23-32x2jLt.

9-3-15. Pycnidia distally imperfect; sporules acute, continuous,

23— 33x3jLi.

8- 10-15. Spore bodies ranging from prominent acervuli to innate

pycnidia; sporules acute, continuous, 24-36x2-3jLi.

Fungi Columhiani 1526 on Fraxinus pennsylvanica lanceolata

from Kansas, issued as Cylindrosporium fraxini E. &. K., bears

sporules which are obtuse, tapering toward one end, becoming

1-septate 33-45x3jii and apparently belongs in this duplex. The

character of the spore body in such groups as this appears to de¬

pend upon its position in the leaf. The beginning of a spore body

is a spherical mass of hyphae in the leaf tissue. A wall develops

on the periphery of the hyphal mass and under this protection a

sporuligerous or hymenial layer is developed on the inner surface

of the wall which gives rise to the reproductive bodies. Normally

the pycnidium is globular and the entire inner surface of the wall

is lined with hymenium except a small opening for the discharge

of the sporules, the ostiole. When, however, the primary hyphal

mass is in contact with the epidermis or by destruction of interven¬

ing host tissue becomes pressed against it no wall or hymenium is

formed by the portion in contact with the epidermis. This results

in a pycnidium the distal portion of which is defective. If the

Davis — Notes on Parasitic Fungi in Wisconsin — XI. 291

deficiency is considerable the pressure of the mesophyll tends to

flatten the remainder and produce an acervular structure. This

is more likely to happen in the thinner leaf of Fraxinus nigra. In

the normal pycnidium the length of the sporule may equal the

radius of the pycnidial cavity but in the acervular condition it is

limited to the distance from the hymenium to the epidermis.

Fig. 2. Diagram to illustrate influence of position of pycnidium with

reference to epidermis on its development. Drawn by E. Dopp Jr., from a

sketch by the author.

I have seen no specimen that corresponds to the description of

Cylindrosporium fraxini (E. & K.) E. & E. (Journ. Mycol. 1:128),

Cercospora fraxini E. & K. loc. cit. 1:2). As noted above Fungi

Columbiani 1526 issued under this name appears to belong to the

group of forms considered above.

Of the parasite of Fraxinus or eg ana of which Fungi Columbiani

4415, 4719, and 4816, issued as Cylindrosporium fraxini (E. & K.)

E. & E., are examples, I have seen no description. It causes red¬

dish brown spots with a paler margin of more or less circular out¬

line quite variable in size and sometimes confluent. The acervuli

are epiphyllous, subcuticular, and soon erumpent. The sporules

are cylindrical, usually curved or undulate, becoming about 4-sep-

tulate, mostly 40-60x2-3 jn. The epiphyllous, subcuticular habit

especially seems to remove this from the forms considered above.

As the range of species of Uredinales in North American Flora

Vol. 7 is based on specimens in the Arthur herbarium there are

sometimes considerable restrictions. For instance the range of

Puccinia aletridis B. & C. (Dicaeoma (?) aletridis (B. & C.)

Kuntze) is given as ^‘The states bordering the Atlantic Ocean and

Gulf of Mexico from Massachusetts to Texas”. Its occurrence in

northern Indiana was recorded by Burrill {Parasitic Fungi of Illi¬

nois pt. 1, p. 195) and in Wisconsin by Trelease {Preliminary List

of Wisconsin Parasitic Fungi, p. 25). It has been collected in Wis¬

consin as recently as 1922.

292 Wisconsin Academy of Sciences, Arts, and Letters.

Additional Hosts

Syncliytrium on Acalypha virginica has been collected at Lone

Rock, Arena, and Prairie du Sac. Only resting sori were present

which resemble those of 8. cellular e Davis. The sori are first whit¬

ish, then pale yellow, then reddish brown as the wall develops.

The contents are sordid white. Those measured were 80-135/x in

diameter.

Bremia lactucae Regel. On Lactuca scariola integrata. Madi¬

son.

Peronospora parasitica (Pers.) Tul. On Cardamine pennsyl-

vanica. Lone Rock.

Sphaerotheca mors-uvae (Schw.) B. & C. On Bihes lacustre.

White Lake.

Microsphaera alni (Wallr.) Wint. On Quercus Mcolor. Chip¬

pewa Falls.

Erysiphe cichoracearum DC. On Verbena stricta. Blue River.

Phyllachbra graminis panici Shear. On Panicum Scrihnerianum.

Prairie du Sac.

Darluca filum (Biv.) Cast. On TJromyces pyriformis parasitic

on Acorns Calamus. Arena.

The Stagonospora with 7-septate sporules referred to 8. inter-

mixta (Cke.) Sacc. was collected at Blue River on Phyllachora in¬

fected leaves of Cinna latifolia. The pycnidia are pale and thin

walled throughout except the ostiolar ring. The sporules are vari¬

able in length the longest one seen being 90x4/x.

Stagonospora albescens Davis. On Carex vesicaria. Arena. In

this collection about 2 dm. of the apical portion of the leaves is

dead and white with a ferruginous margin at the base. The

sporules are 43-57x7-10^, 6-septate. With pycnidia of a Coniothy-

rium with elliptical to ovate fuscous sporules 7-10x3%-4/li.

Septoria graminum Desm. On Poa pratensis. Madison (Geo.

P. Weber).

Septoria bromi Sacc. var. el3anina n. var. ad interim. On large

indefinite areas that are first yellowish then brown with the death

of the tissues; pycnidia scattered, globose or somewhat flattened.

Davis — Notes on Parasitic Fungi in Wisconsin — XI, 298

dark walled, innate, TO-lOO/x in diameter; sporules straight or

slightly curved, acute, continuous, 30-45x1 %-2/x. On leaves of

Elymus virginicus, Arena, Wisconsin, July 13, 1922.

Septoria glumarum Pass. On Secale cereale. Madison (G. P.

Weber). It is said that this is not distinct from Septoria nodorum

Berk.

Septoria atropurpurea Pk. On Aster lateriflorus. Arena and

Blue Kiver. Of the latter collection it was noted: “On the thin

leaves of this species the spots are more or less angular and with¬

out a colored margin. ^ ’

Entomosporium thuemenii (Cke.) Sacc. On “Paul’s English

Double Hawthorn”. Kacine (Mrs. W. H. Crosby). Reported as

serious in its effects on the host.

Gloeosporium rihis (Lib.) Mont. & Desm. On Bihes lacustre.

White Lake.

Monilia cinerea Bon. On fruit of Pyrus melanocarpa. Arena.

Ramularia uredinis (Voss) Sacc. With Darluca parasitic on

Melampsora bigelowii on Salix nigra. Edgerton.

Ramularia occidentalis Ell. & Kell, has been collected at Chetek

on a leaf of Bumex verticillatus that bears also the species that has

been referred to R. pratensis Sacc.

Ramularia repens Ell. & Evht. On Aralia racemosa. Wyoming.

Ramularia asteris (Phil. & Plowr.) Bubak on Aster azureus.

White Lake.

Tuberculina argUlacea n. sp.

Sporodochia epipyllous on definite orbicular spots which some¬

times become confluent and irregular, especially near the leaf mar¬

gins, numerous, more or less prominent, mostly 50-100/^ in diam¬

eter ; eonidia hyaline, oblong, 8-16x21/2-4/^.

On Caeoma-infected leaves of Rubus allegheniensis or allied

species. Madison and Blue Mounds. On Rubus occidentalis (cult.)

Madison, Wisconsin. May and June. This was abundant at Madi¬

son in 1921, apparently parasitic on the Caeoma. The masses of

eonidia are argillaceous to whitish or, when fresh, honey colored.

294 Wisconsin Academy of Sciences ^ Arts, and Letters.

Cercospora caricina Ell. & Dearn. On Car ex lupulina. Blue

Elver. In this collection the slender conidia range up to 120jLi in

length.

Cercospora nasturtii Pass. On Badicula sylvestris. Edgerton.

In this collection from Lake Koshkonong the spots are mostly

small (about 1 mm.) round, white and arid.

On withered leaves of Badicula palustris. Blue Kiver.

Cercospora zehrina Pass. On Trifolium duhium. Mazomanie.

The publication of this name seems to antedate that of 0. helvola

Sacc.

Cercospora granuliformis Ell. & Hoi. On Viola sagittata. Arena.

Cercospora clavata (Ger.) Pk. On Asclepias Meadii. Arena.

Coleosporium campanulae (Pers.) Lev. Uredinia on Campanula

aparinoides. Blue River. Confined to a single restricted station

where it was abundant and destructive. The uredospores are but

17-22/x long.

Puccinia graminis Pers.

Uredinia on Cinna arundinacea. Saxon. Telia on C, arundina-

cea, Wonewoc. On 0. latifolia, Madison and Cadott.

Puccinia pruni-spinosae Pers. Telia on Prunus nigra. Cadott.

An Aecidium on Linaria canadensis has been known in Wiscon¬

sin for a dozen years or more. Because of the relationship of the

host and the resemblance to the Aecidium on Pentstemon it has

been suspected that it is connected with Puccinia andropogi Schw.

but the connection has not been demonstrated and no record has

been made for this reason. It occurs along the Wisconsin River

from above Mazomanie to Lone Rock and probably further.

Urocystis agropyri (Preuss) Schroet. On AgrosUs alba. Madi¬

son. (W. H. Davis)

Entyloma compositarum Earl. On Boltonia asteroides. Rich¬

land County opposite Muscoda and at Blue River.

Doassansia sagittariae (West.) Pisch. On LopJiotocarpus calcy-

cinus. Blue River. The parasite was not found on Sagittaria in

this locality. In the irregularity of the spore balls this resembles

Davis— Notes on Parasitic Fungi in Wisconsin — XI. 295

the forma confiuens that occurs on Sagittaria heterophylla. Evi¬

dently the smut does not recognize Lophotocarpus as being gener-

ically distinct from Sagittaria.

Sclerotium (f) glohuliferum Davis. On Glyceria grandis.

Chetek.

Additional Species

In 1921 Plasmopara cuhensis (B. & C.) Humphrey was collected

in gardens at Madison on Cucumis sativa^ CucurMta maxima and

Cucurhita pepo in September.

Peronospora myosotidis DBy. A very scanty collection on upper

leaves of Myosotis laxa in July at Arena is supposed to be of this

species but the material was not sufficient for proper examination.

Pleospliaerulina hriosmna Pollacci. On Medicago sativa (cult.)

Madison. (P. R. Jones)

Phyllosticta oakesiae Dearn. & House. On Oakesia sessilifolia.

White Lake. In Notes” VIII, p. 418 a collection of this was re¬

ferred to Phyllosticta cruenta (Fr.) Kickx. As this occurs in Wis¬

consin the development is usually imperfect and I suspect it of

being a state of Diplodia uvulariae Davis.

Phyllosticta ludwigiae Pk. On Ludvigia polycarpa. Blue River.

In this collection the spots are usually elongate and the pycnidia

prominent and often defective.

Phyllosticta verhenicola Martin. On Verbena stricta. Blue

River. I have not seen an authentic specimen of this species and

Dr. Seaver informs me that there is none in the Ellis herbarium.

In this collection the spots are mostly somewhat larger than the

description indicates and somewhat cinereous in color, the sporules

6— 8x2— 3 /X.

Phyllosticta. ambrosiae n. sp.

Spots brown, suborbicular, immarginate, 3-12 mm. in diameter,

mottled with small white arid areas on which the usually solitary

pycnidia are located; pycnidia epiphyllous, dark brown, ostiolate,

globose and about lOO^i in diameter to oval and 150ju. long ; sporules

hyaline, bacilliform, 3-6xlft. On Ambrosia trifida. Days Mills,

Wisconsin, July 30, 1921.

296 Wisconsin Academy of Sciences , Arts^ and Letters.

Sclerotiopsis concava (Desm.) Shear & Dodge. On Frag aria

virginiana. Madison. Steironema ciliatum. Blue River. In the

latter collection with the conidial form Hainesia lythri (Desm.)

Hoehn. Connected with Pezizella lyth^d (Desm.) by Shear &

Dodge. (Mycologia 13: 135 et seq.)

Stagonospora sparganii (Fckl.) Sacc. In leaves of Sparganium

eurycarpum. Spring Green.

In the provisional list Panicum depauperatum was given as

a host of Septoria graminum Desm. In this specimen the pycnidia

are on indefinite pale leaf areas ; they are black, globose to ellipti¬

cal and lenticular, 100-150/x in diameter with firm walls composed

of very small cells ; the sporules are usually curved, sometimes very

strongly so, 50-70xl-l%/x. I am now referring this to the South

American Septoria tandilensis Speg. In his description the author

notes resemblance to Puccinia coronata. The Wisconsin specimen

came to me labeled Puccinia emaculata. The immersed pycnidia

are sometimes laterally compressed.

Septoria cenchrina n. sp.

Spots linear, cinereous, variable in size, often confluent ; pycnidia

amphigenous, brown-black, globose to depressed-globose to elliptical

with a thick wall which is often defective distally, 60-11 5/x in

diameter; sporules filiform, continuous or indistinctly septulate,

straight or more or less strongly curved, hyaline, 30-100xl%-3/>i.

On leaves of Cenchrus carolinianus. Spring Green, Wisconsin,

July 19, 1921. The longer sporules grow out through the widely

open mouth of the pycnidium after the manner of those of Cylin-

drosporium and the species approaches that genus but the Septoria

condition seems to be the normal state.

Septoria didyma Fckl. What is perhaps a short-spored form of

this species is characterized as follows: Spots circular or sub-

circular, brown with a raised purple border, becoming arid, about

1 mm. in diameter ; pycnidia few, pale, globose, epiphyllous, subepi-

dermal, inconspicuous, 50-70/x in diameter; sporules hyaline, ob¬

long, usually curved, uniseptate, 14-27x3/m. On leaves of Salix

longifolia. Lone Rock.

Septoria aparines EU. & Evht. On Galium asprellum. Prairie

du Sac. In the collection that I am recording under this name

the pycnidia are scattered over the lower surface of the leaf

Davis — Notes on Parasitic Fungi in Wisconsin — XI, 297

or sometimes confined to a lateral half. The leaf or half leaf be¬

comes pale yellow and finally dead and brown. The pycnidia are

somewhat flattened, about 100x75/a and extend through to the

epidermis of the upper leaf surface. There are no pycnidia on the

stems. The sporules are continuous, slightly curved, acute,

30-60x1 I have labeled this forma asprelli.

Septoria erechtitis EIL & Bvht. On Erechtities hieracifolia. Blue

Eiver.

Of a collection on Salix lucida the following notes were made:

Spots angular to orbicular, oblivaceous to brown, becoming cinere¬

ous with a dark border, paler below, 1-3 mm. in diameter ; acervuli

mostly hypophyllous, small, scattered, subcuticular; sporules min¬

ute, hyaline, l-Sxl-l^/x abstricted from hyaline, vertical, parallel

hyphae, 15~20xlju,. Cadott, Wisconsin, September 19, 1922. This

seems to be a microconidial state but for the purpose of filing I

have labeled it Gloeosporium egenmn n. sp.

Collet otrickum pisi Pat. On Pisum sativum (cult.) Marshall.

(P. R. Jones)

Oolletotrichum violarum n. sp.

Spots circular to angular or irregular in outline, arid with a

slightly raised margin, l-~6 mm. in diameter, sometimes confluent,

often lacerate ; acervuli amphigenous, small, black ; setae marginal,

black, variable in size (up to 165x5ja) and number, acute; sporules

hyaline, oblong, somewhat curved or at least inequilateral, acute

at each end, continuous, 18-20x3 ju,. On leaves of Viola scahrius-

cula, Spring Green, Wisconsin, July 3, 1922. I have not seen a

specimen of the fungus on Viola rotundifolia collected by G. W.

Clinton and referred to as a variety of Vermicularia eoncentrica

P. & C. (V. pecMi Sacc. nec V, eoncentrica Lev.) by Peck in 29th

Report, p, 48.

Septogloeum subnudum n. sp.

Spots suborbicular, reddish brown with an olivaceous border,

2-5 mm. in diameter, becoming confluent; acervuli few, small,

inconspicuous, epiphyllous, subcuticular; sporules hyaline, fusoid-

oblong, straight, inequilateral or usually curved, 20-40x7-11/1., with

a septum toward each end. On Smilax kerhacea^ Sauk City, Wis¬

consin, August 23, 1922. The development of the parasite seems

298 Wisconsin Academy of Sciences, Arts, and Letters.

to be poor in this collection. One sporule was seen having a third

septum in the middle.

RamiClaria canadensis Ell. & Evht. On Carex sp. indet. Edger-

ton. In this collection the subulate conidiophores range up to 65/x

in length, the conidia are fusiform, acute at each end, 16-23x3i/^-4ju.

In a collection on an undetermined species of Carex made at Madi¬

son in 1912 the conidia are oblong, acute, at least at one end,

20-30x4-6ja. In both collections the conidia develop a median

septum.

Cladosporium caducum n. sp.

Spots hypophyllous, at first pale olivaceous, becoming yellow,

finally brown, indeterminate, suborbicular, 1-4 mm. in diameter,

when old showing on the upper surface ; ponidiophores hypophyl¬

lous, scattered or exceptionally in twos or very small fascicles, deep

brown, ascending to erect, 3-4 septate, sometimes branched, often

paler and torulose apically, 50-80x3 i/^-4/x ; conidia acro-pleuro-

genous, fuligenous or brown, limoniform, apiculate or acute and

narrowly truncate at the extremities, sometimes catenulate and

the chains sometimes branched, continuous, 10-15x4-7jtA. On leaves

of Betula nigra along the Wisconsin river. The spots are usually

multiple, the infected leaves fall and considerable defoliation is

sometimes caused. This seems near Cladosporium but I have

not seen septa in the conidia. They resemble those @f Monilia in

form. Although the parasite is abundant on the bottom lands and

its effects conspicuous I have found it difficult to get good speci¬

mens as the conidiophores are very inconspicuous and soon fall,

away. The best specimens were obtained in July.

Cladotrichum leersiae Atk. On Leersia oryzoides. Blue River.

In this collection oval, pale-avellaneous immarginate spots are pro¬

duced which are sometimes confiuent. The conidia are 13-17/x

in length and sometimes germinate without the formation of a

septum.

Napicladium arundinaceum (Cda.) Saec. On Phragmites com¬

munis. Madison.

Cercospora crassoides n. sp.

Spots orbicular, alutaceous with a broad dark purple border,

less distinct on the lanose lower surface of the leaf ; conidiophores

epiphyllous, sometimes a few on the lower surface, ferruginous, sub-

Davis — Notes on Parasitic Fungi in Wisconsin — XI. 299

erect, more or less geniculate, septate, sometimes branched,

60-70x6-7)a; conidia ferruginous, lanceolate, 6-8 septate, tapering

into a flagelliform distal portion about as long as the body,

100-1 65xl3-7/>i. On leaves of Froelichia floridana, Lone Rock,

Wisconsin, July 23 and 25, 1921. The spots are sometimes very

abundant but mostly sterile. It is not unlikely that vertical septa

appear at full maturity and that this therefore is an Alternaria

like A. crassa (Sacc.) Rands (Cercospora crassa Sacc.) which it

resembles in its conidia.

Cercospora cucu7'l)itae Ell. & Evht. A parasite on leaves of

Cucurhita maxima (cult.) collected at Madison, I have referred to

this species although it is quite different from the type in appear¬

ance. In the Wisconsin material the spots are suborbicular, brown,

with a distinct darker border above, whitish and immarginate

below, 2-5 mm. in diameter, sometimes confluent; conidiophores

scattered or in small tufts, amphigenous, fuscous, somewhat curved,

sometimes septate, simple, more or less denticulate especially near

the apex, 70-130x3-4ju, ; conidia flagelliform, straight or the slender

distal portion curved, septate, hyaline, 87-123x3/x. It was collected

in October.

Uromyces perigynius Hals. var. altiporulus n. var. Uredinia

brown, erumpent, elliptical to linear 0.2-1 mm. long; uredospores

brown, globose to elliptical, ovate, ohovate or oblong, wall brown

1-1%/x thick, echinulate, 12-23xl2-15/>i, germ pores two, variously

situated in the upper half of the spore; telia similar but darker;

teliospores obovate to subfusoid, brown, rounded or often conical

at the apex which is strongly thickened (up to 10/x), pedicel hyaline

as long as the spore or longer. On leaves, bracts and perigynia

of Carex Gi^ayii, Blue River, Wisconsin, August 9, 1922. I am

keeping this collection separate because the germ pores are uni¬

formly further from the equator than in JJ. pe^dgynius as I have

seen it and as it is described. Often one pore lies higher than the

other, sometimes both are subpolar.

Happening to be in a locality in which Aecidium allenii Clint,

was abundant on Shepherdia canadensis attempt was made to get

a clue to the alternate stages. The infected Shepherdia was

confined to the small valley through which the waters of Fish

creek flowed into Green Bay. The first step was to find a host

having the same habitat as the Shepherdia and Carex ehurnea was

300 Wisconsin Academy of Sciences, Arts, and Letters.

hit upon as such a plant. Examination of the old leaves showed that

they had borne a rust the previous year. Then began the watching

of the Carex eburnea plants in the vicinity of infected Shepherdia

plants. Soon a few uredinia appeared on the culms speedily fol¬

lowed by abundant telia on culms and leaves. The following

autumn a trip to the locality was made and abundant telial material

on the Carex secured and seeds of Shepherdia which were kept in a

box outdoors during the winter. It was during this winter, while

attending a meeting of the American Association for the Advance¬

ment of Science that it was learned that Aecidium allenii had been

connected with a grass rust in Colorado. However the following

spring attempts were made to secure plants of Shepherdia cana¬

densis but without success. The seed would not germinate and no

nursery was found that could supply plants. Some plants of

Shepherdia argentea were obtained and attempts made to infect

them but without success. Plants of Artemisia dracunculoides were

also exposed to infection without result. Attempts to infect Shep¬

herdia argentea using teliospores of Puccinia coronata on Calama-

grostis canadensis also failed. The next autumn another trip to

the locality was made and in addition to telial material small plants

of Shepherdia canadensis were secured which were brought to

Madison and heeled in for the winter in what was thought to be a

safe situation. On looking for them in the spring it was found that

trucks had been running over them during the winter but two of

the plants proved to be still alive. These were potted and taken

into the greenhouse and one of them used for the infection experi¬

ment. The weather was very warm and no spore germination was

observed in slide cultures and it was still hotter in the greenhouse.

Nevertheless attempts were made to secure infection but without

result. Finally in despair, the Shepherdia plant was transplanted

to the garden and before leaving it the wisp of wet Carex leaves

that had been suspended over it was drawn across two of the leaves

on both surfaces. As it happened it turned quite cold that night

and typical Aecidium allenii developed on the two leaves that had

been smeared and on those leaves only. Again the locality was

visited the next autumn and telial material and small plants of

Shepherdia canadensis obtained and again the following spring

telial germination was not secured. Still an attempt to infect

in the greenhouse was made which finally resulted in a single but

well developed aecidial spot bearing numerous cups. Examining

this one day it was thought to be ready for plucking and pre-

Davis — Noies on Parasitic Fungi in Wisconsin — XI. 301

serving but as there were a few peripheral cups that were not yet

open it was decided to leave it one more day. The next day it was

found that some creature with epicurean taste had carefully eaten

out each cup except three or four that were not open. The evidence

seems to indicate that there is in Wisconsin a rust bearing aecidia

on Shepherdia canadensis and uredinia and telia on Carex ehurnea.

Puccinia caricis-shepherdiae n. sp.

Aecia; “Spots large, indefinite, yellowish; peridia hypogenous,

elongated, cylindrical, white, nestling among the tomentum of tJie

leaf; spores bright oransre, subglobose, 1-1000 in. in diameter.’’

Aecidium allenii Clint, in 24th Bept. of the New York State

Museum, p. 93. Uredinia few, mostly culmicolous; uredospores

elliptical, deep brown, ;Vall l%x2%/x thick, finely echinulate,

germ pores two approximately equatorial, 23-33xl7-27jw,. Telia

culmicolous and foliicolous, elliptical to linear, rupturing the epi¬

dermis, dark brown ; teliospores brown, clavate, rounded to rounded-

conical at the apex which is strongly thickened (up to 13/x) more or

less constricted at the septum, the lower cell narrower and much

paler than the upper, 44-63xl7-23/>i, pedicel hyaline, the length of

the spore or shorter. This seems closely related to Puccinia prings

heimiana Kleb. but the uredo and teliospores are somewhat larger

and the pedicels of the latter longer and more firmly attached. On

Carex ehurnea, Fish Creek, Wisconsin.

[In 1923 an attempt was made to infect Shepherdia canadensis

with teliospores from Carex ehurnea outdoors to see if the infections

would not be more abundant than had been secured in the green¬

house. Old leaves of Carex ehurnea bearing telia were obtained at

Fish Creek May 14th. Kusted leaves were placed under and on a

plant of Shepherdia canadensis in an open plot from time to time.

Two leaves bearing aecia were removed June 28th. During my

absence after July 3d the plant was watched by Dr. E. A. Baird

who removed two leaves bearing aecia July 23d and one July 31st.

A plant of Shepherdia argentea exposed to infection in the same

manner bore no aecia. No infection of this species occurred in the

greenhouse in previous years. So far as the evidence goes at

present S. argentea does not bear this Aecidium.]

In June, 1922 a few uredinia were found on a leaf of Bumex

altissimus growing beside the railroad at Madison. Later they

were found in more abundance along the same railway line at

302 Wisconsin Academy of Sciences, Arts, and Letters.

Arena and in September telia were f.ound also. The rust agreed

with the description of Puccinia punctiformis Diet. & Hoi. and Dr.

H. S. Jackson has identified it with that species through com¬

parison with authentic material in the Arthur herbarium. As this

had been known only from California, Mexico and Guatemala it

was presumably a waif in Wisconsin and perhaps will not be able

to maintain itself in this climate.

Galinm triflorum as a host of Puccinia punctata Lk. should be

stricken from the provisional list', the rust that has been collected

on that host being Puccinia troglodytes Lindr. Collections have

been made at Neopit, Athelstane and White Lake.

Puccinia tumidipes Pk. On Lycium halimifolium (cult.) Madi¬

son (Edward Kremers, com. R. B. Streets).

University of Wisconsin Herbarium,

Madison, Wisconsin, March, 1923.

Index to ‘‘Notes^’ IX, X, XI.

302a

INDEX TO “NOTES” IX, X, XI

Names of Fungi in italics.

Acalypha virginica, 276, 292

Acer Negundo, 282

Acorus Calamus, 292

Aecidium allenii, 299

Aecidium dicentra e, 279

Aecidium fraxini, 259

Agropyron tenerum, 259

Agrostis alba, 278, 294

Albugo bliti, 265

Albugo Candida, 265

Albugo portulacae, 265

Albugo tragopogonis, 265

Allium canadense, 281

Alnus crispa, 257

Amaranthus hybridus, 265

Amaranthus retroflexus, 265

Ambrosia, 289

Ambrosia psilostachya, 259, 276

Ambrosia trifida, 254, 295

Amelanchier oblongifolia, 288

Amelanchier spicata, 288

Anemone canadensis, 271

Anemone quinquefolia, 266, 271

Anemone virginiana, 259, 283

Apocynum androsaemifolium, 277

Aralia racemosa, 293

Artemisia ludoviciana, 259

Artemisia serrata, 256

Asclepias Meadii, 294

Ascochyta lophanthi lycopina, 277

Ascochyta necans, 274

Ascochyta pisi, 277

Ascochyta pteridis, 273, 274

Ascochyta salici/oliae, 288

Aster azureus, 293

Aster lateriflorus, 257, 276, 277, 280,

293

Aster paniculatus, 259, 277

Aster sagittifolius, 283

Aster Tradescanti, 256, 257, 277

Aster oma gentianae, 273

Aster omella astericola, 281

Baptisia bracteata, 258

Basidiophora entospora, 256, 269, 276

Betula nigra, 276, 298

Bidens vulgata puberula, 278

Bifusella linear e, 252

Boehmeria cylindrica, 287

Boltonia asteroides, 294

Botrytis epichloes, 261

Bremia lactucae, 256, 266, 292

Cacalia atriplicifolia, 288

Cacalia reniformis, 288

Campanula aparinoides, 294

Cardamine pennsylvanica, 292

Carex, 298

Carex chordorrhiza, 277

Carex eburnea, 257, 301

Carex Grayii, 299

Carex grisea, 253

Carex intumescens, 259

Carex longirostris, 259

Carex lupulina, 294

Carex tribuloides, 281

Carex vesicaria, 292

Carya cordiformis, 253, 261, 283

Carya ovata, 253

Celtis occidentalis, 285

Cencbrus carolinianus, 296

Gercospora ampelopsidis, 289

Gercospora antipus, 278

Gercospora arctostaphyli, 253

Gercospora brunnea, 289

Gercospora caricina, 253, 258, 294

Gercospora clavata, 294

Gercospora crassoides, 298

Gercospora cucurbitae, 299

Gercospora davi&ii, 275

Gercospora diffusa, 278

Gercospora epigaeae 275

Gercospora epigaeina, 275

Gercospora euonymi, 262

Gercospora flagellifera, 258

Gercospora fraxini, 291

Gercospora gain, 289

Gercospora granuliformis, 294

Gercospora helvola, 294

Gercospora medicaginis, 278

Gercospora mississippiensis, 274

Gercospora molluginis, 285

Gercospora moricola, 261

Gercospora nasturtii, 294

Gercospora oxybaphi, 258

Gercospora platyspora, 275

Gercospora prenanthis, 289

Gercospora racemosa, 289

Gercospora ratibidae, 286

Gercospora rudbeckiae, 289

Gercospora saniculae, 275

Gercospora six, 275

Gercospora smilacina, 275

Gercospora smilacis, 274-275

Gercospora stomatica, 258, 275

Gercospora tabacina, 289

Gercospora teucrii, 262

Gercospora velutina, 258

Gercospora verbenae-strictae, 286

Gercospora viciae, 258

Gercospora zebrina, 294

Gercosporella apocyni, 253

302b Wisconsin Academy of Sciences, Arts and Letters,

Cercosporella cana, 257, 274

Cercosporella celtidis, 285

Cercosporella mirahilis, 274

Cercosporella pyrina, 257

Cercosporella reticulata, 261

Cerotelium urticastri, 279

Chenopodium album, 252, 267

Cicuta bulbifera, 277

Cicuta maculata, 252

Cinna arundinacae, 294

Cinna latifolia, 292, 294

Cirsium discolor, 256

Cirsium muticum, 266

Cladochytrium maculare, 265

Cladosporium astericola, 285

Cladosporium caducum, 298

Cladosporium carpophUum, 261

Cladosporium simplex, 285

Cladosporium suhsessile, 253, 257

Cladosporium triostei, 257

Cladotrichum leersiae, 298

Claviceps nigricans, 280 "

Clintonia borealis, 263

Coleosporium campanulas, 294

Colletotrichum graminicolum, 257

Colletotrichum pisi, 297

Colletotrichum violarum, 297

Convolvulus spithamaeus, 261

Cordyceps clavulata, 260

Crataegus Oxyacantha, 293

Cryptomyces pteridis, 273

Cucumis sativa, 260, 295

Cucurbita maxima, 295, 299

Cucurbita Pepo, 295

Cylindrosporium apocyni, 253

Cylindrosporium caryogenum, 283

Cylindrosporium clematidis, 253

Cylindrosporium fraxini, 290, 291

Cylindrosporium guttatum, 283

Cylindrosporium scdicifoliae, 288

Cylindrosporium toxicodendri, 283

Cylindrosporium vermiforme, 257

Cyperus Schweinitzii, 258

Dactylis glomerata, 278

Darluca filum, 277, 292

Depazea gentianaecola, 272, 273

Desmodium canadense, 259, 274

Desmodium illinoense, 274

Didymaria platyospora, 275

Didymellina iridis, 253

Diplodia uvulariae, 295

Doassansia ranunculina, 275

Doassansia sagittariae, 294

Echinochloa crusgalli, 286

Eleocharis palustris, 280

Elymus bracbystachys, 276

Elymus virginicus, 293

Entomosporium thuemenii, 293

Entyloma calendulae, 254

Entyloma compositarum. 254, 259, 294

Entyloma linariae gratiolae, 262

Entyloma ranunculi, 278

Eocronartium muscicola, 260

Epichloe typhina, 261

Erechtites hieracifolia, 297

Erigeron annuus, 257, 274

Erysiphe cichoracearum, 276, 292

Eupatorium urticaefolium, 278

Evonymus atropurpureus, 260, 262

Fragaria virginiana, 278, 296

Fraxinus americana, 278

Fraxinus nigra, 257, 259, 260, 289, 290

Fraxinus oregana, 291

Fraxinus pennsylvanica, 257, 260, 279.

285, 289, 290

Froelichia floridana, 299

Fusicladium caryigenum, 261

Fusicladium cerasi, 261

Fusicladium depressum, 275

Fusicladium effusum, 261

Fusidium pteridis, 273

Galium asprellum, 276, 296

Galium triflorum, 302

Gaylussacia baccata, 278, 279

Gentiana Andrewsii, 272

Gentiana puberula, 273

Geranium maculatum, 266

Geum canadense, 256

Geum strictum, 263

Geum virginianum, 263

Gloeosporium aridum, 257

Gloeosporium caryae, 253

Gloeosporium egenum, 297

Gloeosporium fraxineum, 257

Gloeosporium leptospermum, 273

Gloeosporium necans, 273

Gloeosporium ohtegens, 273

Gloeosporium pteridis, 273

Gloeosporium ribis, 293

Gloeosporium salicis, 256—257

Glyceria grandis, 295

Glyceria nervata, 261

Gnomonia ulmea, 256

Gratiola virginiana, 262

Hainesia lythri, 296

Halenia deflexa, 264

Helenium autumnale, 276

Hepatica acutiloba, 271

Heterosporium gracUe, 253

Hydrocotyle americana, 255, 264

Hypoderma lineare, 252

Hypoxis hirsuta, 283

Ilex verticellata, 260

Iva xanthifolia, 287

Koeleria cristata, 254, 281

Lactuca Scariola integrata, 256, 292

Lactuca spicata, 266

Laportea canadensis, 279

Lathyrus palustris, 258

Lecanium corni, 260

Index to Notes’’

IX, X, XL

302

Leersia oryzoides, 298

Lepachys pinnata, 286

Leptothyrium conspicuum, 272

Leptotkyrium gentianaecolum, 272

Lespedeza frutescens, 258

Linaria canadensis, 294

Lonicera Sullivantii, 278

Lophodermium linear e, 252

Lophotocarpus calycinus, 294

Ludvigia polycarpa, 295

Lycium halimifolium, 302

Lycopus americanus, 287

Lycopus uniflorus, 264, 282, 287

Lycopus virginicus, 277

Lysimachia terrestris, 257, 264

Macrophoma arens, 281

Marssonina fraxini, 253, 289

Marssonina necans, 273

Marssonina populi, 277

Marssonina potentiUae, 253, 278

Marssonina potentUlae tormentUlae, 257

Marssonina rhahdospora, 257

Marssonina thomasiana, 260

Medicago lupulina, 278

Medicago sativa, 295

Melampsora americana, 255

Melampsora arctica, 255

Melampsora bigelowii, 293

Melica striata, 256, 257

Melilotus alba, 275, 281

Mentha arvensis canadensis, 289

Microsphaera alni, 292

Mollisia earliana, 253

Mollugo verticillata, 286.

Monilia cinerea, 293

Morus rubra, 261

Myeosphaerella ruhi, 272

Myosotis laxa, 295

Napicladium arundinaceum, 298

Oakesia sessilifolia, 295

Oenothera biennis, 268

Oenothera rhombipetala, 277

Oryzopsis asperifolia, 251

Ovularia monosporia, 289

Ovularia ohliqua, 289

Oxybaphus hirsutus, 258

Panicum depauperatum, 296

Panicum Scribnerianum, 292

Panicum virgatum, 276

Parietaria pennsylvanica, 280

Pedicularis canadensis, 264

Peronospora aXsinearum, 267

Peronospora alia, 268

Peronospora arthuri, 268

Peronospora calotheca, 268, 276

Peronospora chamaesycis, 268

Peronospora corydalis, 267

Peronospora effusa, 267

Peronospora ficariae, 267, 276

Peronospora floerkeae, 268

Peronospora leptosperma, 256, 268

Peronospora myosotidis, 295

Peronospora ohovata, 267

Peronospora parasitica, 268, 292

Peronospora polygoni, 267

Peronospora potentUlae, 268

Peronospora rubi, 256, 268

Peronospora schleideni, 267

Peronospora silenes, 267

Peronospora trifoliorum, 268

Peronospora urticae, 267

Peronospora viciae, 268

Petasites palmatus, 264

Pezizella lythri, 296

Phacidium taxi, 280

Phleospora anemones, 283

Phleospora oxyacanthae, 274

Phleospora sdlicifoliae, 288

Phleospora ulmi, 256, 277

Phoma alliicola, 281

Phragmites communis, 298

Phyllachora ambrosiae, 276

Phyllachora graminis, 276

Phyllachora graminis panici, 276, 292

Phyllachora melicae, 256

Phyllachora oryzopsidis, 251

PhyUactinia corylea, 276

PhyUosticta ambrosiae, 295

Phyllosticta apocyni, 277

PhyUosticta atriplicis, 252

Phyllosticta congesta, 280

Phyllosticta cruenta, 295

Phyllosticta destruens, 288

Phyllosticta dioscoreae, 260

Phyllosticta fraxinicola, 260

PhyUosticta grossulariae, 256

Phyllosticta innumerabilis, 288

PhyUosticta ludwigiae, 295

PhyUosticta melaleuca, 288

PhyUosticta oakesiae, 295

Phyllosticta phaseolina, 260

Phyllosticta phomiformis, 256

Phyllosticta pyrolae, 281

PhyUosticta quercea, 284

Phyllosticta steironematis, 281

PhyUosticta ulmicola, 288

PhyUosticta verbasdcola, 281

Phyllosticta verbenicola, 295

Phyllosticta virginiana, 288

Physalis heterophylla, 278

Physalospora ambrosiae, 276

Physocarpus opulifolius, 251

Physoderma vagans, 265

Phytophthora infestans, 266

Phytophthora thalictri, 266

Phythophthora thalictri, 266

Piggotia vaccinii, 272

Pisum sativum, 297

Plasmopara acalyphae, 251, 266

Plasmopara australis, 267

302d Wisconsin Academy of Sciences, Arts and Letters,

Plasmopara cephalophora, 267

Plasmopara cuhensis, 260, 295

Plasmopara geranii, 266

Plasmopara halstedii, 267

Plasmopara humuli, 266

Plasmopara Ulinoensis, 280

Plasmopara Tcellermani, 287

Plasmopara ohducens, 251, 266

Plasmopara pygmaea, 266, 271

Plasmopara ribicola, 266

Plasmopara viburni, 267

Plasmopara viticola, 267

Pleosphaerulina briosiana, 295

Poa annua, 277, 279

Poa pratensis, 278, 292

Polygonum Persicaria, 279

Populus balsamifera, 277

Populus grandidentata, 257

Populus tremuloides, 253, 257

Potentilla anserina, 278

Prenanthes, 289

Prenanthes alba, 264, 289

Protomyces andinus, 269

Prunus americana, 261

Prunus nigra, 294

Prunus pennsylvanica, 280

Prunus virginiana, 288

Psedera, 252

Pseudopeziza singularia, 276

Puccinia absinthii, 259

Puccinia agropyri, 259

Puccinia aletridis, 291

Puccinia andropogonis, 254

Puccinia anemones-virginianae, 259

Puccinia asteris, 277

Puccinia atropuncta, 275

Puccina caricis-shepherdiae, 301

Puccinia cryptotaeniae, 289

Puccinia gigantispora, 259

Puccinia graminis, 259, 279, 294

Puccinia Jcoeleriae, 254

Puccinia Tcoeleriae-liatridis, 254

Puccinia liatridis, 254

Puccinia magnusiana, 254

Puccinia peridermiospora, 259, 279

Puccinia polygoni-amphibii, 279

Puccinia pruni-spinosae, 294

Puccinia punctiformis, 302

Puccinia pustulata, 254

Puccinia simillima, 254

Puccinia troglodytes, 302

Puccinia tumidipes, 302

Puccinia zygadeni, 275

Pucciniastrum arcticum americanum, 254,

255

Pucciniastrum myrtilli, 279

Pyrola elliptica, 281

Pyrus ioensis, 257

Pyrus melanocarpa, 293

Quercus bicolor, 256, 284, 292

Radicula palustris, 294

Radicula sylvestris, 294

Ramularia asteris, 257, 293

Ramularia canadensis, 298

Ramularia celtidis, 285

Ramularia desmodii, 274

Ramularia dispar, 278

Ramularia effusa, 278

Ramularia fraxinea, 278

Ramularia lysimackiae, 257

Ramularia occidentalis, 293

Ramularia pratensis, 293

Ramularia repens, 289, 293

Ramularia tanaceti, 285

Ramularia tenuis, 261

Ramularia uredinis, 278, 293

Ramularia variata, 289

Ramularia virgaureae, 257, 258, 259, 278

Ranunculus recurvatus, 276, 279, 280

Ranunculus septentrionalis, 276, 277, 279,

280

Rhus Toxicodendron, 283

Rhytisma linear e, 252

Ribes lacustre, 292, 293

Ribes oxyacanthoides, 256

Ribes prostratum, 266

Ribes triste, 266

Rubus allegheniensis, 251, 256, 282, 293

Rubus canadensis, 257

Rubus hispidus, 257, 264

Rubus idaeus aculeatissimus, 254

Rubus occidentalis, 254, 255, 293

Rubus triflorus, 254, 255, 263

Rubus villosus, 264

Rudbeckia, 289

Rudbeckia laciniata, 264

Rumex altissimus, 301

Rumex verticillatus, 293

Salix alba vitellina, 256

Salix amygdaloides, 278

Salix longifolia, 296

Salix lucida, 257, 277, 297

Salix nigra, 293

Schizonella melanogramma, 259

Scirpus atrovirens, 263, 272

Sclerospora graminicola, 269

Sclerotiopsis concava, 296

Sclerotium deciduum, 259

Seder otium globuliferum, 295

Scolecotrichum graminis, 278

Scrophularia leporella, 256

Secale cereale, 293

Septocylindrium caricinum, 253

Septocylindrium concomitans, 278

Septogloeum ampelopsidis, 252

Septogloeum convolvuli, 261

Septogloeum querceum, 283

Septogloeum subnudum, 297

Septoria acerella, 282

Septoria ampelopsidis, 252

Index to *^Notes’’ IX, X, XI.

302e

Septoria angularis, 283

Septoria annua, 277

Septoria aparines, 296

Septoria aquilegiae, 252

Septoria atriplicis, 252

Septoria atropurpurea, 277, 293

Septoria hesseyi, 289

Septoria bromi elymina, 292

Septoria cacaliae, 288

Septoria caricinella, 277

Septoria cenchrina, 296

Septoria chenopodii, 252

Septoria comitata, 282

Septoria commonsii, 256

Septoria didyma, 296

Septoria erechtitis, 297

Septoria fumosa, 283

Septoria gei, 256

Septoria glumarum, 293

Septoria graminum, 292, 296

Septoria irregularis, 283

Septoria lycopi, 282

Septoria negundinis, 282

Septoria nocti florae, 256

Septoria nodorum, 293

Septoria oenotherae, 277

Septoria polaris, 277

Septoria populi, 257

Septoria rubi, 272, 282

Septoria salicifoliae, 288

Septoria scrophulariae, 256

Septoria sii, 277

Septoria solidaginicola, 277, 283

Septoria stachydis, 256

Septoria tandilensis, 296

Septoria umbelliferarum, 253

Shepherdia argentea, 301

Shepherdia canadensis, 299, 301

Silene nivea, 256

Smilax herbacea, 297

Solidago latifolia, 261, 283

Solidago nemoralis, 257

Solidago patula, 277

Solidago serotina, 258, 278, 285

Sparganium eurycarpum, 296

Sphaeria gentianaecola, 273

Sphaerotheca humuli, 251

Sphaerotheca mors-uvae, 292

Spiraea salicifolia, 288

Stachys palustris, 256

Stagonospora albescens, 281, 292

Stagonospora atriplicis, 252

Stagonospora intermixta, 292

Stagonospora meliloti, 282

Stagonospora sparganii, 296

Steironema ciliatum, 281, 296

Stipa spartea, 262

Strophostyles helvola, 260, 279

Synchytrium anemones, 264

Synchytrium asari, 264

Synchytrium aureum, 255, 263, 276

Synchytrium cellulare, 262, 287

Synchytrium cinnamomeum, 280

Synchytrium decipiens, 264

Synchytrium globosum, 263

Synchytrium nigrescens, 280

Synchytrium pulvereum, 264

Synchytrium scirpi, 263, 272

Taenidia integerrima, 275

Tanacetum vulgare, 285

Taxus canadensis, 280

Teucrium canadense, 262

Thalictrum dasycarpuip, 266, 278

Thalictrum revolutum, 266

Trifolium dubium, 294

Triosteum aurantiacum, 257

Tuberculina argillacea, 293

Typhula muscicola, 260

Ulmus americana, 288

Ulmus fulva, 256, 277

Ulmus racemosa, 256, 288

TJrocystis agropyri, 278, 294

JJromyces appendiculatus, 279

Uromyces graminicola, 254

Uromyces hedysari-paniculati, 259

Uromyces minutus, 259

Uromyces perigynius altiporulus, 299

Uromyces pyriformis, 292

Urophlyctis major, 265

Urophlyctis pluriannulata, 265

Ustilago hypodytes, 262

UstUago sphaerogena, 286

Ustilago striaeformis, 2.1 S

Venturia cerasi, 261

Verbascum Thapsus, 281

Verbena stricta, 286, 292, 295

Vernonia fasciculata, 276

Vicia americana, 268

Vicia angustifolia segetalis, 277

Viola conspersa, 264

Viola pallens, 264

Viola pubescens, 264

Viola sagittata, 294

Viola Bcabriuscula, 297

Vitis bicolor, 288

Vitis vulpina, 288

Zizia aurea, 265

THE CYTOLOGY AND PHYSIOLOGY OF VENTURIA

INEQUALIS (COOKE) WINTER

Charles N. Fret

Introduction

During the last century our present conceptions of the nature

and habits of the fungi have been developed. Fries 1819-22 began

the systematic classification of the fungi. De Bary (1853) and

Kiihne (1856) demonstrated the parasitic nature of the fungi and

laid the foundation of modern pathology. Pasteur (1858-60),

Raulin (1869-70) and Nageli (1882) studied the nutrition of fungi.

In 1791 Bulliard described the asci as female organs. He believed

that the asci were fertilized by the bursting of the paraphyses.

It was de Bary (1863), however, who first studied and described

the sexual organs of the fungi and observed fertilization. His

critical observations form the basis of our knowledge as developed

at the present time. De Bary (1863-70) discussed the origin of

the fungi, the function of the archicarp and antheridium, which he

considered as functional sex organs, and designated the ascus

a spore mother cell. He believed that the archicarp should be re¬

garded as the female sex organ and the antheridium as the male

and that there occurred a material union of a male cell, or a part

of its protoplasm and nuclear content, with a cell of the archicarp.

Prom the archicarp the ascogenous hyphae originated, the latter

developing the ascus which he designated a spore mother cell.

According to de Bary it is possible that the lines of descent for the

Ascomycetes came from divergent forms. The two groups, the

Phycomycetes and Ascomycetes, converge, the Mucorineae and Pero-

nosporeae of the former being comparable to Eremascus among the

Ascomycetes. Due to the insufficient knowledge of his time de

Bary concluded the relationship and phylogeny of the higher fungi

would remain in doubt until further research establishing more

definite evidence as to their origin and character. He was struck

by the similarity between the sex organs of the Plorideae and the

304 Wisconsin Academy of Sciences ^ Arts, and Letters.

Ascomycetes and in his lectures declared that a relationship might

exist between the two groups.

Tulasne (1867) described the fusion of the sex organs of Pyro-

nema conflnens. Janczenski (1871) discovered the origin of the

ascogenous hyphae in Ascoholus furfuraceus and established their

relation to the asci, thereby verifying the predictions of de Bary.

In 1883 Kuhlman substantiated the statements of Janzewski as to

the origin of the asci. Sach (1875) developed views similar to de

Bary’s. He noted the similarity of the archicarp of the fungi

to the procarp of the red algae, and believed the fungi were derived

from the red algae.

Opposed to the views of de Bary were those of Brefeld (1879) and

Van Tieghem (1891). Brefeld maintained that the fungi pos¬

sessed no sexual characters. He attributed only a vegetative

character to the fusions. The ascus was considered comparable to

a sporangium of the lower fungi, the spores being produced endo¬

genously in the ascus as in the sporangium. The Ascomycetes

were derived from the Phycomycetes through such forms as

Thamnidium and Mortierella. His theory was based on the assump¬

tion that the sporangium through evolution had become an ascus.

As we shall see later, this theory rests on poor evidence. Due to

the work of Harper and others, it was clearly shown that the two

organs are not homologous. In the Oomycetes the spores are pro¬

duced by progressive cleavage, whereas in Ascomycetes the spores

are delimited by astral radiations from the centrosome-like body

of the ascus nuclei.

Van Teighem (1891) expressed the view that the archicarp is

perhaps only an ascogenous hypha differentiated at an early period,

and that the antheridia are part of the enveloping perithecium or

serve as respiratory organs.

In 1894 Dangeard, while investigating Peziza vesiculosa, dis¬

covered fusion of nuclei in the ascus. He maintained that the sex

organs were no longer functional and that the ascus may be con¬

sidered an egg in which fertilization occurred. The archicarp and

antheridium might be regarded as vestigial organs. Later he

changed his views somewhat, stating that the ascogonium and

ascogenous hyphae constitute a gametophore derived from a

gametangium which formerly functioned in the production of

motile gametes, but lost this function in adapting itself to the land

habit. This conception involves the relationship of the Aseomy-

cetes to the Oomycetes. He assumes that the ascogonium and

Frey— Physiology of Venturia Inequalis.

305

ascogenous hyphae are outgrowths from a parthenogenetic egg and

the ascus is the result of this vegetative growth, an organ which has

no equivalent in the lower fungi.

Harper (1895) began his studies of the Ascomycetes with Peziza

stevensonia and Ascoholus furfurmeous and later investigated

Sphareotheca castagnei. He found that the asci arise from ascoge¬

nous hyphae substantially in the manner described by de Bary

(1865-66), Janzewski (1871), and Kuhlman (1883), and described

the fusion of two nuclei in the ascus of Peziza stevensonia to form

the primary ascus nucleus. The fusion nucleus grows and either

migrates to the periphery or remains in the center of the ascus. He

followed the processes involved by the three successive divisions of

the fusion nucleus resulting in the formation of eight nuclei.

Harper found the chromosome number to be eight and believed

that in the first division, which differs in appearance from the

other two, a reduction in the chromosome number occurs. Fur¬

ther research indicated that the process of spore formation in

Ascoholus furfuraceus was similar.

De Bary had decsribed the formation of the antheridium and the

ascogonium from the mycelium of Sphaerotheca castagnei. Har¬

per (1896) found that the antheridium and ascogonium are pro¬

duced on neighboring hyphae. Each sex organ has one nucleus at

the beginning of development and each cell is cut off from the

mycelium by a wall. When the ascogonium has completed its de¬

velopment the antheridial hypha elongates, its nucleus divides, one

daughter nucleus passes to the tip of the cell and a partition wall

is laid down between the two nuclei. The antheridium lies against

the oogonium, the wall between the two organs is dissolved at the

point of contact, and the male nucleus pushes through the pore

and fuses with the egg nucleus. Sterile hyphae grow from the

female stalk and surround the sex organs after the completion of

nuclear fusion. Similar hyphae grow from the stalk cell of the

antheridium; the antheridium itself disintegrates after fertiliza¬

tion.

The nucleus of the ascogone after fusion with the male nucleus

divides and a wall is laid down which separates the two daughter

nuclei. The upper cell nucleus divides again and a second wall is

formed. Five or six uninucleate cells may be formed but the penul¬

timate cell is binucleate. The two nuclei of the penultimate cell

fuse, forming the primary ascus nucleus. Harper holds that this

fusion is vegetative, the sexual fusion occurring in the ascogonium.

306 Wisconsin Academy of Sciences, Arts, and Letters.

Three successive nuclear divisions of the primary ascus nucleus

result in the formation of the eight nuclei of the spores.

The sex organs of Erysiphe communis, according to a later paper

by Harper (1896), are borne on different hyphae, and the uninu¬

cleate oogonium is separated from the mycelium by a basal cell.

The uninucleate antheridium is similar, but smaller and also has a

basal cell. The nucleus of the antheridium divides and a wall is

laid down between the daughter nuclei. The nucleus of the end

cell of the antheridium passes into the oogonium through a pore

formed by the dissolution of the cell wall at the point of contact

between the two sex organs. The sterile hyphae arise from the

stalk cell of the oogonium. The nucleus of the oogonium, formed

by the fusion of the egg and the male nucleus, then divides. The

daughter nuclei immediately divide again. The oogonium elon¬

gates and walls are formed between the nuclei, giving rise to a

chain of five to seven cells. The chain of cells so formed is crooked

and the penultimate cell has two nuclei and from this cell the

ascogenous hyphae arise. The ascogenous hyphae branch, each

branch forming two or three cells. One large intercalary cell in

each branch has two nuclei; the two nuclei of this cell fuse and

then the cell develops into an ascus. The remaining cells of the

ascogenous hyphae degenerate.

The cells of the ascogonium of Ascoholus furfuraceus according

to Harper (1896) are uninucleate in the early stages, but later

seem to be connected by openings in the walls between them. The

ascogonium becomes a bow-like row of cells. The nuclei in each

cell divide many times. The fourth cell from the apical end of the

ascogonium is the largest and from it ascogenous hyphae arise.

The nuclei of the adjacent cells flow into the ascogenous hyphae

and the empty ascogonium disintegrates. The nuclei gather at the

tip of the ascogenous hyphae. The ascogenous hyphae branch, and

nuclei pass into the branches which are then separated from the

ascogenous hyphae by walls, thereby making the cells so cut off

ascus mother cells from which asci arise.

In 1897 Harper described in detail the cytological phenomena

connected with ascus development and spore formation in Erysiphe

communis, Peziza stevensoniana and Ascoholus furfuraceus. Fusion

of two nuclei to form the primary ascus nucleus takes place. The

process is eventually the same in the three organisms. Harper

does not regard the fusion in the ascus as sexual, for in Ascoholus

furfuraceus cases occur of four nuclei fusing to form the ascus

Frey — Physiology of Venturia Inequalis.

307

nucleus. He believes that this fusion, since it involves more than

two nuclei, is not sexual. The ascus is a new structure, the prod¬

uct of the asexual generation. The spores are cut out of the cyto¬

plasm by kinoplasmic radiations from the central body. Harper

found that reduction occurs in the first division of the ascus nucleus

of Erysiphe communis.

The process of spore formation is similar in Peziza stevensonia

and also in Ascoholus furfuraceus. The kinoplasm forms the spin¬

dle fibres and the astral radiations which cause the delimitation of

the spores. The nucleus may be active in directing the process of

spore delimitation as it moves to the periphery of the ascus. The

central body may be only a distributing point of nuclear activity

although it appears to be able to form new kinoplasmic fibres by

its own activity. The nature of the kinoplasmic changes in form¬

ing a cell wall are not known. The chromosome number does not

change but there seems to be a reduction in quantity. The chromo¬

somes always remain attached to the central body by fibres.

Harper (1899) studied spore formation in the sporangia of cer¬

tain Phycomycetes. Brefeld (1873) held that the jelly of the

Mucorineae, in which asexual spores are imbedded, is homologous

with the epiplasm of the ascus, and that the sporangia of the

Zygomycetes illustrate an ancestral type from which the ascus has

developed. Harper ’s results utterly failed to corroborate Brefeld ’s

theory. In Synchytrium, Pilobolus and Sporodinia the spores are

cut out by progressive cleavage of the cytoplasm, as a result of

which all the cytoplasm of the sporangia is used in the formation

of the spores. There is, therefore, no structure in the molds com¬

parable with the epiplasm of the ascus. Spore formation in the

ascus of Lachnea scutellata was found to be similar to the corre¬

sponding process in Peziza stevensoniona and Ascoholus furfuraceus

except in slight details. The process in Sporodinia is spore forma¬

tion by progressive cleavage; the process in Lachnea scutellata is

free cell formation.

Tulasne (1866) had observed a fusion of oogone and antheridium

by means of pores in Pyronema confiuens. De Bary (1863) could

not find the actual fusion,’ though he felt certain that it' occurred.

Harper (1900) figures an oogonium of Pyronema confiuens with a

trichogyne applied to the antheridium. At the point of application

of the trichogyne to the wall of the antheridium a pore is formed

by dissolution of the cell walls. The nuclei of the antheridium pass

into the trichogyne and through a pore in the wall separating the

308 Wisconsin Academy of Sciences, Arts, and Letters.

trichogyne from the oogonium. The opening in the basal cell wall

of the trichogyne then closes. The male and female nuclei in the

oogonium then fuse in pairs. As there are more egg nuclei than

male nuclei many of the former fail to find mates and are later

disorganized. The female nuclei aggregate in the center of the

oogonium previous to fusion, but after fusion migrate to the

ascogenous hyphae which are produced from bud-like outgrowths

from the oogonium at the time of the formation of a pore in the

basal wall of the trichogyne. The ascogenous hyphae grow and

branch profusely, their ultimate branches giving rise to the asci.

At first each branch contains two nuclei. These nuclei are not sis¬

ter nuclei but come from different nuclei in the ascogenous hyphae,

each nuclei having descended from one of the original pair in the

ascogenous hyphae. Possibly each one is a direct descendant of

one of the daughter nuclei of the fusion nucleus and the two nuclei

in the ascus therefore represent two distinct lines of descent.

The two nuclei in the branch, which is curved, forming a shep¬

herd’s crook, divide simultaneously and form four nuclei. The

spindles are so placed that a pair of nuclei are left in the crook,

one from each spindle, so they are not sister nuclei. The other two

nuclei, each the daughter of a different mother nucleus, migrate

from the spindles, one going to the apical region of the branch,

the other to the basal region. Two partition walls are then laid

down making three cells out of the branch, the apical cell contain¬

ing the single apical nucleus, the middle cell containing two nuclei,

and the basal cell containing the nucleus that migrated to the basal

region. The middle cell lies in the crook of the branch and is some¬

what dome-shaped. It now forms the ascus, the two nuclei in it

fuse to form the primary ascus nucleus. The nuclear fusion is

followed by three successive divisions that are similar to those al¬

ready described by Harper for other Ascomycetes.

The results of Harper’s (1905) study of Phyllactinia corylea

substantiated his earlier work. The sexual apparatus is formed

where two hyphae cross or lie close to each other. The oogonium

and antheridium become applied and spirally twist about each

other. A portion of the walls between them is dissolved and the

single nucleus of the antheridium migrates through the pore into

the oogonium. It fuses with the somewhat larger nucleus of the

oogonium and the pore through which it entered closes. After

nuclear fusion, or just previous to the migration of the male

nucleus, the stalk cell of the oogonium produces paraphyses to form

Frey — Physiology of Venturia Inequalis.

309

the perithecium. The stalk cell of the antheridium may also form

paraphyses. The ascogonium increases in size, the fusion nucleus

divides and a chain of three to five cells is formed. The end cell

of the chain is uninucleate, the penultimate cell is generally bi-

nucleate but may contain more than two nuclei. Branches arise

from the penultimate cell and perhaps from some of the others;

these branches are the ascogenous hyphae. The asci are formed

after the ascogeneous hyphae have become septate. Each ascus

arises from a terminal cell or as a lateral outgrowth from an inter¬

calary cell of an ascogenous hypha. The cells that produce asci

are at first binucleate. The ascogenous hyphae are multinucleate

before they become septate which makes it possible to conceive

that the two nuclei in the ascus are not sister nuclei. Following

nuclear fusion in the ascus the ascogonium and the sterile cells of

the ascogenous hyphae degenerate.

About the time of degeneration of the oogonium the perithecial

envelope has made considerable growth. The wall formed by the

perithecial hyphae consists of three layers: a peripheral, or outer

layer of cells; a protective lignified layer, beneath the peripheral

layer ; and a layer of thin walled nutritive cells in the interior sur¬

rounding the asci. Harper has given a detailed account of the

nuclear behavior in the ascus and the development of the spores;

these are in agreement with his observations on other forms.

After the fusion of two nuclei to form the primary ascus nucleus,

the cytoplasm of the ascus appears spongy but has no large vacu¬

oles. Directly after fusion the chromatin of each of the fusing

nuclei remains independent (retains its individuality), each group

of chromatic structures consisting of about eight strands. The

central bodies finally approach and unite, the nucleoli fuse, the

two groups of chromatin strands intermingle and fusion is com¬

plete. At this time the ascus and the perithecium are about half

grown. A synapsis stage now follows; later the chromatin fibres

gradually become looser and a spireme appears. Harper always

found eight chromatin strands at this stage. A period of rest in¬

tervenes previous to spindle formation, and the nucleus migrates

from the base to the middle of the ascus. Each strand of the fibres

then elongates, becomes bent and forms a chromosome. The chromo¬

some contracts and thickens, leaving a fibrillar connection with the

central body. The chromatin bodies seem to maintain a definite

connection at all times with the central body. The central body

now divides, the daughter bodies separate and a spindle is formed

310 Wisconsin Academy of Sciences, Arts, and Letters.

between them. Eight chromosomes are found in the complete equa¬

torial plate. In the three successive nuclear divisions in the ascus

eight chromosomes always appear. The daughter nuclei of the

first division are smaller than the mother nucleus. They divide at

once or after a short period of rest. At the end of the third divi¬

sion each ascus contains eight nuclei. Only two spores are found

in the ascus; the other six of the eight nuclei resulting from the

three divisions of the fusion nucleus disintegrate. After the third

division the polar asters persist, especially those connected with

the two nuclei that are to form the spore nuclei. A nuclear beak

is formed as in Erysiphe and the spores and delimited by the activ¬

ity of kinoplasmic fibres that radiate from the central body. The

remaining fibres now disappear ; the central body breaks away from

the plasma membrane of the young spore. The nucleus is re¬

formed, becomes spherical, the central body lying against its mem¬

brane, and a spore wall is formed. Harper holds that the primary

ascus nucleus contains quadrivalent chromosomes, which are sep¬

arated into univalent chrosomes in the course of the three succes¬

sive nuclear divisions. The fusion of nuclei in the ascus is a vege¬

tative process and the ascus may be regarded as a spore mother cell.

Harper considers that there is a true alternation of generations in

the Ascomycetes.

I have taken up the work of Harper in detail as it represents

the development of de Bary’s fundamental conceptions in addition

to Harper ^s own interpretation of the phylogeny and morphology

of the fungi. His work developed the whole field of fungous mor-

thology far beyond the grasp of the men of de Bary ’s time and his

views have been, until recently, accepted as the best interpretation

of the cytological processes concerned in the life history of the

fungi. Dangeard (1903-04) on the other hand, inclines toward

the views of Brefeld, regarding the sex organs as non-functional

and the ascogenous hyphae as parthenogenetic outgrowths of func¬

tionless sex organs. The only nucleus fusion is in the ascus. He

maintains that the union of gametes in the ascus has all the char¬

acteristics of a sexual fusion and the ascus all the characteristics

of a sexual organ. The chromosome number doubled by fusion in

the ascus is reduced by the first division, and the two succeeding

divisions are homotypic.

Many contributions by various workers have been added to the

literature substantiating one or the other of these views, but in the

main supporting the ideas of Harper. Kecently, however, a paper

Frey — Physiology of Venturia Inequalis.

311

by Clausen (1912) appeared in which he sets forth some new views.

Clausen adopts an entirely new viewpoint. His theories combine

the ideas of de Bary, Harper and Dangeard, and, if substantiated,

will form a distinct advance in our knowledge of the sexuality of

the fungi. Clausen retains the idea that the ascogonium and

antheridium are sexual organs and that only one nuclear fusion

occurs. This takes place in the ascus and may be considered com¬

parable to the fusion occuring in the basidia and in teleutospores.

The sexual act initiated in the sex organs is completed in the ascus.

It may be well to review in more detail the ideas of Clausen in

order that a complete understanding of the various theories may

be gained. The views of Blackman and Welsford (1912) are op¬

posed to those of Clausen and will be reviewed later. Clausen

(1905) described the formation of the sex organs in Boudiera.

Short branches arise from the vegetative hyphae which branch

dichotomously. The branches then twist about each other forming

spiral coils, two branches in a coil, one forming the ascogonium and

trichogyne, the other the antheridium. The ascogonium contains

two or more nuclei. Ascogenous hyphae arise from the ascogonium

after fusion with the antheridium has taken place. Nuclear be¬

havior and the formation of the asci is similar to that of Pyronema

confluens. Clausen (1906) states his belief in the sexuality of the

fungi and opposes the views of Dangeard.

In a paper published in 1907 Clausen seems to have changed his

views completely. He finds that the sexual organs of Pyronema

confluens function, but nuclear fusion does not take place in the

ascogonium. The male nuclei pass into the ascogonium, in the man¬

ner described by Harper, and a male nucleus pairs with a female

nucleus but no fusion occurs. Conjugate divisions of the nuclei

may occur as they migrate into the ascogenous hyphae. The ascus

is formed in the manner described by Harper. The nuclei that fuse

to form the primary ascus nucleus are, perhaps, direct descendants

of the male nucleus on one side, and the female nucleus on the

other, which paired in the ascogonium.

Clausen (1908) states that sexual fusion occurs in Saproleginia

monoica. Degeneration of all the nuclei of the oogonium except

those at the periphery occurs. The peripheral nuclei divide mitoti-

cally and from the daughter nuclei uninucleate eggs are formed.

The antheridia apply themselves to the oogone, a pore is formed

and nuclear fusion takes place between the male nucleus and the

312 Wisconsin Academy of Sciences ^ Arts, and Letters.

egg. Each egg has a centrosome which may function like the cen¬

tral body of the Ascomycetes described by Harper.

In a later work Clausen (1912) confirms the results of his earlier

research on Pyronema confluens. He agrees with Harper as to

migration of nuclei from the antheridium to the ascogonium, but

does not believe that nuclear fusion takes place in the ascogonium.

The nuclei pair and migrate into the ascogenous hyphae. The

ascogenous hyphae becomes septate and the cell nearest to the asco¬

gonium may have eight pairs of nuclei, the other cells only one pair.

The penultimate cell lying in the hook usually forms an ascus and

the two nuclei lying in the hook fuse to form the primary ascus

nucleus. In some cases the apical cell at the extremity fuses with

the stalk cell and another ascus is formed. The cell usually form¬

ing an ascus may in some cases by budding produce a hook-like

structure which forms another ascus in the usual way.

The first division in the ascus is heterotypic and the nuclei con¬

tain twelve chromosomes after the division. The second division

is homotypic, twelve chromosomes appear at the equatorial plate

and twelve pass to the poles. During the third division the astral

rays appear at the poles, and twelve chromosomes pass to each pole.

Clausen finds no centrosome as described by Harper in Phyllactinia.

He believes the delayed fusion in Pyronema confluens may be due

to toughness of the nuclear membrane. Clausen adheres to the

idea of sexuality in Erysiphe and Sphaerotheca and holds that the

ascus is a spore mother cell as stated by Harper. In accordance

with the views of Clausen the Ascomycetes are similar to the

Basidiomycetes and their phlogeny must be conceived from a new

standpoint, preferably from some oomycete. It is simpler to con¬

sider the Ascomycetes from the viewpoint of Clausen but to do so

we must disregard the work of so many investigators who believe

they have observed nuclear fusion in the ascogonium, especially the

critical work of Harper, that more evidence is necessary before

these facts can be considered established.

Another point of view which may be considered as a reversion to

the ideas of Brefeld has been brought out by Blackman and Wels-

ford (1912) working with Polystigma rubrum. They maintain

that the sexual organs have degenerated and that the ascogonium

develops but does not function. The asci arise from the vegetative

hyphae and the development then proceeds as described by Harper,

Clausen, and others.

Frey — Physiology of Venturia Inequalis.

313

It is necessary at this point to take up the work of other investi¬

gators that we may determine to what extent the theories proposed

by the leading workers or pioneers have been substantiated. Nichols

(1896) holds with Harper that sexual fusion occurs in the Ascomy-

cetes, but it may not occur in all of them. In Ceratostoma no sex¬

ual fusion occurs and in Teichospora aspersa and Teichosporella sp.

the ascocarp is formed by the division of a single hyphal cell of the

mycelium. The ascus arises from a single central cell.

Wager (1899-1900) described the formation of a single sexual

nucleus in the ooplasm of the oogonium of Peronospora parasitica.

The remaining nuclei of the female organ pass into the periplasm

and degenerate. Both male and female nuclei undergo mitosos

before fusion and a central body is present. The zygote is uninuc¬

leate. Stevens (1899) holds that the oogonia and antheridia of

Albugo bliti are multinucleate when formed but later the oogonium

is differentiated into oosphere and periplasm. All the nuclei pass

into the periplasm and undergo mitosis. The nuclei that lie at the

boundary between ooplasm and periplasm give one daughter

nucleus to the ooplasm, the other to the periplasm. Fertilization

occurs in this species ; the male nuclei enter the ooplasm by means

of the antheridial tube and fuse in pairs with the female nuclei.

Several sex organs are produced on Cladonia crispum. Baur

(1898) thinks fertilization is necessary to produce asci. Baur

(1901) states that he observed fertilization of the carpogonium by

spermatia in Parmelia acetabulum. Parmelia and Cladonia accord¬

ing to Baur (1904) have ascogenous hyphae arising from the carpo¬

gonium. He observed fusion of spermatia with trichogynes in

Anaptychia and Endocarpion but could discover no fusion in

Solarina and declared it to be apogamous.

Juel (1902) holds that Dipodascus, which has multinucleate sex

organs arising side by side from a single hypha, has functional

oogonia and antheridia. One large male nucleus passes into the

oogonium by a pore and fuses with a large nucleus in the oogonium.

The oogonium produces a single ascogenous hypha, the latter pro¬

ducing a single ascus. The nuclei arising from the fusion nucleus

decrease in size with each division and finally pass into the ascus.

Juel thinks that Dipodascus is comparable to Albugo in the forma¬

tion of the egg. Sexual fusion occurs in Gymnoascus reesii and

Gymnoascus candidus according to Dale (1903). The ascogenous

hyphae arise from cells which have paired and fused.

314 Wisconsin Academy of Sciences, Arts, and Letters.

Barker (1903) describes sexual fusion in Monascus. The sex

organs are small branches arising from the same hyphae and lie

side by side. The oogonium consists of three cells; the stalk cell^

central cell, and trichogyne. The male nuclei pass into the tricho-

gyne by a pore and fuse in pairs with the female nuclei. The

ascogenous hyphae arise from the central cell by budding but later

grow back into it due to the pressure of the perithecial hyphae.

The asci therefore appear to originate within the central celL

Barker tries to show that relationship exists between the Ascomy-

cetes and the Oomycetes. He believes that the nuclei left in the

periplasm of the oogonium of Albugo Candida are comparable to

those which degenerate in the central cell of Monascus.

Guilliermond (1903-04) points out that nuclear fusion accom¬

panies cell fusion in some of the yeasts. This fusion he considers

as sexual. He finds that conjugation of conidia and nuclear fusion

precede spore formation in the Schizosaccharomycetes and Zygo-

sacchromycetes. Two cells are connected by a tube and in the tube

nuclear fusion occurs. This is followed by one division and the

daughter nuclei migrate, one going to each of the two original cells.

Four spores are formed after the nucleus has made two successive

divisions.

Maire (1903-04) and Dangeard (1903) hold that all Ascomy-

cetes have four chromosomes. Guilliermond finds that the chromo¬

some number in the nuclei of several Ascomycetes, Peziza vesiculo-

sis and Peziza catinus, is not the same; therefore, the chromosome

number varies with the species and the statements of Maire amd

Dangeard are invalid. Harper (1896, 1905) finds that the chromo¬

some number varies with the species. He considers that there is

a true alternation of generations in the Ascomycetes, the division

in the ascus being purely vegetative, and states that it is impos¬

sible to determine when the two reductions in chromosome occur.

It may be in the first and second division or in the second and third.

The chromosomes are quadrivalent as they pass to the poles in the

first division and bivalent in the second. The chromosome number

remains constant during the three divisions, according to Harper

(1905). The three divisions must, therefore, be necessary as two

reductions must occur, and possibly arose in connection with in¬

hibited cell and nuclear division due to increase in the nutritive

material of the ascus. Fraser and Welsford (1908) hold that the

first division of the primary ascus nucleus is the “meiotic” or re¬

ducing division, the synapsis occurring after the first contraction.

Frey — Physiology of Venturia Inequalis.

315

The third division is also a reducing division compensating for the

vegetatve fusion in the ascus. The tetravalent chromosome num¬

ber before the first division in Otidea aurantia is four and follow¬

ing the first division each nucleus contains two chromosomes. In

Peziza vesiculosa the first and second divisions constitute a

“meiotic” phase. Eight chromosomes travel to the pole after the

first division and on the spindle of the second, or homotypic divi¬

sion, four chromosomes appear and the resulting nucleus contains

four chromosomes. Following the third, or ^ ‘ bracymeiotic ” divi¬

sion, the nucleus contains four chromosomes which are regarded as

univalent.

Fraser and Brooks (1909) hold that the first division in the ascus

of Humaria granulata is synaptic and eight chromosomes migrate

to each pole. In the second division four chromosomes appear and

four migrate to each pole. The union of chromosomes takes place

after the sexual, or pseudo-apogamous fusion in the ascogonium.

The chromosomes of the asexual fusion, the fusion in the ascus, re¬

main apart until the heterotypic division is complete and then pair

in the prophases of the homotypic, or second division, separating

in the “ brachymeiotic ”, or third division in the ascus. Ascoholus

furfuraceus is similar to Humaria granulata except that in the for¬

mer vacuoles play an important part in delimiting the spores.

In Lachnea stercorea the nuclei fuse in pairs in the ascogonium

and the chromosome number of the fusion nucleus is double that of

the vegetative cell nucleus. In the second fusion in the ascus the

chromosome number is doubled again making eight chromosomes in

the primary ascus nucleus. The first division in the ascus is het¬

erotypic and four univalent chromosomes appear at each pole. The

second division is homotypic and four chromosomes pass to each

pole. During the prophases of the third division there are still

four chromosomes. In the metaphase the four chromosomes are

still present but they do not divide and two chromosomes pass to

each pole. The third division is, therefore, brachymeiotic and dif¬

fers from meiosis in that there is seldom a synaptic contraction

visible. When contraction does occur the reduced number of

chromosomes is present in the prophase. If pairing occurs in the

second prophase, reduction occurs in the second division. A sec¬

ond contraction never occurs during brachymeiosis. The absence

of any pairing in the third division may indicate that the asexual

fusions have little effect on the fungus. Perhaps the nuclei are

too closely related. The spores are delimited after the manner

316 Wisconsin Academy of Sciences, Arts, and Letters.

described by Harper with the exception that no fusion of kino-

plasmic fibres was observed.

Olive (1905) finds that the central cell of Monascus described

by Barker, is the nurse cell. The ascogenous hyphae arise from

the trichogyne and grow into the central cell and form asci in the

interior. Schikorra (1909) states that Monascus has paired nuclei

and nuclear fusion takes place only in the ascus.

According to Trow (1895) there is no similarity in the develop¬

ment of Saprolegnia diviea to that of some of the Ascomycetes

which have been described. He finds that the fertilized nucleus

undergoes division into oospores directly after fusion. No nuclear

division or fusion occurs in the sporangium. In the oogonium and

the antheridium each nucleus undergoes one reduction before

fusion.

Trow (1901) finds that the ooplasm of the mature oogonium of

Pythium ultimum contains one nucleus, the remaining nuclei pass

into the periplasm and degenerate. One male nucleus from the

antheridium fuses with the nucleus in the ooplasm; fusion of the

nuclei, however, is delayed until the oospore wall is formed.

Christman (1905) described the process of fertilization in

Caeoma nitens. Pustules form containing hyphae with short thick

cells, each cell having a single nucleus. The single nucleus divides,

the cell elongates and a distal cell is cut off which dwindles in size.

The basal cell keeps on growing and inclines toward another basal

cell. Fusion occurs by the formation of a pore in the upper part

of the adjacent sides making the apical ends continuous. The two

nuclei lie in the apical end and conjugate divisions follow. Two

daughter nuclei wander back into their respective cells, but the

original nuclei remain side by side, move to the distal end and are

cut off by a cell wall. The cell formed is an aecidiospore mother

cell which may divide repeatedly. In the paired basal cells the

process may be repeated.

Faull (1905) finds that the ascus of Sordaria fimicola does not

always arise from the penultimate cell, but in many cases from the

ultimate. In Sordaria humana and Podospora acerina the asci

arise from the terminal cells of ascogenous hyphae. He holds that

the ascus is homologous to the zoosporangium of the Oomycetes.

Faull (1911-12) states that only one nuclear fusion occurs in the

Laboulbeniales. In Lahoulbenia chaetophora and Lahoulhenia

gyrinidarium no antheridia are produced. By a series of divisions

the binucleate ascogenic cells are produced. The ascogenic cells

Frey — Physiology of Venturia Inequalis,

317

bud, and the buds form the asci. The nuclei of the ascogenic cells

divide simultaneously and one of the daughters of each mother

nucleus passes into the ascus, the pair left in the ascogenic cell

may divide again. The two nuclei that enter the ascus fuse to form

the primary ascus nucleus.

Miyake (1901) states that the multinucleate oogonium of

Pythium de laryanum is differentiated into ooplasm and periplasm.

A nucleus from the periphery migrates into the ooplasm to func¬

tion as a female nucleus, which previous to fertilization divides.

Only one antheridial nucleus functions.

It is interesting at this point to take up the yeasts, a group of

Ascomycetes which have asci but no definite sex organs. Some

yeasts, according to Guilliermond (1912), form the ascus by the

fusion of two vegetative cells. A bud or beak is formed which

fuses with the bud of another cell and the nuclei migrate into the

tube and fuse. There is considerable variation in the manner of

fusion and in the number of spores formed in the ascus. Wager

and Peniston (1910) state that during spore formation in the

Saccharomycetes the nuclear vacuole and network disappear. The

nucleolus becomes closely surrounded by chromatin granules and

then divides into two nearly equal, or equal, daughter nuclei each

of which consists of a portion of the nucleolus with the surrounding

chromatin. Each of the nuclei divides again to form the four

spore nuclei of the ascus.

Guilliermond (1911, 1917, 1918) states that pairings of cells

may occur in some forms of yeasts by the cells forming a beak and

that nuclear fusion occurs, followed by migration of the fusion

nucleus into one of the cells, leaving the other empty. The egg

then forms an ascus. From the similarity of the developments of

yeasts to Endomyces and Eremascus he thinks these forms are affili¬

ated.

From the evidence available no definite phylogeny can be traced

for the yeasts. It appears that only one nuclear fusion takes place

and that the oogonium itself becomes the ascus. In such simple

related forms as Gymnoascus reesii and Gymnoascus candidus Dale

(1903) finds that sexual fusion occurs. The ascogenous hyphae

arise from cells which have paired and fused.

Observations on Saccharomyces cerevisiae and related forms

made by the writer indicate that dumb-bell forms are quite com¬

mon in old cultures, especially if the culture solutions are of high

acidity and unbalanced physiologically. Whether these forms have

318 Wisconsin Academy of Sciences, Arts, and Letters.

any reproductive function has never been definitely determined.

In nearly every case examined the cell contents were vacuolate and

had apparently degenerated. When the cultures were dried no

spores appeared and there was no indication of budding from the

dumb-bell forms when nutrient solutions were added.

Investigations of the Basidiomycetes have brought forth some

very interesting results, those of Christman (1905) having been

considered, especially in relation to the ideas of Clausen on Ascomy-

cetes. Blackman (1904) finds that in Pragmidium violaceum

binucleate cells are formed in the aecidium. The spermatia are

considered functionless. In the aecidium a group of special cells

form the fertile cells; the fertile cells divide, each one cutting off

a sterile cell above. The nuclei that' enter the fertile cells migrate

from undifferentiated mycelial cells and the binucleate fertile cell

then produces aecidiospores which are binucleate. Nuclear fusion

does not occur until the teleutospore is formed. Fusion is followed

immediately by reduction and the formation of sporidia. Similar

processes were observed in Gymnosporangium clavariaeformae.

Blackman and Fraser (1906) state that the fertile cells of

Phragmidium violaceum are female gametes which are fertilized

by vegetative cell nuclei instead of spermatia. In Puccinia poarum

nuclear migration begins before the fertile cells are formed. The

nuclei migrate from one vegetative cell to another and later

to the fertile cells. Two vegetative cells may form a fertile cell by

fusion. In Melampsora rostrupi basal cells in the aecidium pair

and form aecidiospores which are binucleate. Puccinia mal-

vacearum is apogamous. From all the forms studied they conclude

that nuclear fusion does not occur in the aecidiospore. The con¬

jugate nuclei fuse in the teleutospore and reduction occurs

during the formation of sporidia, a process which may be consid¬

ered a true alternation of generations. In Humaria rutilans no

male cells function, but the female nuclei of the multinucleate

aseogonium fuse in pairs. Whether this process can be considered

analogous to that of Phragmidium violaceum is questionable.

Bensaude (1918) investigated the nuclear behavior in certain

Basidiomycetes. She finds that the mycelium is heterothallic, a

-f- spore and a — spore must develop in sufficient proximity in

order that fusion of the cells of the hyphae can be accomplished.

Binucleate cells are formed but' fusion of nuclei does not take place

until the basidium stage is reached. Fitzpatrick (1918) and Gil¬

bert (1910-11) (1921) have found that the hyphal cells of the

Frey — Physiology of Venturia Inequalis.

319

Basidiomycetes usually have a single nucleus in the early stages;

later two nuclei may be found. The spores are uninucleate. The

cells immediately beneath the hymenial layer always contain two

nuclei. Fusion of nuclei occurs in the basidium. Gilbert (1921)

states that the young spores of Dacromyces are one celled and

uninucleate ; later division occurs and an eight celled spore is

formed. Uninucleate hyphae are produced from these cells.

The spermagonia and ascogonia of Polystigma ruhrum are func¬

tionless according to Blackman and Welsford (1912). They de¬

scribe the formation of functionless spermatia on terminal hyphae.

They state that large uninucleate cells surround the ascogonium

but its cells are multinucleate. The ascogonium degenerates and

perithecial hyphae arise in the neighborhood of the ascogonium,

formed perhaps by vegetative hyphae, or special hyphae arising

from or about the base of the degenerating ascogonium. The spe¬

cial hyphae are finger-like and grow together in a conical mass,

the apex usually directed toward the lower epidermis of the leaf.

The nuclei of the cells are arranged in pairs. From these hyphae

the ascogenous hyphae arise, the nuclei of the latter arranged in

pairs. The asci probably arise from the penultimate cells of the

ascogenous hyphae for in these cells nuclear fusion seems to occur.

There is evidence of nuclear fusion when the ascogenous hyphae

are differentiated.

Welsford (1907) finds no antheridia taking part in the sexual

fusion of Ascoholus furfuraceus. The female sex organs consist

of a row of 6-10 cells. The middle cell is at first uninculeate but

later becomes multinucleate and develops the ascogenous hyphae.

The remaining cells of the ascogonium connect with the middle cell

by pores through which the nuclei migrate and fuse in pairs in the

ascogenous hyphae. The penultimate cells of the ascogenous

hyphae contain two nuclei which fuse and from this cell the ascus

is formed in the usual way.

Fraser (1913) states that the archicarp of Lachnea creta devel¬

ops on one of the larger filaments ; it forms two or three coils and

becomes septate, each cell becoming multinucleate. The archicarp

becomes differentiated into three regions; the stalk, the coiled

ascogonium, and the septate trichogyne which elongates and

branches. Antheridia are not present. Large pores form between

the cells of the multinucleate ascogonial cells and nuclei migrate

from one cell to another. The ascogenous hyphae arise from the

cells of the ascogonium and asci are formed. Keduction occurs

320 Wisconsin Academy of Sciences, Arts, and Letters.

during the first division in the ascus and in the telophase of the

third division.

Many investigators fail to find antheridia in the forms they

studied. Eamlow (1906) holds that Theleholus stercoreus is apoga-

mous and no antheridia are present. Cutting (1909) failed to find

antheridia in Ascophanus carneus. He also states that any cell

of the multinucleate and multicelled ascogonium may produce

ascogenous hyphae.

Eamlow (1914) holds that the nuclei of Ascophanus carneus d©

not fuse but are paired in the ascogonium and wander into the

ascogenous hyphae paired. Each cell of the ascogenous hyphae

receives a pair of nuclei but only one cell develops into an ascus.

Observations indicate that Ascoholus immersus is similar, except

that each ascus contains sixteen spores.

Overton (1906) states that the ascogenous hphae of Thecotheus

pelletieri may arise from any cell of the ascogonium. The cells of

the ascogonium are not connected by pores. The ascus may con¬

tain sixteen or thirty-two spores. This seems to indicate that three

divisions are not necessarily the limit and the number of divisions

may have no special significance.

According to Brown (1908) the antheridium of Pyronema con-

fluens is functionless, one fusion of nuclei occurs in the ascogonium.

He holds (1910) that only one nuclear fusion occurs in Leotia

luhrica, that being the fusion in the ascus.

The filaments of Helvella elastica form ascogenous hyphae ac¬

cording to McCubbin (1910). The nuclei of the cells in the fruit¬

ing body are paired. This organism is of great interest as no sexual

organs function in the production of asci.

Carruthers (1911) holds with Harper that two nuclear fusions

are necessary. In Helvella crispa the chromosome number in the

vegetative nuclei is two and in the ascogenous hyphae four. The

chromatin does not mingle during the first part of meiosis; only

in the spireme stage is there a union and then perhaps an imperfect

one. The chromosome number in the primary ascus nucleus is

eight. Two contractions occur during the first, or heterotypic

division and chromatin is extruded. The first and second divisions

constitute a meiotic phase. The third division is simply a separa¬

tion of nuclei which fused in the ascus. Sixteen nuclei may be

formed but only eight spores are produced.

Frey — Physiology of Venturia Inequalis.

321

Bachman (1912-13) finds that the archicarp and spermatia of

Collema pulposum function. The trichogyne grows toward the

spermatia, coils about it and fertilization is accomplished.

Fraser (1913) describes the trichogyne of Lachnea creta. She

holds that this form is apogamous, no antheridium having been

found.

Nienburg (1914) opposes the views of Blackman and Welsford

on Poly stigma ruhrum. He states that the archicarp has a coil at

its base and a chain of long multinucleate cells forming a tricho¬

gyne. A long multinucleate cell, the antheridium, applied itself

to the long uninucleate cell, the ascogonium, the walls dissolve and

the nuclei migrate into the ascogonium where one increases in size

and becomes the male nucleus. When this male nucleus is equal

to the female nucleus in size the remaining cells of the archicarp

have degenerated. The ascogenous hyphae arise from the ascogo¬

nium and contain paired nuclei but the development of the aseus

could not be followed. The trichogyne is not a sexual organ, but

may be regarded as a vegetative or nutritive structure. Nienburg

did not observe fusion in the ascogonium or in the ascogenous

hyphae and believes the only fusion that takes place is in the ascus.

Brooks (1910) described the development of Gnomonia erythros-

toma. He finds spermatia are produced but they no longer func¬

tion. Several hyphae entwine and a perithecium is formed. Later

a trichogyne appears. A coil of deeply staining cells forms the

ascogonium which, however, degenerates. The ascogenous hyphae

arise de novo, probably from the vegetative hyphae. The only

nuclear fusion takes place in the ascus. Eeduction occurs in the

first division

Moreau (1919) working with Pelligeracees could find no sperma¬

tia which possessed the power to function as male cells. The

ascogonium is multinucleate and no connection between the

spermatia and the ascogonium was observed. The fusion in the

ascus is the only nuclear fusion taking place.

Investigations of the development of Pyronema confluens were

made by Brown (1915). He states that the form he studied pro¬

duces a trichogyne which, however, does not fuse with the antheri¬

dium. He observed no fusion of nuclei in the ascogonium or in the

ascogenous hyphae. The only fusion is in the ascus.

Welsford (1915) states that the reason paired nuclei are found

in the ascogonium and ascogenous hyphae is due to better nutri¬

tion which causes rapid division. Nuclei in cells filled with large

322 Wisconsin Academy of Sciences, Arts, and Letters.

quantities of food material proceed to divide, and before the nuclei

can migrate another division occurs. This accounts for the figures

observed by Clausen. There is, therefore, no significance to be at¬

tached to nuclei in this condition. In poorly nourished mycelia the

paired nuclei are absent, as the nuclei have time to move apart

before another division takes place.

Killian (1915) described the origin of the archicarp in Venturia

inaequalis. He finds that a single coiled hyphae enlarges in the

perithecium and forms an ascogonium. A trichogyne is produced

which protrudes from the perithecium and grows toward the

antheridium, but remains surrounded by a layer of cells. The two

organs come into contact and pores are formed through which the

male nuclei pass into the trichogyne. The walls of the ascogone

are dissolved and the male nuclei pair with the female nuclei and

aggregate in the end or basal cells of the ascogonium. Further ob¬

servations were not made as the organism offers considerable diffi¬

culty for study.

Cryptomyces pteridis represents a type in which the trichogyne

no longer functions. Pairing of equivalent cells in the fruiting

body occurs and one of the cells may be regarded as an egg. The

nucleus from the male cell migrates into the oogonium through a

pore, but fusion with the egg nucleus does not take place for some

time. The cell containing the egg does not produce ascogenous

hyphae, although it elongates in an attempt to do so, but itself be¬

comes the ascus. The process is essentially like that described by

Guilliermond for yeasts. Only one nuclear fusion occurs.

The phylogeny of the Ascomycetes has been the subject of con¬

siderable controversy. Sachs (1868) and de Bary (1870-1899)

noted the similarity of the sex organs of the Ascomycetes and the

Plorideae. Brefeld (1875) held that the Ascomycetes were derived

from the Phycomycetes. He believed the sporangium evolved into

an ascus, a view which Harper’s work has shown to be extremely

improbable. De Bary, (1881-84) states that the Ascomycetes may

have originated from the Peronospora. He suggests that certain

forms like Eremascus may have been derived from the Mucorales.

Bucholz (1912) states that Endogone is a Phycomycete. The

structure developed from the fusion of two unequally differentiated

sex cells he designates a ^^zygosporocarp”. Clausen (1912) pre¬

fers the view that the Oomycetes gave rise to the Ascomycetes

Dodge (1914) in a very able discussion of the literature concludes

Frey — -Physiology of Venturia Inequalis,

323

that the Ascomycetes must have arisen from the Florideae. He

bases his conception on the similarity of the archicarp, especially

the trichogyncj to the corresponding organs in the Florideae. The

difficulty in accepting this view arises from the lack of evidence

as to the origin of the ascus, no equivalent organ having been found

in the Florideae^ unless we consider ooblastema filaments as homol¬

ogous to ascogenous hyphae. Speculations^ at present, are only

worth while in so far as they stimulate further research. It is

entirely possible that a large number of forms are present in na¬

ture, still undiscovered, which will supply the missing links. If

such forms cannot be found our theories, like those on the origin

of man, will ever remain the subject of dispute. The search for

homologies in the evolution of the sex organs offers the most plaus¬

ible method of attack. One would expect the vegetative organs to

respond to immediate environmental changes whereas the genera¬

tive organs or cells may not fundamentally alter.

Atkinson (1916) in a very detailed discussion attacks the views

of Sachs on the origin of the Ascomycetes. He finds it impossible

to accept the view of the origin of the Ascomycetes as being de¬

rived from the red algae. There is, according to him, no homology

existing between the archicarp and the procarp. The ascus might

develop from forms similar to Dipodascus by branching of the

zygogametangium. By reducing the number of spores in the ascus

of an organism of this type forms like Eremascus and Endomyces

are produced. The trichogyne, Atkinson contends, is not a func¬

tional organ in most of the Ascomycetes studied, and may have de¬

veloped rather as an outgrowth, or beak of the oogonium, a condi¬

tion often found in certain oomycetes when the female organ is

stimulated. In the Laboulbeniales the trichogyne is highly devel¬

oped but does not always function. The septa prevent nuclear

migration and thus it occurs that many Ascomycetes' that retain

the sexual organs may not have nuclear fusion. Many Ascomy¬

cetes that retain functioned gametangia do not possess a trichogyne.

In cases in which the sperm enters the trichogyne there is no evi¬

dence that this process is necessary to initiate disintegration, as it

may proceed without it and cannot be cited as proof of fertilization.

In such forms as Monascus he regards the antheridium as an

elongated terminal cell of a hyphae and homologous with a chain

of conidia. This conception harmonizes such anomalous forms as

Collema in which the trichogyne fuses with a spermatium or

conidium.

324 Wisconsin Academy of Sciences, Arts, and Letters.

There is no evidence, according to Atkinson, to substantiate the

view that ooblastema filaments are phylogenetically related to the

ascogenous hyphae of the Ascomycetes. Pore formation is not

connected with the presence of a trichogyne or antheridium as it

is formed in Ascobolus, Lachnea creta and Polystigma ruhrum

forms in which one or both of these organs are absent. In the

fusion of ooblastema filaments with auxiliary cells the diploid

nucleus of the former never fuses with the haploid nucleus of the

auxiliary cell but they actually repel each other.

Atkinson doubts whether the nuclear fusion in the ascus may

be regarded as purely vegetative and believes it has greater signifi¬

cance. He cites Dangeard (1897) and W. H. Brown (1909) on

Pyronema to substantiate his views that the sex organs do not al¬

ways function. In Lachnea scutellata Brown (1911), the antheri¬

dium is absent, as also in Ascophonas carneus, and Ascobolus im-

mersus. Ramlow (1914) and W. H. Brown (1910) maintain that

no nuclear fusion occurs in the ascogonium of Leotia and Paul

(1911-12) working on Laboulbenia holds the same view. The

archicarp is absent in some forms and vegetative cells form

ascogenous hyphae as in Gnomonia erythrostoma Brooks (1910)

and Helvella elastica MucCubbin (1910). In Polystigma rubrum

the archicarp, according to Blackman and Welsford (1912), does

not function. In Collema pulposum observed by Bachman (1913),

according to Atkinson, there is no evidence that the nucleus of the

spermatia migrate to the ascogonium, although disintegration fol¬

lows. The archicarp of Gnomonia erythrostoma Brooks (1910)

has also ceased to function. If we accept the views of Clausen

(1912) these results have no significance as only one fusion occurs

and the sex organs are no longer necessary.

It is not difficult to bring up negations but the articles quoted

by Atkinson to support one statement refute his claims in another.

His argument that the trichogyne is of no fundamental importance

from a phylogenetic standpoint and that the sex organs of many

Ascomycetes are functionless or atrophied makes one doubt the

wisdom of placing so much faith in the immutability of certain

organs of the Phycomycetes as he apparently has. Progression and

degeneration may go on together, and to discard one view or fact

because it does not harmonize with preconceived notions does not

solve the matter. More evidence is required before a genralization

can be made.

Frey — Physiology of Venturia Inequalis.

325

From the literature reviewed one obtains no definite or uni¬

formly accepted ideas of the phytogeny or cytology of the ascomy-

cetes. The followers of de Bary have not succeeded in establishing

his views beyond dispute. Harper’s ideas are in line with those of

de Bary and opposed to those of Brefeld and Dangeard. Clausen

agrees with Harper except as to the fusion in the ascogonium.

Fraser and Welsford, and Blackman and Welsford have taken a

middle ground. On the one hand they resurrect the ideas of Bre¬

feld as to the function of the sex organs. They agree that two

nuclear fusions occur and describe the three divisions in the ascus.

The difficulty of following nuclear behavior and the development

of the sex organs may be the reason for much disagreement. A

large number of forms have been studied, but it is possible that

we have only begun the cytological work. The work of Harper

indicates the complexity of the problem.

If we accept the views of Clausen it must be only upon incon¬

trovertible evidence as the double fusion, if it occurs, may be con¬

sidered an attempt in evolution which not other plants have under¬

taken. It would simplify the matter greatly to find that only one

nuclear fusion occurred in the life history of the Ascomycetes, but

mere simplicity should not mislead us nor prevent us from accept¬

ing conditions as they exist.

The divisions in the ascus have been studied in great detail by

Harper (1905) and Fraser and Welsford (1909). Only a small

number of forms have been studied and more research may bring

out many more interesting phases. In some of the yeasts and lower

Ascomycetes only two divisions in the ascus are recorded whereas

in Thecotheus , (Overton 1906), Ehyparobius, (Barker 1905), and

Ascoholus immersus, (Ramlow 1914), four and five divisions of

the primary ascus nuclei may take place.

De Bary’s idea that the sex organs of the fungi are functional

becomes more plausible as cytological evidence accumulates in re¬

gard to the life history of the Lichens and Florideae. The law

of function in a structure so important as the sex organs can only

be viewed as due to a long cycle of evolutionary changes, to nutri¬

tional disturbances or to a sudden mutation. Paleobotanical

studies indicate that the sex organs suffer little change through

long cycles of time, whereas the vegetative organs may respond

quickly to new environmental influences. Loss of function or

suppression of an organ can scarcely be explained by the meager

knowledge of physiology at our disposal. Such changes may be

326 Wisconsin Academy of Sciences, Arts, and Letters.

influenced by nutrition, moisture, and temperature changes, but

there is no evidence to indicate that such change of suppression

is permanent.

It may be possible to view the ascus as homologous to the ooblas-

tema of the red algae, but the fusion of the auxiliary cell with the

filament brings in a phase of development which is not eEisily har¬

monized with the development of the ascus in the Ascomycetes.

Morphological Observations

Venturia inaequalis (Cooke) Winter is an Ascomycete classi¬

fied as a Pyrenomycete. Lindau (Engler and Prantl) places it in

the order Sphaeriales, family Pleosporaceae. Fries (1819) named

the conidial stages of the organism Spilocoea pomi. Fuckel trans¬

ferred the fungus to the genus Fusicladium. Cooke (1866) de¬

scribed the stage in which asci are formed and named the organism

Sphaerella inaequalis. Winter transferred the fungus to the

genus Venturia naming it Venturia inaequalis. Aderhold (1897)

connected the Fusicladium stage with the perithecial stage known

as Venturia inaequalis by Winter and, not aware that Winter had

transferred the organism, placed it in the genus Venturia naming

it Venturia inaequalis (Cooke) Aderhold.

Venturia inaequalis is parasitic on the fruit and leaves of species

of Pyrus and produces conidia during the spring and summer.

As the leaves die and fall to the ground the mycelium, which is

almost lacking and exists chiefly under the cuticle, penetrates the

leaf tissue and during November in Wisconsin perithecial forma¬

tion begins. The perithecium is not embedded in a stroma. The

ascospores are found in March and April but do not ripen prepara¬

tory to discharge until May, discharge becoming active about the

time the blossoms of the apple are opening.

The fungus is saprophytic from the time the leaf falls until

the perithecium is formed. It is not known if the mycelium con¬

tinues to function after the ascospores have been formed.

The mycelium is septate and branches irregularly. The un¬

inucleate cells are from ten to forty microns in length and five to

eight microns in diameter. The conidia are produced on eonidio-

phores when the latter are from two to thirty microns in length.

In cultures four or five conidia may be borne in a cluster at the

apex of the conidiophore. As observed on the leaf only one coni-

dium is borne at the apex of the conidiophores. As each coni-

Frey — Physiology of Yenturia Inequalis.

327

dium is formed and abstricted the conidiophore grows further and

produces another conidium, leaving a shoulder or node where

the former was cut olf. In culture several shoulders may be

grouped; apparently the interspaces between the shoulders are

extremely short, causing the conidia to appear in groups. The

conidia are two-celled in most cases, each cell being uninucleate.

The conidia measure about seven to ten microns in diameter and

twenty-eight to forty microns in length. The color of the conidium

is olive or brown.

Perithecial development, as previously stated, begins in autumn.

The first evidence of perithecia was obtained from leaves fixed

November 14, 1916, At that time the perithecia were approxi¬

mately forty to seventy-five microns in diameter. Sections showed

that considerable internal differentiation had occurred as each

perithecium contained several large cells which stained more heav¬

ily than the surrounding tissue. The perithecia are spherical in

form, each consisting of a dense mass of closely interwoven dark

colored hyphae. The diameter of a mature perithecium varies but

may exceed 150 microns. The perithecium may be formed in any

part of the leaf but generally on the side lying uppermost. The

perithecium is generally embedded in the spongy parenchyma and

may be seen with the hand lens as it pushes up forming a small

papilla or dome just before it discharges its spores. The spores

are discharged through the opening known as the ostiole, but

when ripe and suddenly wetted the whole upper portion of the

perithecium may be lifted off. The discharge is produced, so far

as could be ascertained, by the hydration of a colloidal gelatinous

mass surrounding the asci which swells enormously.

The organism was grown in culture and observations were made

to determine the course of development. Oatmeal agar proved

to be a satisfactory medium for growth. Potato agar failed to pro¬

duce perithecia. It was necessary to keep the cultures in the re¬

frigerator to produce perithecia. The observations on the effect

of temperature were first made by Jones (1913). A tube was inoc¬

ulated and allowed to remain at room temperature for several

weeks and then placed in the refrigerator which was held at a

temperature of about 8° C.

Various fixatives were tried but Flemming’s medium proved to

be the most satisfactory. Urea was added to increase the rate of

penetration when the perithecia were large and gelatinous. Fixa¬

tions were made twice each week.

328 Wisconsin Academy of Sciences, Arts, and Letters.

Staining proved to be very difficult as the fungus failed to retain

the stain. Flemming’s triple stain gave good results with young

perithecia. Heidenhain’s iron-alum haematoxylin gave good re¬

sults when counter stained with Orange g. and was used for most

of the cytologieal work.

To determine the development of the perithecium portions of

oatmeal agar containing mycelium of the fungus were placed in

a Van Tieghem cell and kept in the refrigerator. The appearance

of short protuberances on the filaments of the mycelium were

noted, as in figs. 1-6. Two branches arising from the same fila¬

ment' begin coiling about each other forming a compact, coiled

mass. The branches become septate and it was possible to make

out two and three cells. In most cases the coil soon becomes so

intricate that it is impossible to follow its development. After

the coil begins to form it is joined by other branches arising from

the base of the stalk or the filament from which the stalk arose.

Sometimes one branch begins to take a spiral form apparently

without the assistance of another branch. It was also possible

to obtain growth for a period of two weeks in oatmeal agar placed

on a slide and covered with a cover glass, if the slide was placed

in a petri dish and kept in a sterile condition and at low tempera¬

ture, thereby making possible accurate observations of the early

stages. In such cultures one often observes conjugations consist¬

ing of a short branch connecting adjacent or parallel hyphae.

As far as could be determined these structures never gave rise to

perithecia. Sometimes one finds a nucleus in the conjugating tube.

Frequently one finds filaments forming perithecial structures in

great abundance and branching profusely, while the neighboring

filaments exhibit no tendency to branch or to form perithecia.

Sections of the young perithecium, fig. 10, indicate the pres¬

ence of a large coil which stains more deeply than cells of the adja¬

cent hyphae. This coil increases in size as the perithecium grows.

Figs. 7, 8, 9, 10 and 11 show coils within the perithecium with

part of the coil projecting beyond the limits of the wall.

From the similarity of my figures to those of Harper (1905),

Clausen (1912), and Brooks (1910), one could not avoid conclud¬

ing that the large coiled structures staining deeply were archi-

carps. The coil within the perithecium resembles the ascogonium

as described by Harper, and undoubtedly functions as such. There

is evidence that a single branch enlarges to form the ascogonium

as one can often follow the heavy staining cells to the periphery

Frey — Physiology of Venturia Inequalis.

329

of the perithecium, figs. 10, 11a and 14. The tongue-like struc¬

ture emerging from the coil in the perithecium and extending be¬

yond the margin is apparently a trichogyne. During the early

stages the ascogonium and trichogyne seem to be non-septate, fig.

10. After fusion with the antherids the ascogonium becomes

septate and paired nuclei are found in the cells, although some

figures, fig. 14, indicate that the ascogonium may become septate

without the presnce of a trichogyne. Surrounding the trichogyne

are numerous hyphae forming a wall around it. Most of the

apical cells of the antheridial hyphae are swollen, have a bulb¬

like appearance, and arise from the filaments in the neighborhood

of the perithecium. Some of these hyphae are applied to the tri¬

chogyne and seem to fuse with it, figs. 7, 8, 9, 10 and 12. Most

of my figures show these bulb-like cells devoid of nuclei and high¬

ly vacuolate, as if the nuclei had migrated and the cells were dis¬

integrating. Occasionally multinucleate cells were found as in

figs. 7 and 12. These figures seem to indicate that some of these

cells function as antheridia. The antheridium is found closely ap¬

plied to the trichogyne and a pore is formed. Several nuclei lie

immediately within the trichogyne and two nuclei appear to be

within the antheridium in fig. 7. Paired nuclei are seldom found

in the trichogyne. Apparently the nuclei migrate singly toward

the ascogonium, but this may not always be the case for one oc¬

casionally finds paired nuclei in the trichogyne. The nuclei are

approximately the diameter of the vegetative hyphae, 6-8 microns,

and the membrane is sharply differentiated. Occasionally very

little chromatin is visible but a large, dense nucleole is generally

present'. Sections of the perithecium cut after fertilization has

occurred show that the coil has become septate, and that the cells

of the ascogonium have greatly enlarged, stain more densely and

contain two nuclei. Figs. 8, 11 and 12 indicate that the nuclei are

paired. Perithecia found in leaves and those taken from cultures

compare very closely in every morphological detail during all

the stages observed.

In sections made from young perithecia grown on oatmeal agar

one frequently observes the archicarp coiled but non-septate at its

base, as in figs. 7 and 10. The trichogyne is non-septate and ex¬

tends beyond the perithecium. It is not definitely known at what

stage in the development of the coil fertilization occurs. The only

disagreement with this type of structure and the ones described

previously by other workers is the non-septate coil. If fertiliza-

330 Wisconsin Academy of Sciences^ Arts, and Letters.

tion occurred at this early non-septate stage, it would not be neces¬

sary for the nuclei to pass through pores from cell to cell. All

the preparations indicate that fertilization occurs during the non-

septate stage, for later stages show the nuclei paired in the cells

of the coil of the ascogonium, figs. 11, 12 and 13.

An effort was made to discover if nuclear fusion occurred in

the ascogonium. Sections cut from the coils as they formed and

from the perithecium after it is fully mature, including the inter¬

mediate stages, failed to give evidence of funsion. Nuclei are

found in pairs, fig. 12, and later in aggregates of four, fig. 23c.

Figs. 7, 8 and 11 show sections of the perithecium. The ascog¬

onium is coiled, four to six cells being visible, and the nuclei are

paired. Fig. 8 shows the trichogyne still attached and the nuclei

migrating toward the ascogonium each cell of which has paired

nuclei. In none of the preparations of these early stages, how¬

ever, was nuclear fusion apparent. Figs. 10, 11a and 11b indicate

the condition of the structure. The basal or stalk cells are still

visible and extend toward the periphery of the perithecium, while

at the opposite end of the coil the trichogyne emerges. The stalk

may be regarded as the basal remnant of the original branch. At

the time the coil originates it appears as if two branches were

concerned, one enlarging to form the ascogonial coil. What the

function of the other hypha is could not be determined. It may

be nutritive or sexual, but it is impossible to follow the develop¬

ment accurately as the vegetative hyphae enclose the coil and form

a dense covering. The diameter of the perithecium is often 150

microns and one section or series of sections is difficult to inter¬

pret. The ascogonium is coiled and all portions' do not lie in the

same plane. In some figures, 11a and 11b, the coil is shown very

distinctly. The cytoplasm is stained very densely and the nuclei

stand out remarkably well. Each cell is binucleate, the cytoplasm

is slightly granular in each cell. Fig. 12 also indicates the con¬

dition of the ascogonium when the nuclei are paired. As the peri-

thecia enlarge and when they are from eight to nine weeks of

age in cultures, the cells seem to increase in number and sections

often show four nuclei in one cell, figs. 19, 23b and 23c. The

cells of the coil increase in size until the perithecium has reached

its maximum diameter. Following the four nucleate stage a

large number of cells appear and extensive branching is apparent.

There is considerable evidence to indicate that the large cells of the

ascogonium branch, the branches become septate and form the asci,

Frey — Physiology of Venturia Inequalis.

331

figs. 16-20. The large cells of the ascogonium disappear gradually

and lose their capacity to stain more densely than the surrounding

hyphae.

As previously stated, at the time of fertilization the trichogyne

is non-septate. The ascogonium has no definite cell walls. It was

impossible to find sufficient evidence to indicate whether or not

rapid nuclear division occurred in the coil, but a large number

of nuclei are found later, fig. 12, indicating that division occurs.

The evidence points to migration of nuclei from the trichogyne as

initiating the multinucleate condition. There was no nuclear di¬

vision apparent in any sections to indicate that the uninucleate

cells of the ascogonium become multinucleate before fertilization.

In the young trichogyne one can seldom find a nucleus. The as-

cogonial coil contains several nuclei, fig. 10, but after the tri¬

chogyne has fused with the antheridial cells it contains many

nuclei; often a chain of them may be observed in the trichogyne

extending from the apical end to the margin of the perithecium

or beyond, figs. 8-12.

The nuclei of the ascogonium continue to grow in size until the

cells no longer stain heavily, and there is apparent a great in¬

crease in the number of the cells. At this stage the perithecium is

at its maximum size and it is difficult to make accurate observa¬

tions of the various stages of development. Sections cut in March

from material gathered February 27 in the orchard have no heavy

staining material left and the nuclei are not much larger than

the nuclei of the vegetative cells. The contents of the nucleus

which at first stained very deeply become more and more difficult

to stain about the time of ascus formation, figs. 16, 21, 22. A cross

section of the perithecium gives very little indication of differen¬

tiation in any of the tissues not arising from the coil.

There is no evidence to indicate that typical ascogenous hyphae

as described by Harper and Clausen occur. The whole content of

the perithecium, with the exception of a layer of four or five cells

at the periphery, seems to be occupied by asci and gelatinous ma¬

terial. All the large deeply staining cells of the ascogonium have

disappeared, and the space formerly occupied by them is filled

with asci which appear as if they originated from the basal por¬

tion of the perithecium, figs. 24 and 25ab. The only explanation

formulated, and which agrees with the sections observed, is that

apparently there is a breaking up and budding of the large cells

of the ascogonium, forming a large number of small cells, the

332 Wisconsin Academy of Sciences, Arts, and Letters.

initial ascus cells arising from some of these cells. The ascogonium

immediately after fertilization may contain seven or eight cells.

Later stages show the contents staining less densely, vacuoles ap¬

pear and four nuclei are found in some cells. Distortion of all

the cells is apparent and the inner contents of the perithecium are

undoubtedly subjected to enormous pressure by the enveloping

hyphae. By such intimate contact with the enveloping vegetative

hyphae it is possible that the metamorphic tissue within receives

nourishment. The cells of the enveloping tissue are uninucleate

and are pressed into plates or rectangular polygons. At the time

of fertilization they contain considerable food material but later

this disappears.

As the ascogenous cells decrease in size, or rather as they bud

and new cells are formed, the cells so formed are uninucleate oi

binueleate. It has not been possible to determine whether these

cells form asci directly or by budding. The perithecium at this

stage has reached its full size and it would be impossible for further

budding and growth to take place unless some of the enveloping

hyphae are absorbed, which seems, however, to occur at a later

period. Possibly the cells elongate and by a process of rearrange¬

ment without much increase in the space occupied, the asci are

finally developed to their full size.

There is a considerable gap in my series of developmental ob¬

servations extending from the origin of the ascus to spore develop¬

ment. The ascus previous to the time the spores form is elongated

and the nucleus or nuclei lie near the center, figs. 24-25. After

the second nuclear division the ascus elongates still more and the

nuclei migrate toward the periphery. About the time of the third

division the ascus elongates again, the spores are delimited and the

ascus is just wide enough at this time to hold the spore. Its length

is approximately twice that of an ascus in the stage previous to the

first division.

The spores are arranged in a chain and fill the ascus with the

exception of a portion at the base. Practically all of the proto¬

plasm of the ascus is exhausted, having been utilized in spore

formation. A large number of cells forming the inner zone of the

perithecium have been crowded aside, and their contents have dis¬

appeared. Many of the cells appear to be disintegrating. Three

or four layers of heavy-walled cells constitute all that is left of

the perithecium. Sometimes there is a cavity in the region below

the ostiole as the perithecium elongates preparatory to discharg-

Frey — Physiology of Venturia Inequalis.

333

ing spores. The cavity may not be devoid of material as in some

there is evidence of a gelatinous material which may take up

water and swell thereby forcing out the spores. Whether an

enzyme, or enzymes, are necessary to ripen the spores and digest

the walls of the ascus is not known. Attempts to discharge spores

by moistening and warming the perithecium after the spores ap¬

pear to be mature fails to produce a discharge, even if consid¬

erable pressure is applied. When the perithecium is wetted and

then punctured with a needle, considerable discharge occurs. The

conditions for discharge are, however, very complex. During

May the asci contain mature spores but neither wetting nor heat¬

ing to room temperature would discharge the spores. July 6, 1917,

leaves were obtained which had been lying on the sod and had not

completely decayed. They contained perithecia full of asci and

spores. The leaf was taken into the laboratory and placed in a

moist chamber and a slide placed about an inch above the leaf.

A tremendous discharge occurred. Several slides were then pre¬

pared and placed at various temperatures. About 80 per cent

germination at 1-2° C. was obtained, but growth was extremely

slow. At 2-3° C., 96 per cent germination was obtained. At 4°

and 6° about the same per cent of germination resulted, but growth

was more rapid. The optimum germination and growth occurred

at 10-18° C. and then gradually decreased until 22° C. was reached

when very little growth occurred, and the germination was greatly

reduced. It may be well to state that the temperature control was

only accurate to within a degree or more. The experiments con¬

ducted on spore discharge demonstrated rather conclusively that

discharge of spores occurs only during rains or when the leaves

are very wet, and that the discharge is greatest during the first

two hours and gradually lessens. Temperature may influence the

rate of discharge, but owing to lack of data at present it is not

deemed possible to draw definite conclusions.

The ascospores when first delimited appear round or lens¬

shaped. When mature they are two-celled and each ascus usually

contains eight spores. Each cell has one nucleus which is no

larger than the nucleus of one of the cells of the filament. The

wall of the spore consists of two layers. With Heidenhain’s iron-

alum haematoxylin the cytoplasm stains heavily, but the nucleus

is much denser and more heavily stained. Evidently the proto¬

plasm also contains considerable food material.

334 Wisconsin Academy of Sciences, Arts, and Letters,

The ostiole is formed at the apex of the perithecium. The outer

zone of cells becomes disorganized, and some of the cells appear

plasmolyzed or devoid of staining material. Pressure due to

swelling of the gelatinous material within the perithecium may

rupture the upper portion and form the small opening known as

the ostiole. If enzyme activity is necessary to dissolve some of

the tissue, the inability to discharge spores before a certain ma¬

turity is reached might be explained. The gelatinous material

within may require certain chemical changes to increase its swell¬

ing capacity, thereby limiting the period when spores may be dis¬

charged. If this material would simply by swelling cause the

rupture of the perithecium, the discharge would be entirely a

mechanical process. In the ostiole filaments appear as if the dis¬

solution and pulling apart had left some of the fibres intact. The

perithecia at this time are greatly elongated and have the appear¬

ance of forming a bottle neck at the upper end. There is very

little staining capacity left in the tissue comprising the wall;

evidently the cell contents have disappeared.

The nuclear behavior was not followed clearly owing to the

difficulty of staining the chromatin. As previously stated, no nu¬

clear fusion was observed in the ascogonium, although many prep¬

arations of the stages when fertilization occurred and those im¬

mediately following it were made. The nuclei in the ascogonium

after fertilization increase tremendously in size and are paired.

Two and four nuclei are found in a cell. The contents of the nu¬

cleus may stain densely or scarcely at all. No attempt was made

to follow the behavior of the nuclear contents during growth and

division. Some sections prepared at the time the spores are de¬

limited in the ascus indicate that a central body exists whose

fibres function in cutting out the spore, in a manner similar to

that described by Harper (1897) (1905).

The work of Killian (1915) indicates that he found sections

showing the large trichogyne and the coiled ascogonium. My

preparations failed to show the antheridium branched as he figures

it, but some of my sections show lobes of the tip or apical cell of

the antheridium closely applied and partly surrounding the tri¬

chogyne. The nuclei aggregate near the pore and pass into the

trichogyne sometimes in bead-like rows. Most of my sections in¬

dicate that the ascogonium is composed of five to eight cells and

that these cells are formed after fusion with the antheridium.

Killian’s figures do not indicate that the ascogonium is divided

Frey — Physiology of Venturia Inequalis.

335

into cells, but his description leads one to believe that he observed

pores in the cell walls through which the nuclei from the tri-

chogyne passed, indicating that the archicarp is septate at the time

of fusion.

The study of this organism gives further evidence of the correct¬

ness of de Bary’s views in regard to the sexuality of the fungi. It

is impossible to agree with Blackman and Welsford’s idea that

the sexual organs of the fungi do not function. The appearance

of the trichogyne in connection with the perithecium and its con¬

jugation with the antheridium cannot possibly be considered as

non-functional. The trichogyne passes from a non-nucleate stage

to one containing many nuclei at the time of fusion with the an¬

theridium, and the cells of the ascogonium become binucleate,

processes which must be shown to have no relation to the develop¬

ment of the asci if we are to accept the theories of Brefeld, Dan-

geard and Blackman and Welsford.

The figures of Blackman (1913) are similar to the stages ob¬

served in Venturia inaequalis. The similarity of the ascogonium

of Collema to that of some of the Pyrenomycetes suggests that the

Ascomycetes and Lichens may be rather closely related, more

nearly, perhaps, than the Oomycetes and Ascomycetes. Further

observations of the Pyrenomycetes may confirm the theories of

Harper and Dodge as to the origin of the Ascomycetes from the red

algae. The trichogyne is a prominent structure which cannot be

disregarded, especially as it falls in line with the structures found

in the Plorideae. The multicelled ascogonium, the trichogyne and

the ascogenous hyphae and asci have no homologue in the lower

fungi. We do find some ground for comparison when we study the

structure of the sex organs in the Plorideae.

It has not been possible to follow the method of nuclear divi¬

sion or the development of the asci. Perhaps the interpretation of

the processes observed can best be made from the assumption that

the sex organs are functional, and that the nuclei pair in the as¬

cogonium and then migrate into the aseus where fusion occurs. In

some respects part of the sexual process is analogous to certain

phases occurring in the Basidiomycetes where nuclear fusion is

delayed. No evidence of a true sporophytic stage could be found

in Venturia. Harper (1905) suggests that we consider the tetra-

sporange of the red algae as a progenitor of the ascus, the car-

pospores not being comparable to any stage in the Ascomycetes.

In case the so-called nuclear fusion, which, according to some

336 Wisconsin Academy of Sciences, Arts, and Letters.

workers on Ascomycetes occurs in the ascogonium, is found to

be only pairing of nuclei, then new conceptions may be developed.

The problem would be greatly simplified if the complex stages

due to triple fusion were eliminated. Another helpful method

which may aid in unraveling the nuclear history is the study of

new forms. The list of new Wisconsin fungi compiled by the

efforts and researches of Dr. J. J. Davis (1916-1921) continues to

grow, and some of these forms may supply the missing link. At

present we know so little about the effects of a physiological change

caused by the environment on the morphology and sexual activity

of organisms, and whether or not changes in the environment are

immediately reflected in the offspring by morphological changes,

that all speculations are futile. Until our conceptions rest upon

more secure grounds, homologies and similarities may only lead

us into scholastic discussions and far from the real nature of the

underlying causes that find expression in these complex and truly

marvelous forms of life.

It gives me great pleasure to express my gratitude and apprecia¬

tion to Dr. E. M. Gilbert for many helpful suggestions and for the

constructive criticism he has rendered during the course of this

investigation.

Literature

Bacliman, F. M. A new type of spermogonium and fertilization in Collema.

Ann. Bot. 26; 747-60 (1912).

- The origin and development of the apothecium in Collema pulposum

(Bernh) Ach. Archiv. f. Zellforschung 10: 369-430 (1913).

Barker, B. T. P. Sexual spore formation in the Ascomycetes. Ann. Bot.

15: 759 (1901).

- Morphology and development of the ascocarp of Monascus. Ann. Bot.

17: 167 (1903).

Bary, A. de. Tiber die Fruchtentwicklung der Ascomyceten. Leipzig. 1863.

- Morphologie und Biologie der Pilze. Leipzig. 1884.

Barrett, J. T. Development and sexuality of some species of Olpidiopsis

(Cornu) Fischer Ann. Bot. 26: 209-35 (1912).

Baur, B. Zur Frage nach der Sexualitat des Collemaceen. Ber. Deut. Bot.

Gesell. 16: 363-67 (1898).

- IJntersuchungen iiber die Entwicklungsgeschichte der Flechten

apothecien. Bot. Zeit. 62 : 21-44 (1904).

- Die Anlage und Entwicklung einiger Flechten apothecium. Flora 88:

319-32 (1901).

Bensaude, Mathilde. Eecherches sur le cycle evolutif et la sexualite chez

les Basidiomycetes. Paris, 1918.

Frey — Physiology of Yenturia Inequalis.

337

- Sur la sexualite chez les Champignons Basidiomycetes. Compt. Rend.

Acad. Sci. Paris 165: 286 (1917).

Blackman, V. H., and Welsford, E. J. The development of the perithecium

of Polystigma ruhrum D. C. Ann. Bot. 26: 761-67 (1912).

- On the fertilization, alternation of generation, and general cytology

of the Uredineae. Ann. Bot. 18: 323-73 (1904).

Blackman, V. H., and Fraser, H. C. L. Further studies on the sexuality of

the Uredineae. Ann. Bot. 20: 35-48 (1906).

Brefeld, O. Untersuchungen aus dem Gesammtgebiete der Mykologie. Botan*

ische Untersuchungen iiber Schimmelpilze. II. Die Entwicklungsge-

schichte von Pennicillium. Leipzig, 1874. Die Hemiasci und die As-

comyceten, IX Heft 1891, pp. 1-118; 340-51.

Brooks, F. T. The development of Gnomonia erythrostoma Pers. Ann. Bot.

24: 585-605 (1910).

Brown, H. B. Studies in the development of Xylaria. Ann. Myc. 11: 1-13

(1913).

Brown, W. H. The development of the ascocarp of Lachnea scutellata. Bot.

Gaz. 52: 273-305 (1911).

- The development of Pyronema confluens. Amer. Jour. Bot. 2: 289-97

(1915).

Bucholtz, F. Beitrage zur Kenntnis der Gattung Endogone, Link. Beihefte

Bot. Centralbl. 292: 147-225 (1912).

Bulliard, P. Histoire des Champignons de la France. 1791.

Carruthers, D. Contributions to the cytology of Eelvella crispa, Ann. Bot.

25: 243 (1911).

Christman, A. H. Sexual reproduction in rusts. Bot. Gaz. 39: 207 (1905).

- Morphology of Rusts. Bot. Gaz. 44: 81 (1907).

Claussen, P. Zur Entwicklungsgeschichte der Ascomyeeten Boudiera. Bot.

Zeitg. 63: 1-28 (1905).

- Zur Kenntnis der Kernverhaltnisse von Pyronema confluens, Ber.

deut. Bot. Gesell. 25: 586-90 (1907).

- Tiber Entwicklung und Befruchtung bei Saprolegnia monoica. Ber.

deut. Bot. Gesell, 26b: 144 (1908).

- Zur Entwicklungsgeschichte der Ascomyeeten Pyronema confluens.

Zeit. Bot. 4: 1-64 (1912).

Clinton, G. P, Apple Scab. III. Agr. Exp. Sta. Bull. 67: 109 (1901).

- Reproduction des Aseomycetes. Ann. d. Sci. Nat., Serie 6, T. 3. 1876.

Cutting, E. M. On the sexuality and development of the ascocarp in

Ascophanus carneus Pers. Ann. Bot. 17: 399-417 (1909).

Dale, E. Observations on the Gymnoascaceae. Ann. Bot. 17: 591-96 (1903).

■ - On the Morphology and Cytology of Aspergillus repens. Ann. Myc.

7: 215-25 (1909).

Dangeard, P. A. La reproduction sexuelle des Aseomycetes. Le Botaniste

4: 21. 1894-95.

■ - Recherches sur le developpement du Perithece chez les Aseomycetes.

Le Botaniste 10: 1-385. 1907.

- Recherches sur la reproduction sexuelle des Champignons. Le Bot¬

aniste 3: 222-278 (1893).

338 Wisconsin Academy of Sciences, Arts, and Letters.

■ — : - La reproduction sexuelle des Ustilaginees. Compt. Eend. I’Aead. des

Sci. Oct. 1893.

- Etude sur le developpement et la structure des organismes superieurs.

Le Botaniste II serie, 1910.

Davis, B. M. The fertilization of Batrochospermum. Ann. Bot. 10: 49-76

(1896).

Davis, J. J. Notes on parasitic fungi in Wisconsin. Trans. Wis, Acad. Sci.

20: 389-413 (1921).

Dodge, B. O. Methods of culture and the morphology of the archicarp in

certain species of the Ascobolaceae. Bull. Torr. Bot. Club 39: 139-97

(1912).

- The morphological relationships of the Elorideae and the Ascomycetes.

Bull. Torr. Bot. Club 41: 157-202 (1914).

Fanil, J. H. The Cytology of the Laboulbeniales. Ann. Bot. 25: 649. 1911.

Fitzpatrick, H. M. The life history and parasitism of Eocronartium muscicola.

Phytopath. 8: 197-218 (1918).

- The cytology of Eocronartium muscicola. Amer. Jour. Bot. 5: 397-499

(1918).

Gilbert, E. M. Studies on the Tremellineae of Wisconsin. Trans. Wis.

Acad. Sci. 16: 1137-70 (1910).

- Cytological studies of the lower Basidiomycetes. Trans. Wis. Acad.

Sci. 20: 387-9 (1921).

Guilliermond, M. A. Le developpement et la phylogenie des Levures. Eev.

gen. des Sciences du 15 Eaut. 1911.

- Contribution a 1 ’etude cytologique des Ascomycetes. Compt. Eend.

de I’Acad. des Sci. 137: 938-39, 1088 (1903).

- Zygosaccharomyces Nadsonii, Nouvelle especes de levures a conjugaison

heterogamique. Annales de I’Institut Pasteur 34: Fasicule. 1918.

- Sur la division nucleaire des levures. Annales de I’Institut Pasteur

31: No. 3, Mars. 1917.

- Sur la karyokinesic de Peziza rutilaus. Compt. Eend. de 1. Soc. Biol.

56, 412 (1904).

- Les progres de la cytologie des Champignons Prog. Eei. Bot. 4: 389-

542 (1911).

Harper, R. A. Beitrage zur Kentniss der Kernteilung und Sporen Bildung

in Ascus. Ber. dent. Bot. Gesell. 13: 67 (1895).

- Die entwicklung des Peritheciums bei SpJiaerotJieca co^tagnei. Ber.

dent. Bot. Gesell. 13: 475-81 (1895).

• - ^tiber das Verhalten des Kerne bei der Fruchtentwicklung einiger

Ascomyceten. Jahrb. f. Wiss. Bot. 29: 656 (1896).

• - Kernteilung u. freie Zellbildung im Ascus. Jahrb. f. Wiss. Bot. 30:

1-49 (1897).

- Nuclear phenomena in certain stages in the development of the

smuts. Trans. Wis. Acad. Sci. 12: 475 (1899).

- Cell division in Sporangia and Asci. Ann. Bot. 13: 47-525 (1899).

- Sexual reproduction in Pyronema confluens and the morphology of

the Ascocarp. Ann. Bot. 14: 321-400 (1900).

- Sexual reproduction and the organization of the nucleus in certain

mildews. Publ. Carnegie Inst., Washington, D. C. 37: 1-92 (1905).

Frey — Physiology of Venturia Ineqiialis.

339

- Nuclear phenomena of sexual reproduction in fungi. Anier. Nat. 44:

533-46 (1910).

- Binucleate cells in certain Hymenomycetes. Bot. Gaz. 33: 1-25

(1902).

iCkeno, S. Tiber die Sporenbildung und systematische Stellung von Monascus

purpureus. Ber. dent. Bot. Gesell. 21: 259 (1904).

Janczenski, E. Eecherches morphologique sur I’Ascobolus furfuraceus. Ann,

des Sci. Nat. 5 E Serie 15: 197 (1892).

Juell, O. H. tiber Zellinhalt, Befruchtung und Sporenbildung bei Dipodascus.

Flora 91: 47 (1902).

Jones, F. E. Is the Venturia of the Wisconsin wild crab, Fyrus coronarius

identical with the Venturia inaequalis of the cultivated apple? Thesis,

MS. Univ. of Wis. 1914.

Bohlman, O. Zuz Entwicklungsgeschichte der Ascomyceten. Acta Soc. Sci.

Fennicae 13: pt. 43, 1883.

Killian, K. Morphologic, Biologie und Entwicklungsgeschichte von Cryp-

tomyces pteridis (Eebent.) Eehm. Zeit. f. Bot. 10, 48-126 (1918).

- — liber die Entwicklung der Perithecia bei Venturia inaequalis (Cooke)

Ad. Ber. dent. Bot. Gesell. Heft. 1: 164-68 (1915).

Kunkel, O. The production of a promycelium by the aecidiospores of

Caeoma nitens Burrill. Bull. Torr. Bot. Club 40: 361-66 (1913).

Kiilin, J. G. 1858. History of plant pathology by G. Whetzel. 1918.

Levine, M. Studies in the cytology of the Hymenomycetes especially the

Boleti. Bull. Torr. Bot. Club. 40: 137-81 (1913).

Maire, R. La formation des asques chez les Pezizes et devolution nueleaire

des Ascomycetes. Compt. Eend. Soc. Biol. 55: 1401 (1903).

- — Sur la cytologie des Hymenomycetes. Compt. Eend. Acad. Sci. Paris

131: 121-4 (1900).

- Sur les divisiones nucleaires dans I’Asque de la Morille et de quelques

autres Ascomycetes. Compt. Eend. Soc. Biol. 56: 822 (1904).

- Eecherches cytologiques sur le Galactinia succosa. Compt. Eend. de

PAcademie der Sci. Paris 137: 769-71 (1903).

- Eecherches cytologiques sur quelques Ascomycetes. Ann. Mycologici.

3: 123. 1905.

• - Eemarques sur la cytologie de qualque Ascomycetes. Compt. Eend.

Soc. Biol. 56: 86 (1904).

McCubbin, W. A. Development of the Helvellineae. Bot. Gaz. 49: 195-

205 (1910).

Miyake, K. The fertilization of Pythium de Baryanum. Ann. Bot. 15:

653-67 (1901).

Nichols, M. A. Morphology and development of certain pyrenomycetous

fungi. Bot. Gaz. 22: 301-28 (1896).

Nichols, S. P. The nature and origin of the binucleated cells in some

Basidiomycetes. Trans. Wis. Acad. Sci. 15: 30-70 (1905).

Nienburg, W. Beitrage zur Entwicklungsgeschichte einiger Flechtenapothe-

cien. Flora 98: 1-40 (1907).

- Entwicklungsgeschichte von Polystigma rubrum DC. Zeit. f. Bot. 6:

369-400 (1914).

340 Wisconsin Academy of Sciences, Arts, and Letters.

Olive, E. W. The morphology of Monascus purpureus. Bot. Gaz. 39: 56-60

(1905).

- Sexual cell fusions and vegetative nuclear divisions in the rusts. Ann.

Bot. 22; 331-57 (1908).

- Nuclear conditions in certain short cycled rusts. Science N. S. 33:

194 (1911).

Oltmans, F. tiber die Sexualitat der Pilze. Biol. Centr. 21: 437-42 (1901).

Overton, J. B. The morphology of the ascocarp and spore formation in the

many-spored asci of ThecotJieus pelletiera. Bot. Gaz. 42: 450 (1906).

Rainlow, G. Zur Entwicklungsgeschichte von Theleholus stercoreus. Bot.

Zeitg. 64: 85-99 (1906).

- Beitrage zur Entwicklungsgeschichte der Ascobolen. Mycologisches

Centrallblatt 5; Heft 4, Nov., 1919.

Rawitscher, Felix. Beitrage zur Kenntnis der Ustilagineen. Zeit. f. Bot. 4:

674 (1912).

Sachs, J. Lehrbuch der Botanik. 1-632. Leipzig, 1868.

Schikorra, W. Uber die Entwicklungsgeschichte von Monascus. Zeit. f.

Bot. 1; 397 (1909).

Sutherland, G. K. Additional notes on marine Pyrenomycetes. New. Phy-

tologist 13-14: 183-93 (1914).

Stevens, F. L. The compound oosphere of Albugo bliti. Bot. Gaz. 28: 225

(1899).

Trow, A. H. Karyology of Saprolegnia. Ann. Bot. 9: 609 (1895).

- Cytology of Pythium ultimum. Ann. Bot. 15: 267. 1901.

- - On fertilization in the Saprolegniae. Ann. Bot. 18: 541 (1904).

Tulasne, R. Selecta Fungorum Carpologia. 3; 197 (1865).

- Note sur les phenomenes de copulation que presentent quelques cham¬

pignons. Ann. des. Sci. Nat. Bot. 6: 211 (1866).

Van Tieghem, P. Traite De Botanique. 2: 1129-56 (1891).

Wager, H. Sexuality of fungi. Ann. Bot. 13: 29-55; 575-97 (1899).

- Fertilization of Peronospora parasitica. Ann. Bot. 14: 263 (1900).

Wager, H., and Peniston, A. Cytological observations of the yeast plant.

Ann. Bot. 24: 45 (1910).

Welsford, E .J., and Blackman, V. H. The development of Polystigma

rubrum DC. Ann. Bot. 26; 761-67 (1912).

Woronin, M. Zur Entwicklungsgeschichte des Ascobolus pulcTierinum und

einiger Pezizen. De Bary and Waronin Beitrage Morph, u. Physiol, des

Pilze 2: 1-11 (1866).

Yamonouchl, S. The life history of Polysiphonia. Bot. Gaz. 42: 401-49

(1906).

Explanation of Figures

All figures were made with the aid of a camera lucida. A Zeiss number 12

ocular and number 2.3 mm. apochromatic oil immersion objective were used

for figures 7-18. The remaining figures were made with a Leitz 1/^0 oil im¬

mersion and number 10 ocular.

Frey — Physiology of Venturia Inequalis,

341

Plates X and XI

Pig. 1. Beginnnig of coil which results in a perithecium.

Fig. la. Shows conjugation of adjacent filaments.

Pig. 2. Formation of coil.

Fig. 3. Coil enlarging; enveloping hyphae being formed.

Fig. 4. Early stages in coil formation. Observed in Van Tieghem cell.

Pig. 5. Large branch which forms the coiled ascogonium has bulb and club

shaped hyphae applied to it.

Fig. 6. Section of a coil. The nuclei are enlarged.

Pig. 7. Club shaped cells, the antheridia, applied to the trichogyne. A pore

is evident in the trichogyne.

Fig. 8. The ascogonium has become septate and consists of several cells, one

of which is shown. The nuclei migrate down the trichogyne singly and

later pair in the ascogonium.

Pig. 9. Section of a perithecium showing the coil after fertilization has oc¬

curred.

Pig. 10. Section of young coil.

Fig. 11a. Section of perithecium after fertilization showing trichogyne and

coiled ascogonium. The nuclei are paired in the cells of the ascogonium

but not in the trichogyne.

Pig. 11b. Another section of perithecium shown in Fig. 11a. The re¬

mainder of the coil is evident.

Fig. 12. Section of a perithecium. The ascogonium has several cells, each

with paired nuclei. The nuclei of the trichogyne are not paired. The

pore through which the male nuclei entered the trichogyne is visible.

Pig. 13. Section of a perithecium showing paired nuclei in the cells of the

ascogonium.

Fig. 14. Section of young perithecium. The ascogonium is multicelled as

in the previous figures. The trichogyne was not found.

Pig. 15. Section of a young perithecium at the time of fertilization. The

nuclei are paired but the ascogonium has not become septate.

Pig. 16. Section of a perithecium. The paired nuclei appear to be fusing.

The trichogyne is disintegrating. The cells of the ascogonium appear to

be branching in preparation for ascus formation.

Fig. 17. Section of perithecium shown in Fig. 16. Note paired nuclei in the

cells which later form asci.

Fig. 18. Section of perithecium. Branching and budding of the ascogenous

cells is taking place.

Fig. 19. The coil of the ascogonium is reorganizing. New cells are being

formed either by budding or by cell division. Nuclear division is taking

place.

Fig. 20. In place of the coil of the ascogonium there are a large number of

cells. From these compact cells branches arise which later form asci.

Pig. 21. Cells in which nuclear fusion seems to be taking place.

Fig. 22. Section of perithecium showing formation of many small cells, which

later develop asci, from the coil which originally consisted of five or

six cells.

Pig. 23 a, b, c. Similar to Fig. 22.

Pig. 24. Section of mature perithecium. Asci are being formed.

Pig. 25 a, b. Section of perithecium showing asci.

342 Wisconsin Academy of Sciences, Arts, and Letters.

TRANS. WIS. ACAD., VOL XXI

PLATE X

Frey- — Physiology of Venturia Inequalis.

343

TEANS. WIS. ACAD., VOL. XXI

CYTOLOGICAL STUDIES OF TAPHRINA CORYLI NISHIDA

ON CORYLUS AMERICANA

Ella May Martin

Introduction

The cytological work of Dangeard (1894) on Exoascus defor¬

mans, of Ikeno (1901) on various Taphrina forms, and of still

later workers on other Ascomycetes, left unsettled various ques¬

tions concerning the uninucleate or multinucleate condition of the

cells, of the origin of the nuclei which fuse in the ascogenous cell,

nuclear divisions in the ascus, and the formation of the ascospores.

It was with the hope of throwing light upon some of these prob¬

lems in Taphrina coryli Nishida, that the work described in the

present paper was undertaken.

Materials and Methods

Leaves and twigs of Corylus americana infected with Taphrina

coryli Nishida were collected by Dr. J. J. Davis and Dr. E. M.

Gilbert in the vicinity of Madison, Wisconsin, in the spring of the

years 1917, 1920, and 1921, and by the writer in the same location

in 1920 and 1921. During the spring of 1921, material was fixed

at various hours of the day and night in order that the condition

of the nucleus and cell at different times might be studied. The

parts collected were immediately dipped in Carnoy^s fluid for a

few seconds to remove the air, and were then placed in vials con¬

taining the desired fixative. A number of fixing reagents were

used, but the best results were obtained with MerkeUs fixative and

with Flemming’s weak chrom-osmic-acetic acid solution.

Spores were also germinated and kept growing for several days

in sterile aqueous solutions made from hazel leaves. Germinating

spores were fixed in Merkel’s solution and attached to slides by

stippling them into a thin film of egg albumen on the slides. These

were then dried and stained. Imbedded material was cut from

346 Wisconsin Academy of Sciences, Arts, and Letters.

three to seven microns in thickness. Flemming’s triple stain was

used for all preliminary studies, but all results with it were con¬

firmed by the use of Heidenhain ’s iron-alum haematoxylin.

Observations

These studies show that the mycelium of Taphrina coryli Nishida

may infect all parts of the leaf and the cortex of the youngest

twigs of Corylus americana, as an intercellular parasite. It is

found between the cells of the cortex of a one-year-old twig, but it

is not found in older branches; apparently it grows out into the

new cortex each season. In a young diseased leaf, the fungus may

be traced along the veins, between spongy parenchyma and palisade

cells, and frequently a layer of short thick cells is found between

the cutin and the epidermis. In an older, badly diseased leaf,

every available intercellular space is occupied by the fungal

hyphae, and in addition, portions of the upper and lower surfaces

of the leaf may be covered with asci. Instead of the classification

of the mycelium given by Pierce (1900) for Exoascus deformans,

namely vegative, distributive, and fruiting hyphae, the writer will

refer to only vegetative and fruiting hyphae; meaning, by the

latter, asci and subcuticular ascogenous cells, and by the former,

all other mycelia belonging to the fungus.

The vegetative cells of Taphrina coryli form long filaments, par¬

ticularly where they follow the vessels in the leaves and between

the palisade cells. In the air spaces between the spongy paren-«

chyma cells, they broaden out and frequently branch. This dicho¬

tomous branching has been described by Pierce for Exoascus de¬

formans, and is quite characteristic of Taphrina coryli (fig. 1).

Each vegetative cell is long, cylindrical, thin-walled and contains

a fine granular network of cytoplasm with either one or, more fre¬

quently, two nuclei (fig. 2). In a binucleate cell, the nuclei are

small, each containing one prominent nucleole and a very small

quantity of chromatin (fig. 3). This binucleate condition arises in

the vegetative cells, and never in the ascogenous cells, of Taphrina

coryli Nishida. Division figures were seen a number of times in

the vegetative cells (figs. 4, 5). The spindles and centrosomes are

extremely small, as are also the masses of chromatin material which

pass to each pole. Ikeno observed binucleate cells in all forms of

Taphrina studied by him, and Dangeard (1894) held that the

mycelial cells of Exoascus deformans were originally binucleate

Martin — Taphrina Coryli Nishida on Corylus.

347

cells. Miss Adams (1915) and Miss Bitner (1915) found the

binucleate condition only in ascogenous cells, where they found

evidences that pairing of cells had taken place. It seems very prob¬

able, however, from my observations, that the binucleate condition

which is later observed in the ascogenous cells has arisen in the

vegetative cells (fig. 6, 7), although by what process, I am unable

to determine.

As the fungus travels outward between cells of the palisade layer,

the hyphae appear similar to those previously described, but the

cells are usually longer and thinner. But as the mycelium passes

between the epidermal cells, it becomes more difficult to determine

the character of the vegetative cells, for they are only rarely seen

and those observed become very slender as they force their way

between the epidermal cells. Slender vegetative hyphae are also

found in the cortex of the one-year-old twigs.

The Ascogenous Cells and Nuclear Fusion

By the time the fungus has reached the region between the epi¬

dermal cells and the cutin of the leaf, its cells have become much

thickened and, being packed together, have become cuboidal in

form. At times these fruiting or ascogenous cells are found in

layers two or three cells thick, but as the outer layer develops,

the cuticle is ruptured, the fungal cells increase in length and form

asci. In the ascogenous cells, the cytoplasm is often more vacuolate

than that of the vegetative cells, though this condition may vary.

The binucleate condition which arose in the vegetative cells appar¬

ently does not continue long in the ascogenous cells, for most of

these cells have only one large, fusion nucleus.

Nuclear fusion is found quite often in ascogenous cells of ma¬

terial collected at any time of day, but in material collected early

in the morning the nuclei in the majority of ascogenous cells are

fusing. It may be that moisture and low temperature stimulate

the activity of the cell in such a way as to favor nuclear fusion.

The two nuclei approach each other (figs. 6, 7), finally lying side

by side in the cell (fig. 8). Their nucleoles (one in each nucleus)

are at first some distance apart, but the thickened chromatin masses

lie close together. As the two nuclei come into contact with each

other, the nuclear membrane breaks down between them, so that

there is formed a large double nucleus (figs. 9, 10, 11). The two

chromatic systems remain separate for a time as described by

348 Wisconsin Academy of Sciences y Arts, and Letters.

Harper for Phyllactinia corylea (1905), but eventually the elong¬

ated nucleus rounds up and assumes the form of a typical resting

nucleus (figs. 12, 13). The two nucleoles gradually approach each

other and unite, the large nucleole formed by the fusion staining

deeply with safranin, but containing an unstained or lightly

stained area at the center. The chromatic system becomes more

clearly defined and is now a coarse network with thickened granules

at intervals. Both the linin and the chromatin granules stain

deeply with gentian violet. That this fusion may be a case of

endokaryogamy similar to that found in the Basidiomycetes seems

probable. By passing through a number of mitoses, in the vegeta¬

tive cells, it may be that the nuclei which finally fuse have lost

their originally similar characteristics.

Division of the Fusion Nucleus

As the resting nucleus prepares for division, a spirem thread

is gradually evolved from the reticulum of linin and chromatin,

and very soon evidence of the pairing of the threads is apparent

(fig. 14). They lie closely twisted about each other, and, since

this synaptic stage is so clear, this division may be considered the

heterotypic division. The condition of synapsis and that of the

longitudinally split spirem (fig. 15) are frequently found, which

fact may be an indication that the nucleus passes through these

phases slowly. The nucleole, during this time, changes in staining

properties, the outer zone taking much less stain than the inner.

Another stage, showing the spirem thrown into loops which group

at the center and spread out to the periphery of the nucleus, is fre¬

quently found (figs. 16, 17). Fitzpatrick (1918) suggested that

these loops represent chromosomes in Eocronartium muscicola, but

this is probably not the case in Taphrina coryli Nishida, for here

five or six (the number of loops observed) does not represent

either the haploid or the diploid number of chromosomes. The

nucleus elongates considerably (fig. 20), and soon a definite spindle

is noted which is larger than that found in the vegetative cells,

but it, also, has a granular body or center at each end. This en¬

tire spindle is likewise surrounded by a clear area. It is im¬

possible to make out the individual chromosomes; probably each

mass of chromatin represents a number of chromosomes (figs. 21,

22, 23, 24).

These chromatin masses are in such position as to suggest the

equatorial plate stage most frequently, though they appear at

Martin — Taphrina Coryli Nishida on Corylus,

349

other times to be in late metaphases and anaphases. The nueleole

disappears at the time of the spindle formation, but no indications

of the division of a central body to form two daughter centers, as

described by Harper (1899, 1905), are seen. The reorganized

nuclei are typical resting nuclei, each containing one nueleole

(fig. 25).

The Formation of the Basal Cell

Each mature ascus of Taphrina coryli Nishida has a basal cell,

but the time of the formation of the septum varies. Often before

the division of the fusion nucleus is complete, the plasma mem¬

brane in the lower part of the ascogenous cell begins to constrict

(fig. 23). Again, the plasma membrane may not begin to constrict

until after the daughter nuclei are fully formed. It may be that

vacuolar membranes also play a small part in the formation of the

cell plate, for vacuoles are frequently in such a position that vacu¬

olar membranes meet the constricting plasma membrane (fig. 25).

Many cases were observed, however, which suggest that the con¬

stricting portions of the plasma membrane have stretched out to

meet each other, forming a cleavage furrow in which will be laid

down the cell wall (figs. 26, 27). From a study of a number of

such cells, it seems safe to conclude that the process of separation

of the ascus cell proper from the basal cell is initiated either before

or just after the first nuclear division is complete. One of the

nuclei formed by the division of the fusion nucleus passes to the

upper part of the ascus where it later undergoes division, the re¬

peated divisions of its daughter nuclei forming the nuclei for the

eight primary spores. The other nucleus formed from the fusion

nucleus remains in the lower part of the ascus, w^hich becomes the

basal cell. The nucleus of this cell persists for a while and then

disintegrates ; the only remains of it being in the form of chromatin

granules which are left in the cytoplasm for a time (fig. 28). As

the ascus grows older, even the chromatin granules disappear, and

the cytoplasm, which has been quite dense, now becomes more and

more vacuolate (fig. 29), until it, too, completely disappears in the

basal cells of the older asci (figs. 31, 33). These would conform

to the empty basal cells figured by Atkinson (1894) for Exoasctis

deformans, Exoascus cerasi, and Exoascus decipiens. Sadebeck

(1893) in Taphrina Sadeheckii Johns, and Miss Bitner (1915)

in Taphrina coryli also found empty basal cells with the older asci.

350 Wisconsin Academy of Sciences, Arts, and Letters.

The Formation op the Ascospores

As the ascHS elongates, nuclear division proceeds within it. The

second mitosis is probably homoetypic and the next two vegetative

divisions, so that there are produced eventually eight spores for

the eight nuclei. Since there was only one nuclear fusion, there is

necessary only one reduction of chromatin material. Examples of

the vegetative divisions in process are few, but those observed show

the typical spindle with masses of chromatin passing toward the

poles. Three divisions of the ascus nucleus have been mentioned

for some of the Exoascaceae.

Ikeno (1903) in Taphrina cerasi, and Dangeard (1894) in Ex-

oascus deformans, mention three divisions. This would be probable

in types that have no basal cell, but where there is such a cell con¬

taining at some time a nucleus, it is evident that there must be four

nuclear divisions: a division of the fusion nucleus and three suc¬

cessive divisions of the primary ascus nucleus and its derivatives.

Nuclear division is seen more frequently in material collected in

the early morning. Ikeno (1901) mentions many division figures

at the time of the development of the spores of Taphrina Johan-

sonii.

The nuclei formed are small and are often found in a resting

condition. They contain a deeply staining nucleole and chromatin

arranged upon a linin network (figs. 28, 29, 30). In a mature

spore the nucleus occupies about one-third or one-fourth of its

volume, and in the formation of the spores dense masses of cyto¬

plasm collect about each of the eight nuclei. Sometimes this cyto¬

plasm takes up so much stain that the nucleus can scarcely be seen

through it (fig. 35). The plasma membrane of a spore seems to

be formed by the union of membranes of vacuoles; this plasma

membrane secretes a thick, clear wall, darkly stained only at in¬

tervals (figs. 31, 32). Vacuoles are especially conspicuous at the

time of spore formation and are in such a position that each spore

is entirely encircled by them, the inner vacuolar membranes uniting

to form the boundary of the spore. This would conform to the

description given by Brown (1911) for the delimiting of the spores

of Lachnea scntellata. Faul (1912) considers this progressive

method of delimiting the spore the characteristic method for As-

comycetes. He expresses the idea that astral rays would not

naturally come together, but would rather tend to diverge. Fraser

and Brooks (1909) report that vacuoles play an important part in

Martin — Taphrina Coryli Nishida on Corylus.

351

the formation of the spores of Ascoholus furfuraceus. No indica¬

tion of astral rays as described by Harper (1899) in Lachnea

scutellata and (1905) in Erisyphe cichorearum, Erisyphe com¬

munis, and Phyllactinia corylea have been found in Taphrina

coryli. If present they may have been such fine threads as to be

invisible with the magnification used. There are, however, many

suggestions that vacuolar membranes are the active agents in form¬

ing membranes of spores (figs. 31, 32, 33, 34).

The Germination of the Ascospores

Nishida (1912) ascribed to Taphrina coryli an irregular number

of spores, and such is the case when each of the original eight spores

comes to maturity and begins to form conidia. This process might

be called fission, for, unlike the budding of yeasts, the spore length¬

ens and divides into conidia of equal size (fig. 37). When the

spores begin to produce conidia, they become somewhat irregular

in form, the constriction often being present earlier on one side of

the spore than on the other (fig. 35). There must be a softening

of the cell wall, for a spore lengthens and, as the cell wall and

plasma membrane constrict, the whole assumes the form of a dumb¬

bell (fig. 36, A-G). At this time the nucleus begins to stretch out

(fig. 36, G) and eventually divides, thus providing a new nucleus

for each of the conidia so formed. The behavior of the nucleus was

observed in the stained material, but the living spores were ex¬

amined under the miscroscope for the rate of conidia formation.

It was found that spores germinate to form conidia rapidly after

they have been subjected to chilling.

Spores were placed in a hanging-drop of sterile solution made

from hazel leaves, and the hanging-drop preparations were then

placed in sterile Petri dishes. These were left in the refrigerator

at a temperature of 10° C. for twelve hours. When taken out in

the morning and placed in a room at about 20° C., germination took

place almost immediately. Figure 37 shows what occurred in the

course of one-half hour. A spore increased in size (compare figs.

37A and 37B) in the first ten minutes; in the next five minutes the

stages represented in figure 37C, D, and E were passed through;

and in another five minutes the constriction is complete between

the two daughter conidia (fig. 37F). Within a few minutes one

conidium may lengthen and furrow in on one side, and soon a

secondary conidium is partly formed (fig. 37G). All the spores

352 Wisconsin Academy of Sciences, Arts, and Letters.

do not germinate at once to form conidia, bnt one or two may

lag behind and show no indication of germinating even after some

of the others have produced secondary conidia (fig. 38).

The secondary conidia are very small, and it is these small conidia

that produce the vegetative mycelium. The attempt to grow

mycelium on artificial media was not successful. Short hyphae

were produced but soon died. These hyphae were slender, having

a diameter less than the diameter of the conidium and containing

a very vacuolate cytoplasm (fig. 39).

Summary

1. TapJirina coryli Nishida is an intercellular parasite infecting

all parts of the leaves and the cortex of the one-year-old twigs of

Corylus americana.

2. The vegetative cells form long filaments and frequently

branch dichotomously. They serve to distribute the fungus

through the leaf and through the twig.

3. The binucleate condition arises in the vegetative cells, and

not in the ascogenous cells.

4. The division of the vegetative nucleus is mitotic, with small

spindles and centrosomes. It is difficult to recognize individual

chromosomes in the chromatin masses.

5. The binucleate condition continues into the ascogenous cells,

and then nuclear fusion takes place. Nuclear fusion takes place

most frequently early in the morning.

6. The fusion nucleus is a typical resting nucleus, containing

a large nucleole, chromatin on a linin network, and a central mass

of deeply staining chromatin.

7. The division of the fusion nucleus is a heterotypic reduction

division, as is indicated by the occurrence of synapsis and of a

later stage showing the spirem thrown into a definite, looplike ar¬

rangement from the center to the periphery.

8. Granular bodies are present at the ends of the spindle and a

clear area surrounds the entire spindle. It is not definite how

these granular bodies are formed.

9. The separation of the basal cell from the ascus proper takes

place by the constriction of the plasma membrane.

10. One nucleus is left in the basal cell; the sister nucleus

passes into the ascus proper. Gradually the former nucleus dis¬

integrates, but the second and its derivatives divide to form nuclei

for the eight primary spores.

Martin — Taphrina Coryli Nishida on Corylus.

353

11. Vacuolar membranes appear to be the active agents in the

formation of the spores.

12. Conidia are produced by the lengthening of the spore,

followed by its constriction into two cells of unequal size, the

smaller growing to equal the larger before separation. The pro¬

duction of conidia is stimulated by chilling.

13. Primary conidia produce secondary conidia which germ¬

inate on artificial media to form short hyphae.

I wish to express my thanks to Professor E. M. Gilbert, who

suggested this work, for his assistance and encouragement.

Literature Cited

Adams, R. M. 1915. Studies in the life history of Exoascus communis on

Prunus cuneata. 1-35. B. A. Thesis, Univ. of Wis. '

Atkinson, G. F. 1894. Leaf curl and plum pockets. Cornell Agr. Exp.

Sta. Bull. 73: 319-355.

Bitner, A. L. 1915. Studies on the life history of Taphrina coryli Nishida

on Corylus americana. 1-39. B. A. Thesis, Univ. of Wis.

Brown, W. H. .1911. The development of the asocarp of Lachnea scutellata.

Bot. Gaz. 52: 275-303.

Dangeard, P. A. 1894. La reproduction sexualle des Ascomycetes. Le

Botaniste 4: 30-35.

Fanl, J. H. 1912. The cytology of Lahoulbenia chaetophora and L. gyrini-

darum. Ann. Bot. 26: 325-355.

Fisch, C. 1885. Ueber die Pilzgattung Ascomyces. Bot. Zeit. 43: 33-39

and 49-59.

Fitzpatrick, H. M. 1918. The cytology of Eocronartium muscicola. Am.

Jour. Bot. 5: 397-419.

Fraser, H. C. I., and Brooks St. John. 1909. Studies in the cytology of

the ascus. Ann. Bot. 23: 537-547.

Harper, B. A. 1899. Cell division in sporangia and asci. Ann. Bot. 13:

467-525.

- 1905. Sexual reproduction and the organization of the nucleus in

certain mildeTvs. Carnegie Inst. Pub. 37: 1-97.

Ikeno, S. 1901. Studies on the spore formation in Taphrina Johansoni Sad.

Flora 88: 229-236.

- 1903. Sporenbildung von Taphrina Arten. Flora 92: 1-31.

Nishida Y. 1912. On Taphrina coryli Nishida n. sp. Miyake Festschrift,

206, Tokyo.

Pierce, N. B. 1900. Peach leaf curl. Its nature and treatment. U. S. Dept.

Agr. Div. Veg. Phys. & Path. Bull. 20: 11-45.

Sadebeck, R. 1893. Die Parasitischen Exoasceen, eine Monographic. Jahrb.

Hamb. Wiss. Bot. 10: 1-110, also Just. Bot. Jahrb. 21 Jahrg. I Abth.

196-199.

354 Wisconsin Academy of Sciences^ Arts, and Letters.

Explanation of Figures

All drawings were made with a camera lucida at table level. Figures 3, 4,

5y 10, 12, 15, 23 were drawn with a Zeiss 2 mm. apochromatic oil immersion

objective and 12 ocular (X2800) ; fig. 13 with Zeiss 2 mm. oil immersion

objective and 18 ocular (X3600) ; figs. 1, 2, 8, 9, 14, 16-22, 24-29, 35, 38

with Zeiss 3 mm. oil immersion objective and 12 ocular (X1650). A Leitz

oil immersion objective %0 and 4 ocular (X1600) was used with drawings

6, 7, 34, 36, 38, and the others, figs. 30-33, 37, 39, were drawn with a Leitz

7 objective and ocular 5 (X665).

Plates XII and XIII

Fig. 1. Dichotomous branching of vegetative hyphae of TrapMna coryli

Nishida.

Figs. 2, 3. Uninucleate and binucleate conditions shown in cross and

longitudinal sections of vegetative cells.

Figs. 4, 5. Mitosis in vegetative cells.

Figs. 6, 7. Two nuclei are present in young aseogenous or fruiting cells.

Figs. 8, 9, 10, 11. Stages in the fusion of nuclei in aseogenous cells.

Figs. 12, 13. Typical fusion nucleus in resting condition.

Figs. 14-24. Division of the fusion nucleus: fig. 14, synapsis; fig. 15,

longitudinally split spirem; figs. 16, 17, spirem in loops; figs. 18, 19, 20,

thickened spirem; figs. 21, 22, equatorial plate; fig. 23, metaphase and con¬

striction of plasma membrane; fig. 24, anaphase.

Fig. 25. Eeorganized nuclei; one, the primary ascus nucleus, and the

other, the nucleus of the basal cell.

Figs. 26, 27. Further constriction of the plasma membrane to separate

basal cell from ascus proper.

Fig. 28. Binucleate ascus and disintegration of the nucleus in the basal cell.

Fig. 29. Second nuclear division of the ascus.

Figs. 30-34. Asci containing spores. Large vacuoles surround the spores:

fig. 32 represents the ascus in cross section.

Fig. 35. Early stage in formation of conidia.

Fig. 36, A, B, C, D, E, F, G. Successive stages in the budding of the

primary ascospore showing behavior of nuclei and constriction of cell walls.

Fig. 37. A, B, C, D, E, F, G. Successive stages in the formation of pri¬

mary and secondary conidia. Stages A-G represent development taking place

in thirty minutes. Drawn as observed in living material grown on artificial

media made from hazel leaves.

Fig. 38. Conidia and ascospore showing that all the ascospores do not di¬

vide simultaneously.

Fig. 39. Germinating conidia drawn from living material growing on the

same media as the spores shown in fig. 37.

Martin — Taphrina Coryli Nishida on Corylus,

355

TEANS. WIS. ACAD., VOL XXI

PLATE Xn

356

Wisconsin Academy of Sciences, Arts, and Letters.

TEANS. WIS. ACAD., VOL. XXI

PLATE XIII

THE STRUCTURE AND BEHAVIOR OF THE NUCLEUS IN

THE LIFE HISTORY OF PHYCOMYCES NITENS

(AOARDH) KUNZE AND RHIZOPUS

NIGRICANS EHRBG

E. A. Baird

Among the contributions to various phases of the life history

of the Mucoraceae, a number of the writers do not include any

clear statement of the structure of the nucleus or how it behaves

in division. Most of the workers, who have attempted to follow

closely the behavior and structure of the nucleus in zygospore

formation, have described the resting nucleus in the forms studied

as consisting of a nucleole, a nucleoplasm containing little or no

stainable substance, and a nuclear membrane. In the consideration

of nuclear division they have generally described a type of mitosis

with a spindle formation, but in the details of this process the

descriptions are incomplete. Many of these have described numer¬

ous nuclear fusions during the maturation of the zygospore together

with a disorganization of other nuclei that do not fuse.

Dangeard and Leger (1894a) in a cytologieal study of zygospore

formation in Sporodinia grandis and two species of Mucor have

described the structure of the nucleus in the vegetative hyphae and

the sexual branches of these forms as vesicular, consisting of a

surrounding membrane and a centrally placed nucleole, which

are separated by a non-staining ‘‘cytoplasm” containing a little

chromatin. In the young zygospores of these forms they find two

sizes of nuclei.

In another report, Dangeard and Leger (1894b) describe with¬

out figure the structure of the nuclei in Mucor mucedo and M.

racemosus as similar to that described above. In the ripe zygo¬

spore of Sporodinia the small and, later, the large nuclei disappear.

After their disappearance the zygospore contains a varying num¬

ber of deep-staining bodies.

Leger (1895a) figures the nucleus of the vegetative mycelium

as a vesicular body with a centrally placed nucleole. He describes

358 Wisconsin Academy of Sciences, Arts, and Letters.

karyokinetic figures as occurring in the vegetative hyphae but does

not figure nor describe the process of nuclear division. He finds

that the nuclei, evident up to this time, begin to disintegrate as

the zygospore matures and its protoplasm becomes spongy and

filled with oil. This disintegration is a process in which the

nucleole becomes smaller and finally disappears. The nucleus

then becomes vacuole-like. A few nuclei, which are two or three

times larger than those entering the zygospore at the time the

gametes fuse, remain in the zygospores longer than do the other

nuclei, but ultimately disappear. In another paper, without figures,

Leger (1895b) has reported on the conditions he has found in

species of Pilobolus, Rhizopus, Chaetocladium and Mortierella,

together with those forms previously studied with Dangeard. He

describes the structure of the nucleus in the vegetative hyphae

in all the forms as similar to that described by Dangeard and

himself. He indicates a point of interest as to the fate of the

nucleus in the mycelium and the columella in that the nuclei

become reduced to nucleoles which persist after all other traces of

the protoplasm have disappeared.

Istvanffi (1895) describes the nucleus in Mucor sp. He finds

that the nuclei are scattered thruout the entire protoplasm; that

they are elliptical or spherical in shape and usually provided with

a nucleole. In spore formation only one nucleus enters each spore.

During preparation for germination, the spore is observed to

have eight to ten nuclei. In the tips of hyphae where he considers

that the nuclei are youngest, he states that they may consist of only

a small homogeneously-stained body, not exceeding Ifi in diameter.

Harper (1899) in his description of spore formation of Pilo¬

bolus states that the vegetative nuclei of this form divide in the

basal bulb, thus giving rise to the nuclei that enter the sporange

during its development. He also finds that the nuclei divide in

the protospore and in the swelling spore in preparation for germi¬

nation. In none of these references to nuclear division does he

describe the structure of the nucleus or indicate how it divides.

However, he figures several nuclei, each consisting of a dense

central body enclosed by a granular nucleoplasm and a nuclear

membrane. He also illustrates several division figures, each con¬

sisting of an elongated body, at the two poles of which are several

dense granules; the two groups of granules are connected by one

or more fibrous strands.

Baird — History of Phycomyces Nitens (Agardh) Kunze. 359

Swingle (1903) in a report of spore formation of Phycomyces

and Ehizopus describes the nuclei in the sporange of either form

as in a resting condition. In either form each nucleus is ap¬

proximately spherical in shape, and consists of one or two nucleoli,

a finely granular chromatin, and a surrounding membrane. In the

columella of Ehizopus he describes the nuclei as disintegrating.

According to him the process of disintegration consists of the ap¬

pearance of a red-stained mass on one side of the nucleus, followed

by the nucleus taking on the appearance of a shrunken homogene¬

ous mass often irregular in shape and staining much as do the

crystalloids of the protoplasm. In Phycomyces the structure of

the nucleus is similar except that it may have as many as three

nucleoli, and in this form he has described the disintegration of

the nuclei in the mycelium as in the columella of Ehizopus.

Gruber (1901) in a description of nuclear behavior in zygospore

production of Sporodinia grandis agrees with Leger up to the

point at which the protoplasm of the two gametes become mixed.

From this point on he does not find nuclei of two sizes, nor fusions

taking place between paired nuclei, nor evidence that nuclei are

disintegrating, but rather that the nuclei are clearly in evidence

after fourteen days of development of the zygospore. After five

or six weeks and again at the end of six months he still finds nuclei

in the same condition as when the zygospore was formed.

In a later paper, reporting on the sexual process of Zygorhyn-

chus Moelleri, Gruber (1912) describes nuclear fusions as occur¬

ring between the nuclei of the male gamete and an equal number

of those of the female gamete.

Dangeard (1903) describes the nuclear structure and behavior

in a species of Mucor and in Sporodinia. In either of these forms

a nucleus of the gametes consists of a small nucleole and a homo¬

geneous achromatic nucleoplasm. After the fusion of the gametes

the nuclei divide one or more times, then nuclear fusions occur.

A daughter nucleus, thus arising in the zygospore, consists of a

nucleole, a network of granular chromatin, and a nuclear mem¬

brane. In the process of nuclear fusion, the membranes fuse at

the point of contact. At first the two nucleoles rest within the

membrane thus formed and then fuse. Some of the nuclei do not

fuse but disintegrate. In rather mature zygospores, this author

finds large deep-staining bodies throughout the protoplasm, which

he suggests have arisen from mucorine crystals.

360 Wisconsin Academy of Sciences, Arts, and Letters.

Moreau (1911a) and 1911b) in describing cytological studies,

especially in zygospore formation, of species of Mucor, Zygorhyn-

chus, Circinella, Rhizopus and Sporodinia states that a nucleus of

these mucors consists of a chromatin nucleole, a nucleoplasm, and

a nuclear membrane. The nucleole, is either centrally, eccen¬

trically, or laterally placed. In some cases in Mucor a centrosome,

chromatic in nature, is observed on the external surface of the

nuclear membrane. In the columella of Rhizopus he finds a

modification of the nucleus in that there is no nuclear membrane

and the nucleus consists simply of a homogeneous body. According

to Moreau, the nucleus divides mitotically in the vegetative hyphae

and in the zygospores of the forms studied. The process of mitosis

is inaugurated by the disappearance of the nuclear membrane and

of the nucleole. No other stages characteristic of prophases are

described. He describes an equatorial plate stage in which double

chromosomes are borne on a straight spindle, terminated at each

pole by a centrosome. He describes and figures a later stage in

which two daughter chromosomes are in process of moving toward

each pole. Stages in the reorganization of the daughter nuclei

are not described. In the zygospore, the mitotic nuclear divisions

take place, according to Moreau, as if activated by the mixing

of the protoplasm from the two fusing gametes. He also describes

a form of amitotic nuclear division of the homogeneous nuclei

in the columella of Rhizopus. In Mucor, following the nuclear

divisions in the young zygospore, the nuclei fuse in pairs, giving

rise to a large number of fusion nuclei. A number of nuclei fail

to fuse and later disintegrate. In Zygorhynchus the nuclear dis¬

integration takes place before fusion, and in the zygospores of this

form, only two fusion nuclei are formed. The fusion nuclei of all

the forms studied by Moreau are similar in structure. Each con¬

tains a single large nucleole. He states that the fusion nuclei per¬

sist in the mature zygospore and form the basis of the nuclei of the

thallus arising from the germination of the zygospore.

Moreau (1913) has published the results of very extensive cyto¬

logical research of a large number of the Mucoraceae. These later

results are in accord with his previous work. In the case of

Phy corny ces he states that he saw clearly stages in the fusion of the

nuclei in the young zygospore at the time the spiny exospore was

being formed. The fusion of the nuclei in the zygospore, of those

forms for which he describes the process, consists in the fusion of

the two membranes at the point of contact, thus forming one

Baird — History of Phycomyces Nitens (Agardh) Kunze. 361

nuclear cavity. At first the two nucleoles lie separate, but later

fuse. In his description of nuclear division in this paper, he con¬

forms to his previous report.

Miss Keene (1914) in her cytological studies of Sporodinia

grandis finds that the nucleus in this form is granular in structure

and contains a centrally placed nucleole which she considers

chromatin in nature. She believes that nuclear divisions occur in

the tips of the two sexual branches. As the mixing of the two

fusing gametes takes place, nuclear fusions occur. Miss Keene

describes no divisions in the zygospore preceding nuclear fusions,

as was described by Dangeard and Moreau. She describes nuclear

degeneration of unfused nuclei in the zygospore, but her descrip¬

tion of the process is somewhat different from that given by other

workers. According to her, the process is first accompanied by an

enlargment of the nucleole which does not stain as deeply as in

preceding conditions. The nucleus eventually becomes a homo¬

geneous-staining mass. According to her the fusion nuclei are in

evidence in zygospores two and three months old.

Later, Miss Keene (1915) has contributed results of cytological

studies of Phycomyces nitens. She has figured the nuclei in ger¬

minating asexual spores as containing one or two deep-staining

bodies. The resting nucleus is described by her as bounded by a

membrane and containing a deep-staining body, probably the

nucleole, and chromatin granules throughout the nuclear cavity.

She states that in the germinating spores, the young sporange, sus-

pensors, and progametes, the nuclei show conditions that are very

suggestive of division figures. She suggests that in nuclear divi¬

sion figures containing three bodies, two may be chromosomes and

the third the nucleole. In the young zygospore she finds that many

of the nuclei are arranged in groups of twelve to sixteen. Some

of these nuclei fuse in pairs ; others do not fuse. Later the unfused

nuclei of the zygospore coalesce to form one or two large amorphous

masses that persist in the zygospore several months old. Similar

masses are formed within the suspensors. The fused nuclei persist

in the zygospore and are confined to a thin peripheral zone of the

cytoplasm.

Burgeff (1915) has contributed a very interesting description,

without figures, of his studies of the cytology of Phycomyces nitens

and mutants of this species. His observations as to the structure

of the nucleus or as to the facts of nuclear fusion are not fully in

accord with those of either Keene or Moreau. He describes the

362 Wisconsin Academy of Sciences, Arts, and Letters.

nucleus of the vegetative mycelium as consisting of a very small

homogeneous body, and suggests that it is identical with a chromo¬

some. Division of a nucleus in the vegetative hyphae consists

simply in the separation of the chromatin body into two daughter

nuclei. He finds that in spore formation in the sporange from five

to twelve of these chromatin bodies are enclosed within each spore.

He describes no change in the structure of the nucleus during zygo¬

spore formation, except that after the fifth day of development the

nucleus is spongy and contains a single chromatin body. He re¬

ports no nuclear divisions or fusion during the formation and

maturation of the zygospore.

The nuclei in the zygospore preparing to germinate are sur¬

rounded by a membrane but have only one chromatin body and by

the time the germ tube pushes out, eleven to twelve days after sow¬

ing, the nuclei are more or less irregular in outline and provided

with several chromatin bodies and a nucleole. At this stage they

are several times larger than the nuclei of the sexual generation.

In the germ tube and sporange, nuclear divisions occur among the

large nuclei. Burgeff states that all stages in mitosis are difficult

to make out. Clear prophases occur in which the nucleole disap¬

pears. The chromatin is separated into twenty -four (estimated)

chromosomes. Twelve chromosomes move to each pole within the

membrane; no equatorial plate stage is observed. Burgeff char¬

acterizes this as a heterotypic division, following which, he states,

a homotypic division occurs in which distinct chromosomes are

present. Following the homotypic division successive divisions

take place, giving rise to a large number of membraneless nuclei.

These nuclei, with a number of nuclei surrounded by membranes,

become the nuclei of the spores formed within the germ-sporange.

He suggests that the nuclei with membranes are either unfused

nuclei of the gametes or else nuclei that have not passed through

reduction division. In spore formation only one nucleus enters into

the formation of a spore. After spore formation, as they mature,

the nucleus of each spore divides successively so that each mature

spore contains a number of nuclei. In his summary, however,

Burgeff states that he has not observed the division of nuclei with

membranes within the spores of the germ-sporange.

Burgeff (1920) describes the nuclei of the parasitic mould,

Chaetocladium, and of its host, Mucor, as being very similar. The

nucleus of either form consists of a deep-staining body with or

without a clear zone surrounding it. The clear zone is more fre-

Baird — History of Phy corny ces Nitens (Agardh) Kunze. 363

quently observed in poorly nourished hyphae. Burgeff calls the

deep-staining body chromatin, states that it is nucleole like, and

that it is frequently located eccentrically in the nucleus. In well-

nourished hyphae the chromatin body is much larger than in poorly

nourished hyphae. His special study of the nuclei has been in the

galls formed at the point of attack of the parasite. He points out

that the nuclei in the sporangiophores contain a much larger

chromatin body than do those in the vegetative branches ; he

attributes this to the presence of reserve food in the sporangio¬

phores. He states that the Mucor nuclei in the gall divide mitoti-

cally, but gives no figures nor details of the process. He also de¬

scribes crystalloid bodies as arising from degenerating nuclei.

Materials and Methods

The material from which the preparations were made was grown

in petri dishes on potato-glucose agar prepared with distilled water.

Flemming’s medium fixing reagent was used almost exclusively,

after a wide variety of reagents was tried out. The imbedded ma¬

terial was cut in sections 4/x-12/x thick and the preparations stained

with Flemming’s triple stain. Heidenhain’s iron-alum-haematoxy-

lin method of staining was also used, but in no case gave the

minute differentiation obtained by the use of the triple-stain.

For the germination of the zygospores of Phycomyces, zygospores

that had remained on the original substratum, upon which they

grew, in a dark room at an average temperature of 20 degrees C.,

germinated readily when transferred to a non-nutrient 2% agar

medium. The zygospores germinated, forming a single germ-

sporangium within four or five days after transferring. The cul¬

tures of germinating zygospores were grown in a light room, at

room temperature. On account of interruption in the work, the

study of nuclear behavior in the germ sporange and of the sex of

the germ-sporangiospores has not been completed.

Phycomyces Nitens

In the swollen spores of Phycomyces about to germinate there

are one or more vacuoles with several (4-13) nuclei imbedded

within the cytoplasm. After the germ tube has been formed

(fig. 1) the nuclei are distributed throughout the enlarged cell

by the increase of vacuolar volume. The nuclei show no change

in structure up to this period. So far as I have observed the

364 Wisconsin Academy of Sciences, Arts, and Letters.

nuclei show no change in structure until some time after the

germination of the spore. As soon as the first hypha has several

branches, one observes the first changes in the nuclei as de¬

scribed below.

A nucleus in the resting spore and during early stages after

germination consists only of a homogeneous, deep-staining, ap¬

proximately spherical body which I consider to be chromatin in

nature. Its surface is undoubtedly membranaceous, although

no membrane is differentiated. With the triple stain this body

stains deeply with the safranin while the surrounding cyptoplasm

stains orange. The first change that occurs in such a nucleus is

the swelling of the chromatin body accompanied by the formation

of a vacuole within it. In some instances the chromatin becomes

distributed for a time in the peripheral zone. With further growth

of the vacuole the chromatin becomes separated at one or more

points. If separated at one point, the chromatin may present a

somewhat crescent-shaped optical section (fig. 2f). On the other

hand, it may be separated so as to present a somewhat horseshoe¬

shaped section with a chromatin body in the opening of the horse¬

shoe (fig. 2b). Usually the chromatin substance is separated into

from three to six bodies, as the growth of the vacuole progresses

(figs. 2a, 2c, and 2d). Figure 2 illustrates the distribution of such

nuclei associated with vacuoles from a hyphae in the substratum of

a three-day-old culture.

The question naturally arises in this connection concerning the

nature of the membrane surrounding the nuclear vacuole. The

vacuole evidently originates below the surface of the chromatin.

But as soon as the chromatin is separated sufficiently at any point

a thin membrane remains surrounding the vacuole ; the membrane

does not stain with the safranin as does the chromatin but stains

more as does the surrounding cytoplasm. This may be due to the

fact that it is so thin as not to appear differentiated from the sur¬

rounding cyptoplasm, but I am inclined to believe that, while the

membrane originates within the chromatin, its composition be¬

comes changed and so differentiated from the chromatin.

The above-described vesicular body has undoubtedly been inter¬

preted by most workers as the nucleus. Since, as I shall point out

later, such a vesicular body functions as a single structure of the

protoplasm at certain stages in the life history of the plant, per¬

haps it would not be incorrect to consider it a nucleus. However,

on account of its behavior in connection with division and distribu-

Baird — History of Phycomyces Nitens (Agardh) Kunze. 365

tion of the chromatin in rapidly growing mycelium, I consider the

vesicle a vacuole, and each portion of chromatin at its periphery a

nucleus.

The nuclei that have been thus formed are destined soon to be¬

come disassociated from one another and in subsequent nuclear

division give rise to daughter nuclei by a similar process.

There appear to be two different forces that bring about the ulti¬

mate separation of the daughter nuclei. One is the growth of the

vacuole from within as just described. The other force is the

streaming of the protoplasm within the hyphae, giving rise to elon¬

gated division figures and most frequently observed in the hyphae

of the substratum where their tubular form is not uniform in out¬

line and very angular in contour. In such hyphae of living cul¬

tures I have frequently observed that the rate of streaming is not

uniform throughout a given diameter. This character of the

streaming evidently accounts for the fact that the vesicular divi¬

sion figures are sometimes elongated and often curved in the

hyphae of the substratum (figs. 2c, 2d).

Where the enlargement of the vacuole alone is operating, the

daughter chromatin masses and the vacuole with which they are

associated present a spherical figure (figs. 2a, 2e). While the out¬

line of the vacuole is still unobliterated, one often finds the daugh¬

ter nuclei beginning a subsequent division (fig. 2e). In this figure

the daughter nuclei have lagged behind the progress of the vacuolar

membrane and appear within the space bounded by the membrane

that was carrying them apart. Here, too, the original vacuolar

membrane is losing its identity and becoming thickened and evi¬

dently is about to be incorporated into the slimy portion of the

cytoplasm. It is also observed that slimy cytoplasm is being

formed within the cavity of the vacuole between the daughter

nuclei.

In the sporangiophore and sporange one does not usually find

division figures of the nucleus elongated or otherwise distorted,

for evidently the protoplasm in which the nuclei are imbedded is

moving uniformly or is at rest, as is the condition of the protoplasm

of the sporange and sporangiophore as the sporange reaches ma¬

turity. In these structures, therefore, one usually finds the nuclei

associated with approximately spherical vacuoles (figs. 3a and 3b).

Figures of similar form are also found in the gametes, suspensors,

and immature zygospores.

J

366 Wisconsin Academy of Sciences^ Arts^ and Letters.

In some instances a vacuole associated with a nucleus may arise

without bringing about a division of the nucleus. In such a case

the vacuole originated below the surface of the chromatin mass

and gradually enlarges eccentrically, giving rise to a vescicular

structure consisting of a vacuole with a single chromatin body at

its periphery (fig. 3a). In this case the vacuole is not destined to

play a part in nuclear division. In some cases the vacuole en¬

larges uniformly around the chromatin body, leaving the chromatin

suspended within the vacuole (figs. 4f, 4g and 5d). Thus is ap¬

pears that numerous vacuoles of the cytoplasm arise from the

chromatin bodies ; they may or may not have divided nuclei as they

were formed.

In figures 4a~4k are represented a number of nuclei taken from

the two gametes of a sexual apparatus. All show stages in nuclear

division except figure 4d where the vacuole was formed at one side

as described above. Figures 4c, 4f and 4g represent the type of

structure in which the vacuole was formed by progressing on all

radii of the sphere, leaving the chromatin body within the vacuolar

sap. A nucleus thus suspended within the vacuolar sap forms a

second vacuole within itself to bring about division (figs. 4f

and 4g).

During the process of nuclear division strands of protoplasm

are frequently observed, extending across the vacuole from one

chromatin body to another (fig. 4a). They vary considerably in

different figures and do not always appear. Such strands are evi¬

dently formed by the chromatin. In some cases the strands con¬

necting the daughter chromatin bodies appear to be so closely asso¬

ciated with the vacuolar membrane as to cause the membrane to

be somewhat flattened, to conform to the straight lines of the con¬

necting strands, and thus present a vacuole with a broken surface

of flat faces rather than the curved surfaces of other vacuoles

(figs. 4f, 4h and 4i).

In many cases the ultimate destiny of a vacuole of a division

figure is that it' becomes obliterated by an incorporation of the

membrane with the slimy cytoplasm, and the space within becomes

gradually filled with slimy cytoplasm, either reticulate or homo¬

geneous. Figures 4j and 4k represent such groups of nuclei after

the vacuole has thus become obliterated. The group of seven nuclei

represented in figure 4k evidently originated from two sister nuclei.

The three in the lower portion of the group and to the left are

Baird — History of Phy corny ces Nitens (Agardh) Kunze. 367

evidently from one of the nuclei, and in this group each has begun

to form a vacuole.

After the fusion of the gametes and during the growth of the re¬

sulting zygospore, I find no evidence of a pairing or fusing of the

nuclei. During the growth of the zygospore, nuclear division

occurs continuously until the protoplasm finally takes on its resting

condition as described below. All of the nuclear division figures

of the zygospore are of the spherical type. The only observed

change in the protoplasm after the fusion of the gametes is the

increased affinity of the cytoplasm for the safranin and gentian

stains. As a result of this change in staining reaction, the differ¬

entiation of the nuclei is not as definite in the zygote as in the sus-

pensors or the vegetative hyphae. The nuclear division figures of

the zygospore are of the same form as has been described for them

in the sporange (figs. 5a-5e). Figure 5 represents a zygospore

during the growing period while the exospore is being formed and

shows the fragments of the walls of the gametes which fused to

form the first wall of the zygospore.

During the growing period of the zygospore many of the divid¬

ing nuclei are found in groups of four to eight. In figure 5a six

dividing nuclei of such a group, at the same optical level, are

shown in their relative positions. Evidently these have originated

from the same nucleus through two successive divisions, but, since

the vacuoles bringing about the divisions have enlarged but little,

the daughter nuclei of each division remain in close proximity to

each other. On the other hand, the isolation of other daughter

nuclei in the zygospore is accounted for by the greater expansion

of the vacuole separating the nuclei. Figure 5b contains both

isolated and grouped nuclei.

The nuclei of a zygospore which has attained nearly its full size

are found associated with smaller vacuoles than are the nuclei of

a zygospore during the rapidly growing period, or of the vegetative

hyphae. Compare figures 5c and 5e of a full-grown zygospore with

figures 3a and 3b from a sporange, and with figures 6a and 6b from

a growing zygospore.

In the stained preparation of the protoplasm I have found no

evidence of fat, nor structures identified with its formation. How¬

ever, a test with Sudan III of protoplasm crushed out' of zygospores

in all stages of development shows very clearly the presence of fat.

Undoubtedly the fat in the zygospores occupies many of the larger

spaces that appear as vacuoles in the preparations. The fat is

368 Wisconsin Academy of Sciences, Arts, and Letters,

evidently dissolved by xylol or chloroform during the clearing and

imbedding processes.

A mature zygospore two weeks old shows very little of the slimy

portion of the cytoplasm. Figure 7 shows the condition of the

protoplasm in a zygospore from an eleven-day-old culture, in which

the zygospores would vary in age from three to seven days. At

this stage one finds that the protoplasm consists of three distinct

portions, vacuoles, the slimy portion of the cytoplasm, and nuclei.

The vacuoles evidently represent spaces filled with oil, cell sap or

gas. Some of the vacuoles contain a slightly-staining granular

substance, as has frequently been described by other workers, but

this is not so specially characteristic of the vacuoles of the zygo¬

spore as it is of the vacuoles of the vegetative hyphae. The slimy

portion of the cytoplasm is still in evidence but in a much reduced

proportion. Most of the nuclei together with the vacuoles asso¬

ciated with them present a different aspect than has been described

for them heretofore in the life history of the plant. Some of the

vacuoles associated with the nuclei appear as has been previously

described; the others now contain a substance that stains with the

same reaction as do the chromatin bodies at the periphery of the

vacuoles. It appears that the chromatin substance has increased

in volume accompanied by a reduction in the proportion of the

cytoplasm.

In figure 7a a distinct vacuole is present, but the surrounding

membrane has become thickened and stains deeper. Figure 7b

illustrates a nuclear vacuole that has become filled with a stainable

substance. Figure 7d represents the final stage in the develop¬

ment of such a nuclear structure in the mature zygospore, the form

in which most of the protoplasm, except the fat, is found. These

structures are often so closely crowded together that their sur¬

faces are compressed flat against each other (fig. 7d). During this

change the same stain-absorbing elements as are contained in the

chromatin characterize all the material contained in the nuclear

vacuoles. I call these structures ‘^reserve food bodies”. Each

consists of a variable amount of food reserve, which occupies the

space of the nuclear vacuoles, and one or more chromatin masses

which may usually, though not always, be observed as deeper-

staining bodies at' the periphery (fig. 7b).

My interpretation is that during the process of maturation of the

zygospore there are no nuclear fusions. After the fusion of the

two gametes the nuclei continue to divide rapidly, as they do in

Baird — History of Phycomyces Nitens (Agardh) Kunze. 369

the actively growing vegetative mycelinnij until the zygospore is

mature. Then the nuclei^ acting as metabolic centers, proceed to

transform available food into a reserve form of protoplasm which

is stored in the space previously occupied by the vacuolar sap of

the associated vacuole. As for the fat, I am unable to conclude

whether it is a metabolic product of the cytoplasm or of the nuclei.

In preparations of zygospores in which the protoplasm is' in a

resting condition, the proportion of stainable substances and of the

spaces appearing as vacuoles varies considerably. This variation

is undoubtedly due to a difference in the amount of nutrition avail¬

able for the given zygospores. In some zygospores a comparatively

large proportion of the space may be unoccupied by the deeply-

stainable substances (fig. 7). In others the same substances occupy

a much greater proportion of the space within the zygospores. In

some cases this material is located largely in the peripheral zone,

in others distributed more or less irregularly throughout.

In crushing zygospores under water, bubbles always appear

from within the rupture of the inner, leathery spore wall. There¬

fore, I conclude that the clear spaces appearing in preparations

are in some instances occupied by gas resulting from metabolism.

The protoplasts of zygospores five months old are unchanged in

appearance from those that have matured up to the resting stage.

After the zygospores are placed on moist agar for germination, one

finds that many of the dense nuclear bodies become separated from

each other by vacuolated cytoplasm. This change is most evident

in the peripheral zone. In the more central region the nuclear

structures stick together in large masses. In the peripheral zone

there are many cases of a single nucleus with a vacuole or a group

of several such nuclei surrounded by a very dense mass of slimy

cytoplasm. The vacuoles associated with some of these nuclei have

lost a large portion of their deep-staining substance. Figure 8

illustrates a section of a germinated zygospore with a portion of

the germ tube. It contains many of the nuclear structures filled

with food reserve as has been described for the zygospores before

germination; many of them persist during zygospore germination

and the formation of the sporange. In the peripheral zone are

many of the dense cytoplasmic masses surrounding one or more

nuclei. Figure 8a shows such a cytoplasmic mass, wherein one

nuclear structure is still unchanged, and from at least two of the

nuclear vacuoles the reserve food has been dissolved. The later

condition of the nuclear vacuoles is clearly illustrated in figures

370 Wisconsin Academy of Sciences, Arts, and Letters.

8b, 8c and 8d. Such figures correspond very closely to those of

nuclei observed in the zygospore during maturation when the

vacuoles were beginning to receive the reserve substance.

An insufficient number of stages were fixed to warrant reporting

on the process of spore formation in the germ sporange. How¬

ever, it has been determined that nuclei of the type shown in fig¬

ures 8a, 8b, 8c, and 8d pass into the sporangiophore and are found

there before and after spore formation in the sporange. There

appear to be no nuclear divisions in the germinating zygospore or

in the germ sporangiophores either before or after spore formation.

Rhizopus Nigricans

A complete study of the nucleus of Bhizopus nigricans has been

made in the vegetative mycelium, in the sporanges, in the mature

and germinating spores, in the gametes, and in the zygospores dur¬

ing maturation. Since no germination of zygospores has been ob¬

tained, it has been impossible to follow the nuclear behavior

through that process.

It is very evident that nuclear behavior is, in general, the same

in Rhizopus as has been described for Phycomyces in the processes

of nuclear division and in the formation of reserve protoplasm

stored in the vacuoles associated with the nuclei of the zygospores.

The nuclear vacuoles do not become quite as large in Rhizopus

as they do in Phycomyces. Compare figures 9a, 9b, and 9c taken

from the sporange of Rhizopus with figures 3a and 3b taken from

the sporange of Phycomyces; both sporanges were at about the

same stage of development.

In Rhizopus the obliteration of the vacuole, causing the separa¬

tion of the chromatin masses, takes place sooner after the separa¬

tion of the daughter nuclei, and, therefore, in the case of Rhizopus

one finds more frequently the nuclei disassociated from a vacuole.

Figure 9e illustrates a case in which two sister nuclei evidently

were at first associated with the same vacuole and later one of them

formed a second vacuole, which separated the two chromatin

masses still further. A third small vacuole also has formed be¬

tween them. A somewhat similar case is illustrated in figure lib

in which a nucleus from a young zygospore has divided and one

of the daughter chromatin masses is forming a vacuole within an¬

other vacuole. Figures 10a, 10b, and 10c are dividing nuclei

taken from a vegetative hypha.

Baird — History of Phy corny ces Nitens (Agardh) Kunze. 371

Another minor variation in the process of nuclear division, oc¬

curring in Rhizopus as compared with Phycomyces, is that in

Rhizopus the chromatin mass is separated more frequently into

two portions rather than into three, four, or more as in Phycomyces

(figs. 10a, 10b, 10c). In a few cases observed, however, in Rhizopus

the chromatin may be separated into several portions (figs. 13a

and 13b).

Figures 11a and 11c illustrate cases in each of which the vacuolar

membrane has progressed further than the two daughter chromatin

bodies, thus leaving them within the vacuole, and figure 13b shows

one such chromatin body forming a second vacuole within itself.

Figures 13a, 13b, and 13c represent nuclei from a zygospore at a

time when the associated vacuoles are becoming filled with homo¬

geneous reserve substance, as was described for the nuclear vacuoles

of Phycomyces.

Figure 12 represents a spore from the sporange of Rhizopus be¬

fore the spores have shrunk and their walls have thickened. The

nuclear structures observed here apparently shrink as the spore

matures and shrinks. Each structure thus becomes one of the

dense nuclei observed in the mature spore and that appear as soon

as the spore germinates, as was figured for Phycomyces (fig. 1).

Nuclei of Aged Mycelium

The nuclei of the vegetative, aged mycelium behave very simi¬

larly to those in the maturing zygospores. Nuclear division con¬

tinues in the vegetative hyphae up to a time which I conclude is

determined by a change in water balance. For, at a time when no

further growth of mycelium occurs, its nuclear vacuoles take on

the appearance of those which contain reserve food in the zygo¬

spores (figs. 14a and 14b). In figure 14a, taken from a 36-day-old

culture of Phycomyces, the vacuole has become filled with a homo¬

geneously-staining substance, and some of the nuclei associated

with the vacuole are still partially differentiated. Figure 14b rep¬

resents a similar condition of a nuclear structure taken from a

vegetative hypha of a 14-day-old culture of Rhizopus.

As the mycelium becomes dried out in aging cultures, the slimy

cytoplasm does not disappear from the mycelium on a large scale

as it does in the zygospores. However, in the aged mycelium par¬

tially dried out, some of the hyphae appear empty, except for a

few of the deep-staining nuclear structures and a very slight

372 Wisconsin Academy of Sciences, Arts, and Letters.

amount of slimy cytoplasm; other hyphae contain a large amount

of slimy cytoplasm and nuclear structures filled with reserve

food (fig. 16).

Mycelium of Phycomyces forty days old was placed on moist

nutrient agar and after sixteen hours many hyphae had grown out

from the old mycelium. Preparations made from sections of this

material showed that nuclear structures of the type described

above for aged mycelium were abundant in the new hyphae in the

substratum. Some had lost completely the reserve food of the

vacuoles and the nuclei were in a state of active division. Most of

the nuclear structures were in a transitory condition in which the

vacuolar portions had swollen considerably; the reserve food was

partially dissolved and some of the nuclei were beginning division.

Figure 15a shows several nuclei in a very dense cytoplasm which

apparently has been formed from the reserve substance of the

vacuolar portion. Figure 15b shows a nuclear structure in which

most of the reserve food has been dissolved and the vacuole is quite

clear; one of the nuclei is in progress of division and a second

nucleus is apparently unchanged.

The foregoing nuclear behavior fits in well with the growth habits

of these plants. The plant grows rapidly with a favorable water

supply and during the growing period the nuclei divide rapidly

and are distributed into the newly formed portions of the plant.

As soon as a decrease in available water takes place or, at least,

after a certain minimum is reached, growth stops and at the same

time nuclear division ceases. As this change occurs most of the

nuclei are at a certain stage in division. The nuclear vacuoles

cease to expand but apparently proceed to absorb food reserve

from the surrounding cytoplasm, giving rise to nuclear structures,

filled with reserve food, as has been described above in old dried-

out mycelium. When such mycelium is again supplied with wa¬

ter, new hyphae are observed growing out from the aged hyphae.

The nuclei and associated structures resume activity, giving up the

food reserve and dividing as previously described.

Discussion

In the above report it has been difficult to use the term ^‘nucleus”

and convey the full idea that the writer holds as to its form and

structure. It is evident that the fundamental unit of structure

composing the nuclei is a small, homogeneous mass that stains

Baird — History of Phycomyces Nitens (Agardh) Kunze. 373

readily with safranin. This is undoubtedly the body seen and de¬

scribed by many writers as the nucleole. The vesicle that I have

called a vacuole associated with one or more chromatin bodies

seems to have been considered by others the nucleoplasm.

As I interpret the structure of the protoplasm of Phycomyces

and Rhizopus, the homogeneous body is nucleus ; it has the general

nature of chromatin in that it stains readily and is present in all

stages in the life history of these fungi.

Single masses of this chromatin may exist unassociated with

others, as in the spores, where the single body appears to have orig¬

inated from the reassembling of two or more small bodies that had

previously been formed by the division of a single chromatin body,

and in the vegetative hyphae, where individual bodies have be¬

come disassociated from others by the obliteration of a vacuole that

previously had been active in bringing about division. Most fre¬

quently several nuclei are associated with a vacuole and these

nuclei may or may not be connected by slender strands of proto¬

plasm other than the vacuolar membrane. A vacuole thus asso¬

ciated with one or several nuclei seems to have been formed within

the chromatin mass. Usually, the vacuole grows in such a way as

to rearrange the chromatin substance into several bodies around

its periphery. At other times, as it grows, the vacuole forms eccen¬

trically in the chromatin body, leaving most of the chromatin un¬

disturbed.

Although it has been observed that slimy cytoplasm seems to be

formed in the immediate vicinity of the nuclei, as in the germinat¬

ing zygospores and in the propagation of the mycelium from aged,

dried plants, and that at the time this cytoplasm is being formed

the nuclear vacuoles apparently lose their reserve, one can hardly

conclude whether the nuclei are active metabolic centers contribut¬

ing to this metabolism by forming cytoplasm from the reserve of

fat outside of the nuclear structures, or whether the food has been

stored in the nuclear vacuoles, and, later, cytoplasm formed di¬

rectly from it. Possibly both the reserve in the nuclear vacuoles

and the fat outside are used. But it appears evident that the nuclei

are very intimately associated with the processes that bring about

the renewed vegetative condition of the protoplasm.

The nuclei in the mature spores of the sporange are dense, homo¬

geneous bodies. When the spore is first formed in the sporange it

contains several nuclear vacuoles with two or more chromatin

bodies associated with each. The condition of the nuclei in the

374 Wisconsin Academy of Sciences, Arts, and Letters.

mature spore appears to come about through a reverse in the proc¬

ess of nuclear division brought about by the expansion of the

nuclear vacuoles. At about the time the spores are formed there

seems to be a change from expansion of their protoplasm to a con¬

traction, due to a loss of water. At the same time the nuclear

vacuoles shrink in size and the chromatin bodies of each nuclear

structure reassemble to form a single chromatin mass. They re¬

main in this condition until after spore germination, at which

time they resume division.

Although it is impossible to state definitely which structures de¬

scribed by other workers correspond to structures that I have de¬

scribed, it is quite plain that most workers have found many of the

same structures in different mucors.

The non-staining ‘‘cytoplasm’’ of Dangeard and Leger (1894)

is undoubtedly the vacuole that I have described as associated with

most nuclei. Leger (1895 a) believes that in aged hyphae the

nuclei becomes reduced to nucleoli and persist after the rest of the

protoplasm disappears. I find a similar condition in many of the

aged hyphae, considering that the nucleole described by Leger is

identical with what I have described as a nuclear structure, con¬

sisting of one or more nuclei associated with a vacuole which has

become filled with a reserve protoplasm.

Istvanffi (1895) describes the nuclei in the growing tips of the

mycelium as small homogeneous bodies. This conception of a

nucleus is what I hold for the nucleus throughout the life history

of the two forms I have studied.

I find no formations in the two forms I have studied that corre¬

spond to the dividing nuclei in the protospores of Pilobolus as fig¬

ured by Harper (1899), in which he interprets fibrous strands be¬

tween two nuclei as the remains of spindle fibers.

Swingle (1903) has described the nuclei of Khizopus and Phy-

comyces as having two or three nucleoli. These are undoubtedly

what I have described as the daughter chromatin bodies of a single

nucleus. His description of disintegrating nuclei in the columella

of Rhizopus and aged mycelium of Phycomyces corresponds very

closely to my description of the nuclear structures with reserve

material in aged mycelium, which later may become active upon

resumption of growth on the part of such mycelium. Other work¬

ers have frequently referred to similar nuclear disintegration. It

seems probable that the disintegrating nuclei in zygospores men-

Baird — History of Phy corny ces Nitens (Agardh) Kunze. 375

tioned by Miss Keene (1915) are the same as the reserve food

bodies of zygospores described by me.

As to the question of nuclear fusions in the zygospores of the

various forms of the Mucoraceae, a review of the various reports

shows that even in the same species there is little agreement as to

when fusion takes place or how many nuclei take part. Several,

as my review of the literature shows, do not find nuclear fusion at

all in the early formation of the zygospore. Burgefi (1915) de¬

scribes nuclear fusions as taking place at or shortly after zygo¬

spore germination. Many of the workers have described nuclear

fusions as taking place soon after fusion of the gametes. For this

period I have described rapid nuclear division.

Burgeff postulated a theory as a result of his work to account for

the effects brought about by some nuclei fusing and others not

fusing. If the nuclei do not fuse anywhere in the life history of

the plants, as my results seem to indicate, the behavior of the

nuclei in the germ sporange remains to be demonstrated before any

theory to explain the significance of sexual process in the

Mucoraceae can be advanced.

Summary

1. The nucleus in Phycomyces nitens and in Bhizopus nigricans

is a dense, homogeneous, chromatin body.

2. The nucleus divides by a method of fragmentation into sev¬

eral portions ; the complete separation of these fragments is brought

about by growth of a vacuole formed within the body of the mother

nucleus. In some instances the separation of the daughter nuclei

is aided by the streaming of the protoplasm.

4. Vacuoles frequently form eccentrically in a nucleus in such

a way as not to bring about nuclear division.

5. Nuclear division continues during the growth of the fungus

in the sexual generation. In the vegetative hyphae nuclear divi¬

sion is arrested by the discontinuance of water absorption by the

mycelium; in the spores of the sporange, by a reverse of water

absorption to water excretion ; and in the zygospores by maturation,

which is undoubtedly accompanied by cessation of water absorp¬

tion.

5. In zygospore formation no nuclear fusions seem to occur.

6. When nuclear divisions stops in the zygospore and in the

vegetative hyphae, reserve material is stored in the vacuoles asso¬

ciated with the nuclei.

376 Wisconsin Academy of Sciences, Arts, and Letters.

7. The nuclei in process of division in the embryonic sporangio-

spores reverse their progress and the portions reassemble to form

the nuclei of the mature spores.

8. Upon germination of the zygospores of Phycomyces nitens the

reserve material in the nuclear vacuoles contributes to the forma¬

tion of slimy cytoplasm. There is no evidence of nuclear division

in the germ tube or in the zygospore.

9. Upon the resumption of growth of mycelium, the reserve of

the nuclear vacuoles contributes to the formation of new cytoplasm,

as do the similar structures of the germinating zygospores.

Sincere thanks are due Prof. E. M. Gilbert for his stimulating

and encouraging interest and suggestions while this work was being

carried on.

Department of Botany,

University of Wisconsin

BIBLIOGEAPHY

Burgelf, H. (1915). Ilntersuchungeii iiber Variabilitat, Sexualitat und

Erblichkeit bei Phycomcyes nitens Kunze. Flora 108:353-448.

- (1920). XJeber den Parasitismus des Chaetoeladium und die hetero-

caryotishe Natur der von ibm auf Mucorineen erzeugten Gallon. Zeits-

chrift fur. Bot. 12: 1-35.

Dangeard, P.-A. (1903). Les ancetres des Champignons su6rieurs. Bot-

aniste 9: 227-252.

Dangeard, P.-A., and Leger, M. (1894 a). Eecherches sur la structure des

Muconin^es. Botantiste 4: 4-11.

- (1894 b). Eecherches sur la structure des Muconinees. Compt. Eend.

Acad. Sci. Paris 18: 430-432.

Gruber, E. (1901). Ueber das Verhalten der Zellkerne in den Zygosporen

van Sporodinia grandis Link. Ber. Deutsch. Bot. Ges. 19: 51-55.

- (1912). Einige Beobachtungen iiber den Befruchtungsvorgang bei

Zygorynchus Moelleri Vuill. Ber. Deutsch. Bot. Ges. 30: 126-133.

Haider, E. A, 1899). Cell division in sporangia and asci. ^ Annales Bot.

13: 490-503.

Istvanffi, Gy. (1895). Ueber die Eolle der Zellkerne bei der Entwickelung

der Pilze. Ber. Deutsch. Bot. Ges. 13: 452-467.

Keene, M. L. (1914). Cytological studies of the zgospores of Sporodinia

grandis. Annales Bot. 28: 455-470.

- (1915). Studies of zygospore formation in Phycomcyes nitens Kunze.

Trans. Wis. Acad. Sci., Arts, Letters 19: 1195-1220.

Leger, M. (1895 a). Structure et developpement de la zygospore du

Sporodinia grandis. Rev. Gen. Bot. 7: 481-496.

- (1895 b). Eecherches histologiques sur le developpement des Muco-

rinees. Compt. Eend. Acad. Sci. Paris 120: 647-649.

Baird — History of Phycomyces Nitens (Agardh) Kunze, 377

Morean, F. (1911 a). Premiere note sur les Mueorinees. Le noyau au

repos. Le noyau en division. Mitose et amitose. Bull. Soc. Mycol.

France 27: 204-210.

- (1911 b). Deuxieme note sur les Mueorinees. Fusions de noyaux et

degenerescence nucleaire dans la zygospore. Fusions de noyaux sans

signification sexuelle. Bull. Soc. Mycol. France 27: 334-341.

- (1913). Eecherclies sur la reproduction des Mueorinees et de quelques

autres Thallophytes. Botaniste 13: 1-136.

Swingle, D. B. (1903). Formation of spores in the sporanges of Mhizopus

nigricans and Phycomeyes nitens. Bull. U. S. Bur. Plant Ind. 37: 7-38.

EXPLANATION OF FIGUEES

All figures drawn with the aid of the camera lucida.

Figs. 1 — 10, 14a, 15a, 15b and 16 are from Phycomyces nitens.

Figs. 11-13 and 14b are from Ehisopus nigricans.

Magnification of figures 7 and 8, 90 diameters; figure 5, 186 diameters; fig¬

ures 2 and 16, 1050 diameters; figure 5b, 2100 diameters; and all other figures

3250 diameters.

Plates XIV and XV

Pig. 1. Germinated spore.

Pig. 2. Hypha from substratum, from culture 3 days old, grown from spore.

Figs. 2a-2f. Nuclei from hyphae similar to that in fig. 2.

Pigs. 3a and 3b. Nuclei from sporange before spore formation.

Pigs. 4a-4k. Nuclei from gametes showing various stages in nuclear division.

Pig. 5. Young zygospore during growing period, showing portion of a sus-

pensor to the right.

Pig. 5a. A group of nuclei from zygospore shown in fig. 5.

Fig. 5b. Detail of protoplasm from zygospore in fig. 5.

Figs. 5c and 5e. Dividing nuclei from zygospore in fig. 5.

Pig. 5d. Nucleus of same zygopore aroimd which a vacuole has formed.

Pigs. 6a and 6b. Nuclear division figures from a maturing zygospore and

with comparatively large nuclear vacuoles.

Fig. 7. Mature zygospore, about 5 days old. The numerous dense bodies are

the nuclear structures, referred to in text, in which the nuclear vacuoles are

filled with reserve material.

Figs. 7a-7d. Nuclear structures with reserve material.

Fig. 8. Germinated zygospore with dense cytoplasm around nuclear struc¬

tures in peripheral region at the right; nuclear structures with reserve still

unchanged in the central region. A small portion of germ tube appears.

Pig. 8a. Cytoplasmic mass formed around nuclear bodies.

Pigs. 8b-8d. Nuclear structures from which reserve material has been dis¬

solved.

Figs. 9a-9c. Nuclear division figures from sporange before spore forma¬

tion.

Figs. lOa-lOc. Nuclear division figures from vegetative hyphae.

Figs, lla-llc. Nuclear division figures from young zygospore.

378 Wisconsin Academy of Sciences^ Arts^ and Letters,

Fig. 12. Embryonie spore. Nuclear division arrested in progress at about

this stage.

Figs. 12a and 12b. Stages in the maturation of spores, and shrinking of

nuclear vacuoles.

Figs. 13a and 13b. Showing nuclear vacuoles becoming filled with reserve

material in a zygospore.

Fig. 13c. Four sister nuclei from zygospore; one is enlarged somewhat

by a vacuole within, and is becoming filled with reserve material.

Figs. 14a and 14b. Nuclear vacuoles filled with reserve material, from aged

hyphae.

Figs. 15a and 15b. Nuclei from hypha propogated from aged mycelium.

In 15a slimy cytoplasm is forming around the nuclear structure; in 15b the

reserve material has been dissolved.

Fig. 16. Aged hypha after 16 hours on culture medium, after its protoplasm

has renewed activity.

Baird — History of Phy corny ces Nitens (Agardh) Kunze. 379

TRANS. WIS. ACAD., VOL. XXI

PLATE XIV

380 Wisconsin Academy of Sciences, Arts, and Letters.

TRANS. WIS. ACAD., VOL. XXI

PLATE XV

#

%

Wk

IOC

o

Ub

lie

A QUANTITATIVE STUDY OF THE LARGER AQUATIC

PLANTS OF GREEN LAKE, WISCONSIN^

H. W. Rickett

Notes from the Biological Laboratory of the Wisconsin Geological and Nat¬

ural History Survey. XXII.

Introduction

In the summer of 1921 a survey was made of Green Lake, Wis¬

consin, with a view to the estimation of the amount of large aquatic

plants produced by this body of water. This report embodies the

results of the investigation. Besides the gross quantitative data,

the study also furnished information on the distribution of various

plants, their depth relations, and the vegetation of different parts

of the lake.

A similar report has already been made (3) on Lake Mendota.

The methods used and the nature of the information obtained in

the study presently to be described were in general the same as

those in the previous case.

The work was done for the Wisconsin Geological and Natural

History Survey, under the supervision of Professor Chancey

Juday. For assistance in identifying several of the plants col¬

lected, I am indebted to Dr. E. A. Baird, Dr. R. H. Denniston, Dr.

G. M. Smith, and Dr. G. E. Nichols. For much assistance in col¬

lecting, I wish to thank Professor Juday and Mr. L. E. Noland.

Description of Lake

Green Lake is a roughly oval body of water, seven and one-half

miles in length and from one to two miles broad. It is 66 meters

deep at its west end, — ^by far the greatest depth recorded for any

Wisconsin lake. The east end is somewhat shallower. Its waters

are cool, clear (compared, for instance, with those of Mendota),

and of the bright tint which gives the lake its name. The color of

^ This investigation was made in cooperation with the U. S. Bureau of Fisheries, and

the results are published with the permission of the Commissioner of Fisheries.

382 Wisconsin Academy of Sciences, Arts, and Letters.

the water is best seen on cloudy days ; but even at other times, when

most lakes reflect the blue of the sky, the surface of this lake is

often a bright green. The same color is very noticeable when one

is beneath the surface. The lake floor is in most places covered

with a fine marl.

These conditions afford an interesting comparison with those in

Lake Mendota. The waters of the latter are less transparent and

warmer than those of Green Lake, and the bottom is of mud.

There are striking differences in the underwater vegetation of the

two lakes, which may be correlated to some extent with these en¬

vironmental differences.

Various types of shore are found in different parts of Green

Lake. Along most of the north side of the lake the land ends in

smooth boulders of considerable size, behind which the shore slopes

steeply upward for a short distance (fig. 1). A beach of smaller

stones runs out a considerable distance beneath the water. To¬

wards the northwest corner of the lake rises the hill known as

Sugar Loaf (see the accompanying map), whose steep sides de¬

scend below the surface of the water at almost the same pitch,

broken only by a narrow rocky beach at the water’s edge. The

opposite shore of Norwegian Bay, and a long stretch of the south

shore of the lake (fig. 2), descend even more steeply into the water,

and consist largely of more or less sheer rock walls. At both ends

of the lake are wide sandy beaches, backed by gently rising mead¬

ows. Finally, at the mouths of the various streams that enter the

lake, and around the outlet in Dartford Bay, there are extensive

swamps and marshes (fig. 3). At the head of Norwegian Bay is

found a muddy bog.

In general the shore line is very similar to that of Mendota, with

the difference that the sandy beaches of the latter are much more

extensive.

As in Mendota, the shore line is paralleled by a belt of sub¬

merged attached plants, extending in an unbroken line all around

the lake. In Green Lake, however, this flora differs widely in

nature and distribution from that of Mendota. The wide sandy

beaches are not nearly so well covered. The rocks of the shore are

nearly destitute of the tufts of Cladophora that are so character¬

istic of Mendota. The main axis of the lake lies parallel to the

southwest winds that prevail in summer, and the consequent vigor¬

ous action of the waves is perhaps partly responsible for these

FiCi. 1. A l)oiilder-covere(l sliore. Malcolin Bay.

Fig. 2. Precipitous shore. Near Dickinson Bay.

I'fi ii'r 1^‘il liriiiii'iritliihii

Fig. 3. Marshy shore, covered with Typha and Sagittaria. Near Silver

Creek.

Bickett — The Larger Aquatic Plants of Green Lake. 383

facts. On the other hand, vegetation is very abundant in the

deeper waters, and descends to a greater depth than in Mendota.

Besides the main plant belt, consisting entirely of submerged

plants, there are near the marshy shores regions well populated

with emersed and floating forms, which merge gradually into the

flora of the swamps themselves.

Methods

The apparatus was the same as that used on Mendota. The vari¬

ous stations for collecting were reached by means of a rowboat

equipped with a detachable motor. An iron frame, half a meter,

on a side, was let down to the bottom, and all plants falling within

the area thus limited were gathered. Depths up to 3 meters

were reached by ordinary diving. Collections in deeper water (up

to 10 meters) were made by means of a diving hood, supplied with

air by a hand pump in the boat. This device permitted almost as

intimate an acquaintance with submerged plants in their habitat

as can be enjoyed with land plants in theirs. It was found pos¬

sible to stay down from 15 to 20 minutes, and to explore a con¬

siderable portion of the lake floor. The water of Green Lake being

comparatively clear and admitting a fairly bright light to these

depths, this method of survey furnished an accurate idea of the

kinds of plants present, the uniformity of their distribution, and

the downward extent of the plant belt. It is interesting to note

that the decrease in illumination is very rapid in water deeper than

8 or 9 meters. At 7 meters there is what seems to be fairly bright

sunlight ; at 10 meters one is almost in darkness.

A section of shore of uniform general characteristics usually has

opposite to it a section of the plant belt of fairly uniform nature

throughout. Stations were therefore chosen on the basis of the

character of the beach, and of the shallow water flora. The num¬

ber of stations that can be made in this way of course depends

largely upon convenience, since in many places the flora varies

greatly within a small area (for instance, in the densely populated

Stations 1 and 2). The plant belt was divided into 38 stations,

with 3 additional stations representing the marshy bays. This

was about the smallest number which would fairly represent every

type of vegetation and at the same time include the whole circum¬

ference of the lake ; it would not have been possible to collect thor¬

oughly from a larger number. The stations used are shown by

number on the map (fig. 4).

Rickett — The Larger Aquatic Plants of Green Lake, 385

Experience with Lake Mendota profited to make the collection

more systematic and more evenly distributed. An effort was made

to collect from every region at the time of flowering, thus obtaining

nearly the greatest weight attained by the plants, and making

quantitative comparisons between different species and different

localities more valuable. Plants in shallow water flower first; the

time of flowering varies directly with the depth of the water. The

early summer was therefore spent in collecting samples from the

shallow water of the lake; then collections were made in water of

medium depth; finally the deepest flora was sampled, using the

diving hood. In the latter case it was not possible in the time re¬

maining to visit all the stations ; but this does not seriously impair

the accuracy of the results, for the flora is more uniform in deeper

water and the plant belt might here be divided into fewer stations.

One station was therefore taken as representative of a group of

several adjacent ones.

Because of the previous experience in this method of collecting

it was possible, in spite of much stormy weather, to collect a larger

number of samples than in Mendota — 309 as against 221.

It soon became evident that the character of the flora varied at

different depths. For convenience in handling the data, the plant

belt was divided, as in Mendota, into three zones, within each of

which the flora may be taken as fairly uniform, but between which

there are great differences. The limits of the zones were the same

as those used in Mendota, namely : Zone 1, 0-1 meter ; Zone 2, 1-3

meters ; Zone 3, 3 meters to the deepest limit of plant growth.

Samples were brought back to the lakeside laboratory (impro¬

vised from a boathouse), and there each was separated into its

component species, the latter being numbered as sub-samples.

These were weighed and spread out to dry. When air-dry, the

smaller ones were dried in an oven at 60 °C. for 48 hours. A few

trials served to show that loss of water beyond this point was

negligible.

From the wet and dry weights of a number of samples was ob¬

tained the percentage of moisture of each species. About twelve

determinations were made for each species and averaged. In

many cases, differences between the averages of different species'

were not significant, as shown by their probable errors; such spe¬

cies were therefore averaged together and considered as having the

same percentage of moisture. Between some species, however,

there were marked differences. The values for all species are shown

386 Wisconsin Academy of Sciences y Arts, and Letters.

in table 2. It is curious that the plants of Green Lake show in all

cases a slightly higher percentage of water than those of Mendota,

although the method of determination was as nearly as possible the

same in both cases.

The Flora

A list of the species collected is given in table 1.

Most of the genera are the same as those found in Mendota.

There are more species of Potamogeton in Green Lake and a few

genera not reported in Mendota. The list given does not pretend

to include all of the species present in the lake; it shows only the

predominating ones. Identification of rare forms was not thought

to be of value in a quantitative study. Some other species may

therefore be included in the quantitative data, among the ones

named, with those they resemble most closely.

The dominant plant is Chara, — which in Mendota forms only a

small fraction of the total vegetation. Chara grows almost every¬

where in Green Lake, sometimes mixed with other plants, often

forming great masses in which no other form can get a foothold.

Badicula aquatica deserves special mention. This plant was

found only in two places (see table 4). It is usually described as

having two kinds of leaves, the immersed ones pinnately dissected

into capillary divisions, the emersed entire, serrate, or pinnatifid.

In Green Lake the plant seems never to reach the surface and flow¬

ers were not observed ; yet there are these two sorts of leaves, both

under water (it was collected at a depth of two or three meters).

The shape of the leaves would seem, therefore, not to be determined

directly by the medium in which it grows. It may be conditioned

by the intensity of the light.

Drepanocladus pseudo-fluitans, a moss, not collected in Men¬

dota, grows in deep water at low temperatures, and in a few places

forms immense mats or beds of close-growing stems and leaves, into

which one may sink to the knees. Apparently it does not fruit in

the lake, but depends wholly upon vegetative methods of reproduc¬

tion.

Castalia is found in large quantities in Dartford Bay and the

outlet and in the little bay behind Terrace Beach (Station 41).

Nymphaea occurs at the other end of the lake, behind Blackbird

Point; but in the open water of the marshes through which runs

Bickett — The Larger Aquatic Plants of Green Lake, 387

the small stream that enters the lake here, Castalia is abundant,

and there is less Nymphaea.

The marshes at the southwest corner of the lake have a rich and

varied flora. Here one finds Bidens Beckii, various species of

Typha, Sagittaria, and many others. Sagittaria latifolia is found

mostly on land or at the water’s edge; S. heterophylla, with its

long lance-shaped leaves easily mistaken for those of a Carex,

grows half submerged in shallow water. There is a Carex growing

in a similar situation, both here and in several other places; in its

young stages it may be mistaken for Vallisneria.

All the swampy or boggy parts of the shore are fronted by bars

some distance out from shore. These bars have the usual shallow

water flora and in addition large patches of Scirpus, of which sev¬

eral species were observed.

The attached algae form a smaller percentage of the total yield

of the lake than they do in Mendota, owing chiefly to the com¬

paratively small quantity of Cladophora. This plant, when present,

is found on rocks at the water’s edge, or only a few inches be¬

neath the surface. In many places it is replaced by a thin fringe

of Oedogonium. In some of the muddier stations (for example.

Stations 32 and 33), there are quantities of Spirogyra, attached

both to the mud and to rocks. The blue-greens, of which Nostoc

and Rivularia were collected, are attached to the rocks near

the shore and to the stems and leaves of many of the other plants,

especially to the species of Potamogeton. Vaucheria tuberosa was

found in one place (Station 37) in fairly deep water, — 6.5 meters,

averaging as high as 200 grams per square meter (wet weight)

over a small area. It has been described as growing in a similar

location in Lake George, N. Y., by Miss E. Moore (1).

As already indicated, the plant belt extends down to 8 meters

beneath the surface, much deeper than that of Mendota. This is

probably due largely to the greater transparency of the water. At

the water’s edge, there is, in all except the marshy places, a zone

of rocks almost barren of plants. In Mendota, where a similar

rocky beach exists, it is almost always densely covered with Clado¬

phora. In Green Lake there are occasional patches of Cladophora,

frequently a thin strip of Oedogonium, and here and there isolated

plants of Myriophyllum arising from between the rocks. Out¬

side of the border of rocks there is sometimes a thin strip of mud

or sand bearing small, scattered plants, usually Char a, Naias, and

Hetheranthera. Here the water is about 1 meter in depth and this

388 Wisconsin Academy of Sciences, Arts, and Letters.

is the limit of Zone 1. From this point the bottom falls away more

or less gently to about 6 meters, the flora passing from that in¬

cluded in Zone 2 to that characteristic of deeper water. At about

6 meters the slope usually becomes much steeper and the outer

strip of vegetation hangs, as it were, to the brow of a hill. The

plants often cease quite sharply at 8 meters, as if an invisible

boundary were holding them in check; on one side they rise two

or three meters high, and packed closely together; on the other

side there is nothing but the smooth mud sloping away towards

the bottom of the lake.

In many places, however, the slope continues gentle to a much

greater depth; in these cases the vegetation does not come to a

sudden end, but thins out gradually down an imperceptible slope.

In one such station (25) plants were found growing at a depth of

10 meters, though small and stunted in growth. Such gently slop¬

ing places are found around the entire west end of the lake, op¬

posite Woods Bay, at Forest Glen, and thence up to Dartford Bay.

In the west end of the lake and in parts of the east end, this gentle

slope occurs in connection with wide sandy beaches. Figures 5 and

6 show the difference between these two sorts of stations.

Pig. 5. Diagram of portion of lake floor bearing attached plants. Medium slope.

Fig 6. Diagram of portion of lake floor bearing attached plants. Gentle slope.

Bickett — The Larger Aquatic Plants of Green Lake. 389

In a few stations there is no flattening out from 1 to 6 meters,

but the bottom drops steeply beneath the water (Stations 16 and

30). In these cases the vegetation extends only to 4 or 5 meters

beneath the surface. This condition is represented in figure 7.

Fig. 7. Diagram of portion of lake floor bearing attached plants. Steep

slope.

There is a certain degree of correspondence between the type of

shore, the slope of the lake floor, and the vegetation. The type of

slope represented in figure 5 is the most common, and is found

opposite rocky shores of all sorts, such as those shown in figures

1 and 2. The steeper the shore, the steeper the slope of the bottom.

The second kind of slope (figure 6) corresponds to low shores,

either marshy or sandy (figure 3). The last type (figure 7) is

found only opposite high land, such as Lucas Bluff and Sugar Loaf.

In Mendota it was found that there are well-defined patches of

sand in various places, running into fairly deep water; and that

some species were more or less limited to a sandy substrate, others

to a muddy one. No such differentiation exists in Green Lake.

The type of bottom is fairly uniform; it consists of a fine mud

mixed with marl in deep water and with a small amount of sand

in shallow water. The most important difference between stations

is the presence or absence of rocks. Yet there is great regional

variation in the vegetation, and although this cannot be correlated

with any visible soil difference, it is probable that chemical analysis

of soil from different places would tell a different story. In the few

places where there are distinct sandy beaches, such as are more

common in Mendota, these are not nearly so well covered with

390 Wisconsin Academy of Sciences ^ Arts, and Letters.

plants as in the latter lake, nor, indeed, so well as are the muddier

parts of Green Lake; nor do they display any species which do

better there than on mud. The comparative scantiness of their

vegetation may perhaps be explained by the violence of the waves

in this lake ; but it is interesting that species, such as Potamogeton

Bichardsonii and'P. pectinatus, which grow decidedly better on

sand in Mendota, attain their greatest development in Green Lake

elsewhere than on these few sandy places.

Calculations

The data obtained by weighing the plants were treated in much

the same way as in the case of Lake Mendota. The original weights

obtained were reduced to common terms — grams per square meter

— for each sample and all the samples in each zone of each station

averaged together. The results are shown in tables 3, 4, and 5.

The values for all the stations of each zone were then averaged, to

give the weight per square meter of each species for each zone.

Since the stations were of greatly differing sizes, it was judged

best not to give them all the same weight in the average. One of

the smallest stations was selected as a unit, and the other stations

expressed in terms of this. The average weights in each station

were multiplied by a factor for the station, its area in terms of the

unit station, and the resulting figures averaged together, using

the sum of all the factors as a divisor. The dimensions of the sta¬

tions were obtained by measurement on the map. In Zone 1, for

instance, the area of Station 23 was found to be 9, that of Station

36 was 6, Station 1 being the unit. The results of these calculations

are given in table 6.

There were several plants that did not form part of the main

plant belt, being found in scattered patches, and yet were present

in considerable quantities. Such were Scirpus, Carex, Castalia,

Nymphaea, and Cladophora. Of the first four of these, samples

were collected in the usual way; the area of the particular spot

sampled was estimated, in most cases by rowing around it and

expressing it in terms of boat-lengths. From these data total

weights were obtained directly. The same method was used for

Cladophora in some cases, where there were large patches. In

most cases, the fringe of Cladophora being thin, all of the growth

for a certain distance was cleaned off, and measurement made of

the length (instead of the area) of the strip or station from which

Bickett — The Larger Aquatic Plants of Green Lake. 391

the sample came. The sample was converted to grams per meter,

and this value multiplied by the length of the strip to give the

total weight. The details of the data on these scattered plants are

presented in table 7.

The areas of the dilferent zones were measured on the map by

means of a planimeter ; these figures being checked up with data

obtained by the Survey at other times. By multiplying the aver¬

age weights of species (table 6) by the appropriate area, the total

weight of each species in each zone was determined. These figures,

together with those for the scattered plants (table 7) are shown in

tables 8, 9, and 10. Total weights of the species for the whole lake

flora were obtained by adding together the values for the three

zones. These are given in table 11. In each case the total weight

of a species is expressed also as a percentage of the total weight of

all plants in the zone under consideration.

From tables 8, 9, and 10, table 12 was prepared. It shows the

relative amounts of each species found at each depth, expressed as

a percentage of the total weight of the species.

Table 13 summarizes the results shown in tables 6 and 11 and

shows the average yield for each zone and for the lake as a whole,

expressed in various units. The averages for the separate zones

are the same as those in table 6, and hence disregard the weights

of the scattered plants. The latter, however, are included in the

average for the lake as a whole.

Quantitative Eesults

The 309 samples, when divided into their species, gave 1,380 sub¬

samples. This is an average of 4.5 species per sample, — ^which is

nearly equivalent to 4.5 species per square meter. The correspond¬

ing value in Lake Mendota is only 3.5. Samples containing as^

many as twelve species were not uncommon in Green Lake, but a

large number consisted of only one or two species.

The average yield of Green Lake is much smaller than that

found for Mendota. Whether this would hold all seasons is, of

course, unknown. The area of the plant zone is also less than that

of Mendota, the total yield being, therefore, very much smaller

in Green Lake.

The greatest difference between the two lakes is in the shallow

water flora. In Green Lake Zone 1 is only a little more than one-

third the area of that in Mendota. Its yield per unit area is less

than one-third. The total yield is, therefore, very much less.

392 Wisconsin Academy of Sciences, Arts, and Letters.

Zone 2 is of approximately the same size in both lakes, but has

a considerably higher yield in Mendota. Zone 3, on the other hand,

is slightly larger in Green Lake, and considerably more productive.

Putting the facts in a different way, in Green Lake, about 9 per

cent of the vegetation is in water less than 1 meter deep, about 40

per cent between 1 and 3 meters, and more than 50 per cent be¬

tween 3 and 8 meters (table 12) ; whereas in Mendota 30 per cent

is found in the shallowest water, 45 per cent between 1 and 3

meters, and only 25 per cent in water deeper than 3 meters.

About one-half of the entire vegetation (dry weight) is com¬

posed of Chara (table 11). In shallow water (table 3), this is

everywhere fairly abundant except at the east end of the lake

where the bottom is muddy (in the other swampy places there

seems to be more sand mixed with the mud). It is most frequently

associated with Potamogeton heterophyllus and small amounts of

Naias ; often it is mixed also with P. pectinatus, Heteranthera, and

Vallisneria. In Zones 2 and 3 (tables 4 and 5), the distribution of

Chara is about the' same as in Zone 1, but it is usually mixed with

most of the other species. Where it occurs unmixed, it is present

in very great abundance. One sample of Chara, gathered from

0.25 square meter, weighed 2,700 grams (wet).

The various species of Potamogeton form about 20 per cent of

the total flora. Potamogeton occurs abundantly in shallow water

only in a few stations of different characteristics. In Zone 2, how¬

ever, it thrives everywhere except for a few stations along the

north and west shores. Restriction of Potamogeton in these places

may be due to greater wave action. The shores here are of the type

shown in fig. 1. In the northwest corner of the lake, Potamogeton

is for some reason almost entirely replaced by Drepanocladus, here

present in great quantity. In the deepest zone, species of Potamo¬

geton are not abundant, except P. zosterifolius.

Ceratophyllum and Myriophyllum rank next in importance,

each forming about 10 per cent of the vegetation. The former is

found in abundance only in Zone 3, and is there universal. The

distribution of Myriophyllum is similar, but its range is slightly

shallower; it is found more commonly than Ceratophyllum in

Zone 2 and does not extend down quite so far as the latter plant

(table 12). A visit to the bottom 7 or 8 meters below the surface

reveals a forest of almost pure Ceratophyllum; whereas at 5 or 6

meters, while the general appearance of things is the same, the

vegetation is about half Myriophyllum.

Bickett — The Larger Aquatic Plants of Green Lake. 393

Potamogeton pectinatus also forms about 10 per cent of the total

weight of plants. It is distributed unevenly, but is locally very

abundant. Elodea, Vallisneria, Drepanocladus, Scirpus, and

Potamogeton zosterifoUus each form between 2 and 10 per cent

and ail other species less than 2 per cent each.

In Mendota the situation is entirely different. Chara forms a

negligible fraction of the vegetation. Vallisneria almost takes its

place, composing about one-third of all the plants. Several species

of Potamogeton bulk large, — P. amplifolius about 25 per cent,

P. Bichardsonii 10 per cent, P. pectinatus 8 per cent. Taken to¬

gether the species of Potamogeton total about 50 per cent. Myrio-

phyllum forms only 4 per cent of the total and Ceratophyllum

still less.

It has been remarked already that both Myriophyllum and

Ceratophyllum are deep water plants, as far as Green Lake is con¬

cerned; this may explain their relative scarcity in Mendota, where

the whole deep flora is so much less luxuriant. The other differ¬

ences, however, must be attributed to various factors. Tempera*

ture may perhaps hold down the Vallisneria in Green Lake, wave

action the shallow water Potamogetons, and soil differences per¬

haps account for the immense development of Chara.

The distribution of species according to depth is well marked

in some cases, in others less so. The cases of Ceratophyllum and

Myriophyllum have been dealt with already. In Mendota the

greater part of these plants is found in Zone 2. Vallisneria reaches

its greatest abundance in Zone 2 in both lakes, but is hardly found

in Zone 3 in Green Lake, while in Mendota about 25 per cent of it

is found in deep water. Ranunculus reaches deeper water in Green

Lake, half of it being found in Zone 2, and one-fourth in Zone 3;

in Mendota it is confined to Zone 1. The same is true of Chara,

which is nearly evenly distributed between Zones 2 and 3 in Green

Lake, and between Zones 1 and 2 in Mendota. Potamogeton ampli¬

folius and P. zosterifoUus are similarly distributed in both lakes;

but P. Bichardsonii and P. pectinatus are found mostly in deeper

water in Green Lake, while in Mendota their greatest growth occurs

in Zone 1.

The final averages and totals of course conceal a great deal of

regional variation. The varying characters of different stations

may be seen in tables 3, 4, and 5, which present the stations sepa¬

rately. Two general kinds of stations may be distinguished. In

one the vegetation is not very rich and is composed mostly of

394 Wisconsin Academy of Sciences^ Arts, and Letters.

Chara, with small amounts of Potamogeton heterophyllus, Naias,

and Vallisneria; the other contains great quantities of the larger

Potamogetons, often in addition to large amounts of Chara. The

shallow water is most often of the first type. The deeper water is of

the latter sort, grading out into almost pure Ceratophyllum and

Myriophyllum. The medium depths vary the most. The species of

Potamogeton often grow in great abundance mixed with Chara,

Ranunculus, and others. Chara, as already mentioned, frequently

grows alone in large patches. Drepanocladus also, in the few

places where it attains abundance, grows in large patches almost

unmixed with other plants, except that scattered stems of Myrio¬

phyllum are often found arising from it. There are in several

stations absolutely barren patches irregularly distributed amidst

luxuriant vegetation.

In some places samples of great weight were obtained. The

largest came from Station 21, and yielded 2,700 grams (wet) from

0.25 square meter. There were several others almost as big. They

were composed for the most part of Chara. There is, as far as I

know, no single spot in Lake Mendota which yields as much as

this. The smallest sample collected weighed 30 grams (wet) from

0.5 square meter. Most of the samples weighed about 200 to 300

grams (wet) from 0.25 square meter. In Mendota the general

average was somewhat higher.

General Discussion

\

Aside from its presentation of the quantitative data, this paper

cannot do much more than suggest the ecological problems that

await the botanist in this field. Pearsall (2) has attacked similar

questions in the English lakes and in an excellent paper presents

important evidence on the nature of the environmental factors

that affect submerged vegetation. He considers that light is im¬

portant only insofar as it limits the downward extension of the

flora and that temperature has little effect in determining the kind

of vegetation, at least within the limits found in one lake. Most

of the variation in the kind of vegetation proves to be connected

with soil differences, which often correspond to differences in their

physical characteristics.

Nothing has as yet been done on Wisconsin lakes which is com¬

parable to Pearsalls correlation of soil composition and type of

vegetation. With regard to the other factors, light and tempera-

Bickett — The Larger Aquatic Plants of Green Lake, 395

ture, there are abundant records from many Wisconsin lakes, whith

have been made available to me by Professor Juday. A brief com¬

parison of Green Lake and Lake Mendota serves to bring out sev¬

eral interesting pomts.

• The degree of transparency of the water was measured by means

of a white disc, 10 centimeters in diameter, which was lowered into

the water until it disappeared from view ; the depth at which this

occurred being recorded. This depth in Green Lake varied from

2.75 to 6.25 meters, the average being about 4.25 meters, during

June, July and August. In Mendota, the range during the same

months of the same year was from 1.75 to 3.8 meters, the average

about 2.25 meters. When these figures are compared with the

downward limits of the plants in the two lakes, it is evident that

they confirm PearsalPs statement that light is a limiting factor in

this respect; and, further, make it probable that it is the chief

limiting factor.

Pearsall also showed, by an iodine method of measuring the

light intensity, that plants grew in as little as 2 per cent of the

light of the surface, but not in less. According to some figures

kindly supplied me by Dr. E. A. Birge, the light intensity in Green

Lake is reduced to 1 per cent of that at the surface at a depth of

about 8 meters. The same light intensity in Mendota is found at

a depth of about 4 meters. These depths correspond approxi¬

mately with the limits of the plant zone in each case. Birge ’s de¬

terminations were made in an entirely different way from those of

Pearsall, which may partially account for the discrepancy between

the two sets of results.

With regard to temperature, table 14 shows the differences be¬

tween Green Lake and Lake Mendota. The figures are averages of

readings taken through June, July, and August. These differ¬

ences in temperature evidently are not large enough to limit the

plant zone, since plants grow in Green Lake in water 6°C. cooler

than that at which they cease in Mendota; the temperatures at

corresponding depths are also lower in Green Lake. It seems that

the lower temperatures of Green Lake may partly account for the

smaller productivity of its bottom by retarding the growth of

plants to a slight extent.

The effect of low temperatures and low light intensity in retard¬

ing growth is illustrated by the reduced stature of plants in deep

water. Most submerged plants flower at or near the surface. This

is especially true of such plants as Potamogeton, Vallisneria, and

396 Wisconsin Academy of Sciences, Arts, and Letters,

Kanunculus. Teleologically speaking, the plants in the deepest

water should grow the tallest so as to obtain more light for the

manufacture of food and for the formation of flowers and fruits.

An “adaptation” to this effect has, however, not been provided by

Nature. Plants growing 7 or 8 meters below the surface reach

heights of 1 or 2 meters, those in deeper water still less; while

those which are but 3 or 4 meters deep frequently reach the sur¬

face. These conditions are illustrated in flgures 5 and 6.

Another interesting point in this connection is that in Green

Lake many instances were observed of plants flowering before they

reached the surface, even when they were growing in fairly shallow

water. In Mendota this was not noticed, and indeed some of the

larger plants grew 5 or 6 meters to the surface before they flow¬

ered. These facts suggest that there is a minimal light intensity

for the production of flowers, which is of course realized further

beneath the surface in Green Lake than in Mendota ; so that in the

latter lake some plants remain vegetatively active for a longer

period and Anally grow to reach the surface, in spite of the re¬

tarding effects of low temperature and low light intensity. Dif¬

ferent plants probably vary in this respect. The whole argument

does not, of course, concern such plants as Ceratophyllum, which

regularly produces flowers under water.

Literature Cited

1. Needham, James G., Chancey Juday, Emmeline Moore, Charles K. Sibley,

and John W. Titcomb. A biological survey of Lake George, N. Y.

State of New York, Conservation Commission. 1922.

2. Pearsall, W. H. The aquatic vegetation of the English lakes. Jour.

Ecol. 8: 163-199. 1920.

3. Rickett, H. W. A quantitative study of the larger aquatic plants of

Lake Mendota. Trans. Wis. Acad. Sci., Arts, and Let. 20: 501-527.

1922.

Table 1. List of plants collected.

Submerged plants, forming main plant belt.

1. Ceratophyllum demersum L.

2. Chara sp.

3. Drepanocladus.

4. Elodea canadensis Michx.

5. Myriophyllum verticillatum L. var. pectinatum Wallbr.

6. Naias flexilis (Willd.) Rostk. & Schmidt.

7. Potamogeton amplifolius Tuckerm.

8. “ foliosus Raf.

9. ‘‘ heterophyllus Schreb.

Bickett — The Larger Aquatic Plants of Green Lake.

397

10. “ natans L.

11. ‘‘ pectinatus L.

12. Richardsonii (Benn.) Rydb.

13. zosterifolius Schumacher.

14. Radicula aquatica (Eat.) Robinson.

15. Ranunculus aquatilis L. var. capillaceus DC.

16. Vallisneria spiralis L.

17. Heteranthera dubia (Jacq.) MacM.

Emersed or floating plants, growing in scattered patches.

18. Bidens Beckii Torr.

19. Carex sp.

20. Castalia odorata (Ait.) Woodville & Wood.

21. Lemna trisulca L.

22. Nymphaea advena Ait.

23. Sagittaria heterophylla Pursh.

24. latifolia Willd.

25. Seirpus spp.

26. Typha sp.

27. Zizania aquatica L.

Attached algae.

28. Cladophora sp.

29. Nostoc sp.

30. Oedogonium sp.

31. Rivularia sp.

32. Spirogyra sp.

33. Vaucheria tuberosa.

Table 2. Percentage of water in plants.

398 Wisconsin Academy of Sciences ^ Arts, and Letters.

Table 3. Yield hy stations, Zone 1, stated in grams per square meter.

Species are designated by number according to their places in table 1. The

upper line opposite each species show wet weights, the lower one dry weights;

X indicates a trace.

STATIONS

Rickett — The Larger Aquatic Plants of Green Lake.

399

TABLE 3 — Continued

Stations

400 Wisconsin Academy of Sciences, Arts, and Letters,

Table 4. Yield by stations, Zone 2, stated in grams per square meter. For

explanation see heading of table 3.

Stations

1.

2.

3.

4.

5.

6.

7.

8.

9.

10.

11.

12.

13.

14.

15.

16.

Bickett — The Larger Aquatic Plants of Green Lake.

401

TABLE 4 — Continued

Stations

402

Wisconsin Academy of Sciences, Arts, and Letters,

TABLE 4 — Continued

Stations

1.

2.

3.

4.

5.

6.

7.

8.

9.

10.

11.

12.

13

14.

15,

Bickett — The Larger Aquatic Plants of Green Lake, 403

TABLE 4 — Continued

Stations

36

37

38

39

40

10.0

0.7

298.0

44.9

76.0

5.3

140.0

21.1

237.0

35.7

724.0

50.9

1844.0

322.9

270.0

18.4

5.0

0.5

15.8

1.8

8.0

0.5

260.0

23.7

60.0

5.9

40.0

4.8

120.0

14.3

12.0

1.4

12.0

1.4

5.0

0.5

12.0

1.5

8.0

0.9

8.0

1.0

300.0

35.7

118.0

9.3

311.0

30.4

0.6

16.0

9.0

1.1

49.0

6.0

44.0

5.3

138.0

16.7

33.0

5.0

24.9

117.0

11.4

3.3

0.3

430.0

51.2

7.0

0.8

203.0

21.0

387.0

23.0

2.8

230.0

27.7

3.0

0.2

468.0

33.2

24.0

2.4

27.0

3.0

49.0

3.5

2083.0

145.0

143.0

14.1

404

Wisconsin Academy of Sciences, Arts, and Letters,

Table 5. Yield hy stations, Zone 3, stated in grams per square meter. For

explanation see heading of table 3.

Bickett—~The Larger Aquatic Plants of Green Lake.

405

TABLE 5 — Continued

Stations

1,

2,

3,

4.

5,

6,

7,

8,

9,

10.

11.

12.

13.

15.

16.

17.

Wisconsin Academy of Sciences, Arts, and Letters,

TABLE 5 — Continued

Stations

37

38

20.0

1.5

X 2460.0

.... 174.7

67.0

10.0

1.2

53.0

10.4

7.0

0.4

513.0

34.8

X . 587.0

. 56.8

40.0

3.9

20.0

2.0

153.0 X X

14.9 .

X

53.0 370.0

6.4 48.8

40.0

5.7

7.0

0.8

60.0

8.5

47.0

5.3

(

Rickett — The Larger Aquatic Plants of Green Lake. 407

Table 6. Average weights of species, stated in grams per square meter.

(From tables 3, 4, and 5.)

408 Wisconsin Academy of Sciences^ Arts, and Letters,

Table 7. Weight of scattered plants.

Rickett — The Larger Aquatic Plants of Green Lake. 409

Table 8. Total weight and percentage of each species in Zone 1 (0-1 meter).

Area: 1.424 square Tcilometers. From tables 6 and 7.

410

Wisconsin Academy of Sciences, Arts, and Letters.

Table 9. Total weight and percentage of each species in Zone 2 (1-3 meters).

Area: 2.905 square Tcilometers. From tables 6 and 7.

Rickett — The Larger Aquatic Plants of Green Lake. 411

Table 10. Total weight and percentage of each species in Zone 3 (3-8 meters).

Area: 4,334 square Mlometers. From tables 6 and 7.

412 Wisconsin Academy of Sciences, Arts, and Letters.

Table 11. Total weight at all depths and percentage of each species. Area:

8.573 square Tcilometers. From tables 8, 9, and 10.

Bickett — The Larger Aquatic Plants of Green Lake, 413

Table 12. Distribution of species by depth, stated in percentages.

From tables 8, 9, 10, and 11.

414 Wisconsin Academy of Sciences, Arts, and Letters.

Table 13. Average yield of Green Lalce. From tables 6 and 11.

Table 14. Temperature of the water at different depths in summer.

THE EOTIFER FAUNA OF WISCONSIN.— II.

A Revision of the Notommatid Rotifers, Exclusive

OF THE DiCRANOPHORINAE

H. K. Harring and F. J. Myers

Notes from the Biological Laboratory of the Wiscoiisiij Geological and Nat¬

ural History Survey. XXIII.

INTEODUCTION.

At the beginning of the survey of the Wisconsin rotifer fauna a

promise v^as made to publish descriptions and figures of all the

species found. We soon realized that this 'was likely to prove quite

an ambitious program, and especially so in the case of the Notom¬

matid rotifers, the largest and also the most chaotic group of all.

Ever since this family was first proposed, there has been a steady

accretion of new species and a periodic shifting of the old ones,

until it has become a veritable Serbonian bog, carefully avoided by

everybody or, at least, trespassed upon only under compulsion.

We do not make any pretense to superior virtue or to being con¬

sidered exceptions to the rule ; the compulsion was, however,

greater and became, under the circumstances, a seemingly un¬

avoidable necessity. A beginning was made with the review of

the central group of the Notommatids appended to the preliminary

list of the rotifers of Wisconsin published in volume twenty of the

Transactions. In the present paper the remaining species are de¬

scribed, as far as material has been obtainable.

No definition was given in part one of what was meant to be in¬

cluded among the Notommatids, and it becomes necessary to ex¬

plain our conception of the family. No usable diagnosis has ever

been given and none was possible until a more detailed study of

the group became available. The tendency in the past has been

to include in the family Notommatidae nearly all the slow-moving,

plant-feeding, illoricate Ploima, without any serious attempt to

define it more precisely. We have endeavored to maintain this

416 Wisconsin Academy of Sciences, Arts, and Letters.

general idea in so far as it could be done without violating accepted

principles of classification and with proper deference to the neces¬

sarily hypothetical nature of present-day ideas of the interrelations

of the various rotifer groups.

The family Notommatidae may then be characterized as a family

of ploimate rotifers with moderately elongate, prismatic or spindle-

shaped, illoricate or partially loricate body, ending in a short, nor¬

mally two-jointed, tubular foot with two toes, in rare cases fused;

w^th a corona composed of simple cilia, primarily forming a mar¬

ginal wreath composed of strong cilia adapted to the propulsion of

the animal, and enclosing the unciliated apical plate and a buccal

field with short cilia for bringing food to the mouth, which is near

the ventral margin of the corona; the ciliation may be continued

beyond the mouth as a so-called chin. The mastax is a more or

less specialized form of the malleate type, and we have divided

the family into six subfamilies, according to the degree of de¬

parture from the type. The stomach and intestine are usually

without distinct separation; the ovary is nearly always an irregu¬

larly oval, disc-shaped organ with the normal number of nuclei,

eight; the excretory system consists of two lateral canals with 3-5

flame cells, opening into the bladder, which is either an expansion

of the cloaca or a separate, pyriform vesicle discharging into the

cloaca through a short duct. The ganglion is large and saccate;

the retrocerebral organ is very unequally developed in the different

genera. The eyespot is usually cervical, but may be frontal or

absent.

The subfamily Proalinae is characterized by a type of mastax

very closely related to the malleate; the unci are adapted to the

crushing or grinding of the food ; a weak ‘ ‘ piston ’ ’ is usually pres¬

ent, but it is attached to the ventral floor of the mastax, and not

to the fulcrum. The retrocerebral organ is absent or limited to a

rudimentary sac.

The subfamily Notommatinae has a mastax to which De Beau¬

champ applied the name virgate, first used by Hudson and Gosse

with ambiguous definition; it is characterized by the development

on its ventral surface of a powerful muscle, the hypopharynx,

attached to the fulcrum and acting as the piston of a pump, the

entire mastax forming the cylinder of the “pump”, thus enabling

the animal to extract the contents of plant cells or of the bodies of

small animals without swallowing them. A well-developed retro¬

cerebral organ is present in the majority of the genera.

Harring <& Myers — Rotifer Fauna of Wisconsin — II. 417

The subfamily Tetrasiphoninae includes only a single species;

the mastax is notable for the unusual development of the epi-

pharynx; a weak piston is present, but it is not attached to the

fulcrum. The retrocerebral organ consists of a large sac and two

long subcerebral glands.

The subfamily Lindiinae has a highly specialized type of mastax,

for which we propose the name ‘ ‘ cardate ’ ’ ; like the virgate it func¬

tions by suction, but the structure is very different; the mastax

oscillates as a unit on a transverse axis, while a complicated epi-

pharynx supports the mouth. The retrocerebral organ is limited

to a ductless sac surrounding the eyespot.

The subfamily Birgeinae with a single genus and species has a

remarkable type of mastax, characterized by the virtual atrophy

of the normal elements and their replacement by a pair of “pseud-

unci”, hook-shaped organs of epipharyngeal origin, which may

be protruded from the mouth to seize the prey. No retrocerebral

organ is present.

The subfamily Dicranophorinae includes Notommatid rotifers

having a f orcipate mastax ; the entire organ is strongly compressed

dorso-ventrally and adapted to the capture and tearing apart of

prey by protrusion through the mouth. The retrocerebral organ

is usually present, either as sac or glands. This subfamily includes

a very large number of species, the great majority still undeseribed,

and we have not been able to include them in this paper.

The distribution of the various genera among the proposed sub¬

families is indicated in the list below, as well as the species in¬

cluded in each and a reference to the description. This arrange¬

ment will no doubt need modification as more detailed information

becomes available, but if it be permitted to serve as a foundation

upon which to erect a more permanent structure, it will have

answered its purpose and accomplished all that was expected of it.

Family NOTOMMATIDAE.

Subfamily Proalinae.

Genus Proales.

decipiens (Ehrenberg) . Vol. XX, p. 603

sordida Gosse . XX, p. 605

parasita (Ehrenberg) . XX, p. 607

gigantea (Glasscott) . XXI, p. 424

wernecMi (Ehrenberg) . XXI, p. 426

brevipes Harring and Myers . XXI, p. 428

daphnicola Thompson . XXI, p. 430

418 Wisconsin Academy of Sciences, Arts, and Letters.

reinhardti (Ehrenberg) . XXI, p. 431

similis De Beauchamp . XXI, p. 434

minima (Montet) . XXI, p. 435

• doliaris (Eousselet) . XXI, p. 437

Genus Proalinopsis.

caudatus (Collins) . Vol. XX, p. 603

staurus Harring and Myers . XXI, p. 439

Subfamily Notommatinae.

Genus Notommata.

copeus Ehrenberg . Vol. XX, p. 562

pachyura (Gosse) . XX, p. 565

collaris Ehrenberg . XX, p. 568

pseudocerherus De Beauchamp . . . XX, p. 598

falcinella Harring and Myers . XX, p. 570

saccigera Ehrenberg . XX, p. 594

cerherus (Gosse) . XX, p. 572

galena Harring and Myers . XX, p. 574

aurita (Muller) . XX, p. 578

Godonella Harring and Myers . XXI, p. 444

tJiopica Harring and Myers . XXI, p. 446

peridia Harring and Myers . XX, p. 576

lenis Harring and Myers . XX, p. 586

placida Harring and Myers . XX, p. 587

pygmaea Harring and Myers . XX, p. 593

spaxia Harring and Myers . XXI, p. 443

angusta Harring and Myers . XX, p. 580

cyrtopus (Gosse) . XX, p. 582

doneta Harring and Myers . XXI, p. 448

telmata Harring and Myers . XX, p. 584

tripus Ehrenberg . XX, p. 589

■venusta Harring and Myers . XX, p. 591

tithasa Harring and Myers . . XXI, p. 450

contorta (Stokes) . XX, p. 600

silpha (Gosse) . XX, p, 596

trypeta Harring and Myers . XX, p. 602

Genus Taplirocampa.

annulosa Gosse . Vol. XXI, p. 452

clavigera Stokes . XXI, p. 455

selenura Gosse . . .XXI, p. 454

Genus Brilophaga.

judayi Harring and Myers . Vol. XX, p. 612

Genus Pleurotrocha.

petromyzon Ehrenberg . Vol. XXI, p. 459

rohusta (Glasscott) . . . . . XXI, p. 461

Harring & Myers — Rotifer Fmma of Wisconsin — II. 419

Genus Cephalodella.

Group A; eyespot frontal,, double.

catellina (Muller)

angusta Myers . .

epitedia Myers . .

paxilla Myers ....

marina Myers. . .

innesi Myers .

mineri Myers. . . .

elongata Myers . .

Group B; eyespot frontal, single.

gihba (Ehrenberg) . XXI, p. 472

gracilis (Ehrenberg) . XXI, p. 473

sterea (Gosse) . XXI, p. 474

glohata (Gosse) . XXI, p. 475

forficula (Ehrenberg) . XXI, p. 476

panarista Myers . XXI, p. 478

Group C; eyespot cervical.

auriculata (Muller) . XXI, p. 479

exigua (Gosse).., . XXI, p. 481

hoodii (Gosse) . XXI, p. 482

plioata Myers.... . XXI, p. 483

ventripes (Dixon-Nuttall) . XXI, p. 484

phy satis Myers . XXI, p. 484

strigosa Myers . XXI, p. 485

tantilla Myers . XXI, p. 486

compressa Myers . XXI, p. 487

dorseyi Myers . XXI, p. 487

hiulca Myers . XXI, p. 488

elegans Myers . .XII, p. 489

galbina Myers . XXI, p. 490

helone Myers . XXI, p. 490

nana Myers . XXI, p. 491

xenica Myers . XXI, p. 492

Group D; eyespot absent.

nelitis Myers . XXI, p. 493

melia, Myers . XXI, p. 493

megalocephala (Glasscott) . XXI, p. 494

pheloma Myers . XXI, p. 496

tenuior (Gosse)... . XXI, p. 497

retusa Myers . XXI, p. 498

dixon-nuttalli Myers . XXI, p. 498

forficata (Ehrenberg) . XXI, p. 499

intuta Myers . XXI, p. 500

collactea Myers . XXI, p. 501

inquilina Myers . XXI, p. 502

licinia Myers . XXI, p. 503

Vol. XXI, p. 465

. XXI, p. 467

. XXI, p. 468

. XXI, p. 469

. . . . .XXI, p. 470

. XXI, p. 471

420

Wisconsin Academy of Sciences^ Arts, and Letters,

vacuna Myers . XXI, p. 503

speciosa Myers . XXI, p. 504

cuneata Myers . XXI, p. 505

hyalina Myers . XXI, p. 505

papillosa Myers . XXI, p. 506

eva (Gosse) . XXI, p. 507

tenuiseta (Burn) . XXI, p. 508

apocolea Myers . XXI, p. 509

strepta Myers . XXI, p. 509

mucronata Myers . XXI, p. 510

parasitica (Jennings) . XXI, p. 511

eupoda Myers . XXI, p. 512

lipara Myers . XXI, p. 512

Genus Bbrystoma.

caudata (Bilfinger) . Vol. XXI, p. 513

Genus Bousseletia.

corniculata Harring . Vol. XXI, p. 514

Genus TylotrocJia.

monopus (Jennings) . Vol. XXI, p. 516

Genus Besticula.

melandocus (Gosse) . Vol. XX, p. 644

gelida (Harring and Myers) . XX, p. 642

anceps Harring and Myers . XXI, p. 519

nyssa Harring and Myers . XXI, p. 521

Genus EospJiora.

najas Ehrenberg . Vol. XX, p. 634

ehrenbergi Weber . XX, p. 637

therina Harring and Myers . XX, p. 639

tJioa Harring and Myers . XXI, p. 523

anthadis Harring and Myers . XX, p. 641

Genus Enteroplea.

lacustris Ehrenberg . Vol. XXI, p. 526

Genus Eothinia.

elongata (Ehrenberg) . XX, p. 646

tripliaea Harring and Myers . XXI, p. 528

argus Harring and Myers . XXI, p. 530

Genus Sphyrias.

lofuana (Eousselet) . . . Vol. XXI, p. 532

Genus Monommata.

longiseta (Muller) . Vol. XXI, p. 535

grandis Tessin . XXI, p. 538

Earring & Myers — Rotifer Fauna of Wisconsin — II.

421

Subfamily Tetrasiphoninae.

Genus Tetrasiphon.

hydrocora Ehrenberg .

Subfamily Lindiinae.

Genus Lindia.

torulosa Dujardin .

pallida Harring and Myers .

annecta Harring and Myers .

producta Harring and Myers .

Candida Harring and Myers .

tecusa Harring and Myers .

truncata (Jennings) .

fulva Harring and Myers .

Genus Birgea.

Subfamily Birgeinae.

enantia Harring and Myers

Subfamily Dicranophorinae.

Vol. XX, p. 630

Vol. XX, p. 618

. XX, p. 620

. XX, p. 622

. XX, p. 616

. XX, p. 61-4

. XX, p. 624

. XX, p. 626

. XX, p. 628

Vol. XX, p. 610

Genera: Dicranophoriis, Encentrum, Erignatha, Alhertia.

In this regrouping we have endeavored to avoid two extremes,

the one of including so many species in a genus that it becomes

unnatural and lacking in homogeneity, and the other of splitting

up the group into enough genera to make each one absolutely

homogeneous; the latter course would probably require in this

instance at least a dozen new genera. The result must therefore

be accepted as a compromise, which obviously precludes the appli¬

cation of any hard and fast rule. The precept we have tried to ob¬

serve is : when only a single structural feature in one species is in¬

volved, a new genus is not proposed unless the departure is very

striking; if, however, the same modification occurs in several spe¬

cies, in other respects apparently closely related, a new generic

name is introduced.

Some concrete examples may help to make this clear. Notom-

mat a pseudocerherus has a peculiar form of mastax, quite different

from other members of the genus, but it agrees so well with the

normal in every other respect that it has not been separated. Much

the same is true of N. saccigera; the pumping function of the mas-

tax has all but disappeared, but it is so obviously derived from the

normal virgate mastax that it seems advisable to leave it in the

genus Notommata. Three of the smaller species, Notommata

venusta, contort a and Uthasa, have the same type of corona, with-

422 Wisconsin Academy of Sciences, Arts, and Letters.

out evertile auricles, but the mastax is so different in these species

that they do not appear to form a natural group. The genus

Eothinia is separated from Eosphora on account of the form of the

mastax, virgate trophi with regularly denticulate rami, because

this occurs in three species which are very similar in other fea¬

tures. A second genus, Besticula, has been created for four species,

closely related to Eosphora, but with a type of mastax intermediate

between this genus and Notommata, adapted to prehension, but re¬

taining the pumping action unimpaired; there is also close agree¬

ment in the form of the body, retrocerebral organ and eyespot.

The compromise arrived at must be justified by the evidence of

a common ancestry brought out by a detailed study of each group.

It is quite true that extreme species often differ considerably in

their characteristics, but their actual relationship appears in a

clearer light when the entire series of intermediates are taken into

consideration.

There is also something to be said against the very fashionable

subdivision of existing genera; it may, and if carried far enough

does, lead to homogeneous and ‘‘natural’’ genera, but this advan¬

tage is obtained at the price of a more comprehensive view of the

actual relationship of the species concerned. A partial remedy is

then introduced in the form of various higher groupings, tribes,

sections, etc., but the total gain by this process does not seem to

justify the more cumbersome machinery and the burden of addi¬

tional generic and other names.

A misleading typographical error occurs several times in volume twenty; on

page 577, line 22, page 583, line 31, page 585, line 20, page 587, line 7, and

page 590, line 38, for ramus” read uncus”.

Subfamily PROALINAE.

Genus PRO ALES Gosse.

Notommatid rotifers with spindle-shaped, illoricate body, with

a slight constriction behind the mastax separating the head and

abdomen; there is usually a distinct reduction in diameter of the

body at the base of the foot, which has two very short toes.

The corona is an oblique disc with well developed marginal cilia

and two lateral tufts of densely set, long cilia, especially adapted

to swimming; they resemble auricles, but are not retractile. The

apical plate is not usually enclosed by the marginal ciliation and

Harring & Myers — Rotifer Fauna of Wisconsin — II. 423

may be dorsal. The buccal field is large and evenly ciliated; the

mouth is at the ventral margin.

The mastax is a modification of the malleate type ; the piston is

small and not attached to the fulcrum, but to the ventral wall of

the mastax. The fulcrum is short and nearly in a straight line

with the flat, roughly triangular rami, which are usually dentate

on the inner edges and have large basal apophyses. The manu-

bria are as long as in the normal virgate mastax; the unci have

four or more well-developed teeth. The epipharynx consists of

two very irregularly shaped pieces, imbedded in the walls of the

mastax at the sides of the mouth.

The retrocerebral organ is rudimentary or absent. The eyespot

is usually cervical, rarely frontal or absent.

Type of the genus, — Proales decipiens {'EihYenloerg)=Notommata

decipiens Ehrenberg.

We have not seen Proales aureus Zavadovsky, parasitic in col¬

onies of Volvox aureus, or P. quadrangular is (Glasscott) ; Dr. E,

Penard kindly sent us a slide with a balsam-mounted specimen of

his P. latrunculus, but we did not succeed in isolating the trophi

of the contracted specimen. A number of species, so poorly de¬

scribed that there is little hope of identifying them, have been re¬

ferred to this genus. They are listed here only to complete the

record.

Proales aureus Zavadovsky, Uchen. Zap. Moskovsk. Gor. Univ. Shaniavskago,

vol. f, 1916, p. 278, pi. 4, figs. 1-9.

Proales algicola Kellicott, Trans. Amer. Micr. Soc., vol. 19, 1897, p. 48

[= Cephalodella catellina (Muller) ?]

Proales coryneger Gosse, Journ. Eoyal Micr. Soc., 1887, p. 863, pi. 14, fig. 4. —

Hudson and Gosse, Eotifera, Suppl., 1889, p. 24, pi. 31, fig. 10.

Proales inflata Glasscott, Proc. Eoyal Dublin Soc., new ser., vol. 8, 1893, p.

51, pi. 4, fig. 1.

Proales latrunculus Penard, Mikrokosmos, vol. 2, 1909, p. 142, figs. 1-7.

Proales micropus (Gosse).

Furcularia micropus Gosse, in Hudson and Gosse, Eotifera, 1886, vol. 2, p.

46, pi. 19, fig. 2.

Proales micropus Jennings, Amer. Natural., vol. 35, 1901, p. 743, pi. 5, fig.

82.

Proales otJiodon, Gosse, Journ. Eoyal Micr. Soc., 1887, p. 366, pi. 8, fig. 11. —

Hudson and Gosse, Eotifera, Suppl., 1889, p. 24, pi. 31, fig. 11.

Proales prehensor, Gosse, Journ. Eoyal Micr. Soc., 1887, p. 366, pi. 8, fig. 12. —

Hudson and Gosse, Eotifera, Suppl., 1889, p. 24, pi. 31, fig. 12. (—Lecdne

depressa Bryce?)

424 Wisconsin Academy of Sciences ^ Arts^ and Letters.

Proales quadrangularis (Glasscott).

Notops quadrangularis Glasscott, Proc. Eoyal Dublin Soc., new ser., vol. 8,

1893, p. 43, pi. 3, fig. 3. — Hauer, Mitt. Bad. Landesver. Naturk., Freiburg

i. Br., new ser., vol. 1, 1921, p. 184, text fig.

Furcularia quadrangularis Murray, Trans. Eoyal Soc. Edinburgh, vol. 45,

1906, p. 180.

Furcularia glol)ulifera Hauer, Mitt. Bad. Landesver. Naturk., Freiburg i.

Br., new ser., vol. 1, 1921, p. 185

The last-named species is evidently valid, and, judging from the

figures and description given by Hauer, it should be included in

this genus.

PEOALES GIGANTEA (Glasscott).

Plate XVII, figures 6-10.

. Notommata gigantea Glascott, Proc. Royal Dublin Soc., new ser., vol. 8,

1893, p. 80, pi. 7, fig. 1.

? Proales ovicola Giard, Feuilles jeunes Nat., vol. 38, 1908, p. 184.

Proales gigantea Stevens, Journ. Quekett Micr. Club, ser. 2, vol. 11, 1912,

p. 481, pi. 24, figs. 1-5.

The body of the free-swimming animal is nearly cylindric, short

and stout; its greatest width is about one fourth of the entire

length. The integument is very soft and flexible and the outline

constantly changing.

The head is short and broad ; the neck is represented by two or

three indistinct folds, which do not encircle the body completely.

The abdomen is elongate ovoid and slightly constricted at the base

of the foot ; the tail is distinct, but not very prominent. The foot

has two short joints, both very large in diameter, and terminates

in a hemispherical bulb with the two very small toes set far apart ;

on the posterior dorsal margin of the foot there is a prominent

spur, projecting at a nearly right angle with the longitudinal axis

of the body; the toes are abruptly reduced to short, needle-like

points.

The dorsal antenna is a small, setigerous papilla in the normal

position; the lateral antennae have not been observed.

The corona is oblique and weakly ciliated with the exception of

two lateral, auricle-like areas provided with strong cilia adapted to

propelling the animal through the water. The mouth is at the

posterior margin of the corona.

The mastax is closely related to the primitive malleate type. The

incus is nearly straight; the fulcrum is long, slender and slightly

Harring & Myers — Botifer Fauna of Wisconsin — II. 425

decurved at the anterior end. The rami are broad and triangular

with a large basal apophysis; on the right ramus there is imme¬

diately behind the basal apophysis a broad, shear-like, striated and

denticulate blade projecting towards the left and opposing the first

tooth of the left uncus ; it has no counterpart on the left side. The

inner edges of the rami are not denticulate. The right uncus has

six long teeth, gradually decreasing in size from the ventral mar¬

gin; to the first tooth is joined an additional rudimentary tooth,

which is only half the length of the main tooth; the two dorsal

teeth are very slender and joined for their entire length. The left

uncus has five principal teeth, one very slender supplementary,

full length tooth and the tip of a second both joined to the first

ventral tooth; to the last, or dorsal, tooth is joined the tip of an¬

other imperfectly developed tooth. The middle cell of the manu¬

brium is long and very broad and has the usual sigmoid curvature ;

the ventral and dorsal cells are broad and plate-like. The epi-

pharynx consists of two irregular, conchoidal structures, imbedded

in the walls of the mastax at the sides of the mouth. The piston

is rudimentary and attached to the ventral wall of the mastax.

The oesophagus is fairly long and quite slender. The gastric

glands are small and nearer the ventral side than is usually the

case. There is no constriction between the stomach and intestine.

The ovary is very large and in the mature animal contains usually

from one to three developing eggs at the same time. The bladder is

very small. The foot glands are huge and completely fill the foot ;

the;y discharge into large mucus reservoirs, contained in the hemi¬

spherical bulb on which the toes are seated.

The ganglion is rather small and saccate; the minute eyespot is

at the posterior end. A large retrocerebral sac is present, but no

subcerebral glands.

Total length of the free-swimming animal 200/jt; toes Sju.

Proales gig ant ea is parasitic in the eggs of the pond snail,

Lymnaea, possibly in several species. Stevens has given an account

of its development in the eggs of Lymnaea auricularia and it is

possible that the notes in the following articles may be based on

either the eggs or the fully developed animals:

BOMME, L., 1773. Bericht aangaande verscheiden zonderlinge zee-insecten,

gevonden aan de zeewieren, op het strand van 't eiland Walcheren. — Verb.

Zeeuwsch Genootsch. Wetensch. te Vlissingen. Middelburg, vol. 3, pp. 283-

318, 1 pi. (‘^Eaderdiertje’^ in snails’ eggs pp. 302-305)

426 Wisconsin Academy of Sciences, Arts, and Letters.

KAESCH, A. F. F., 1846. Die Entwicklungsgeschichte des Limnaeus stag-

nalis, ovatus und palustris, nach eignen Beobachtungen dargestellt. — Arch.

Naturg., Berlin, Jahrg. 12, vol. 1, pp. 236-276, pi. 9.

ECKEE, A., 1851. Zur Entwickelungsgeschichte der Infusorien. — Zeitschr.

Wis. Zool., vol. 3, pp. 412-415; Froriep’s Tagesber. Fortschr. Nat.-u.

Heilk., Abt. Zool., vol. 2 (for 1852), pp. 273-275.

We have not seen the living animal; the foregoing description

is from material kindly furnished us by the late Mr. John Stevens,

of Exeter, England, who gave the first adequate account of this

species. The free-swimming young female pierces the shell of the

snails’ egg, feeds on the contents and lays its eggs within the shell,,

where the young continue the destruction of the embryo snail.

The fully grown female Proales gigantea is very much larger than

in the free-swimming stage, reaching a size of fully 500/x, and be¬

comes a shapeless, distended bag, hardly recognizable as a rotifer.

A full account, of the development of the eggs is given by Stevens.

PROALES WERNECKII (Ehrenl)erg) .

Plate XVII, figures 1-5.

Cyclops lupula Vaucher, Hist. Conf. d’Eau Douce, 1803, p. 18, pi. 3, figs. 8r,.

11s; not Cyclops lupula Muller.

Notommata wernecTcii Ehrenberg, Abh. Akad. Wiss. Berlin (for 1833), 1834,.

p. 216; Infusionsthierchen, 1838, p. 429. — Oliver, Trans. Tyneside Nat.

Field Club, vol. 4, 1860, p. 263, pi. 14. — Magnus, Verb. Botan. Ver. Prov.

Brandenburg, vol. 18, 1876, p. 125. — Wollny, Hedwigia, vol. 16, 1877, p.

163; vol. 17, 1877, pp. 5, 97. — Balbiani, Ann. Sci. Nat., Zool., ser. 6, vol. 7,.

No. 2, p. 1, pi. 4, figs. 1-18; Journ. Eoyal Micr. Soc., 1879, p. 530, pi. 18. —

Hudson and Gosse, Eotifera, 1886, vol. 2, p. 134. — Debray, Bull. Sci.

France et Belgique, vol. 22, 1890, p. 222, pi. 11. — Eothert, Zool. Jahrb.,.

Syst., vol. 9, 1896, p. 673, figs. A-D; Jahrb. Wiss. Bot., vol. 29, 1896, p.,

525, pis. 8, 9.

Copeus wernecTcii Ehrenberg, Infusionsthierchen, 1838, p. 441.

Proales wernecTcii Hudson and Gosse, Eotifera, Suppl., 1889, p. 23, pi. 32,,

fig. 18. — Eousselet, Journ. Quekett Micr. Club, ser. 2, vol. 6, 1897, p. 415,.

pi. 19, figs. 1-4. — Lucks, Eotatorienfauna Westpreussens, 1912, p. 52, fig.

8. — Voigt, Siisswasserfauna Deutschlands, pt. 14, 1912, p. 89, fig. 157. —

Weber and Montet, Cat. Invert. Suisse, pt. 11, 1918, p. 102, fig. 29.

The body of the free-swimming female is elongate, spindle-

shaped and very slender, its greatest width is about one sixth of

the total length. The integument is very flexible and the outline

constantly changing. The body is very transparent.

The head segment is considerably longer than wide ; it is rounded

anteriorly, and this portion is separated from the head proper by

Harring & Myers — Rotifer Fauna of Wisconsin — II. 427

a slight transverse fold; this corresponds to the rostrum of the

forcipate Notommatids. There is no distinct neck. The abdomen

is nearly cylindric in its anterior half ; from there it tapers grad¬

ually to an inconspicuous tail. The foot is short and relatively

slender, continuing the general spindle-shaped outline of the body ;

it has two joints of nearly equal length. The toes are moderately

long, about one twelfth of the total length, slender, conical and

slightly decurved.

The dorsal antenna is a small setigerous papilla in the normal

position ; the lateral antennae have not been observed.

The corona is oblique and has two strongly ciliated areas corre¬

sponding to the auricles of other Notommatids. The ciliation of

the buccal field does not extend beyond the mouth; the circumapi-

cal band has disappeared, as in the forcipate Notommatids.

The mastax is closely related to the malleate type. The incus is

straight; on the upper side of the fulcrum is a broad rib, which

curves around the end and continues for a very short distance on

the lower side ; the web is thin and lamellar. The rami are triangu¬

lar and decurved at their posterior ends; the inner edges do not

come into contact. The unci have only a single tooth, expanded

into a triangular basal plate, into which it gradually merges with¬

out quite reaching the malleus; this has a very small basal plate

and a rod-shaped main stem, which at the posterior end is curved

diagonally forwards and inwards. The epipharynx consists of two

sigmoid plates with a slender rib on the lower edge; they are im¬

bedded in the walls of the mastax near the base of the rami. The

piston seems to be very weak ; it is attached to the anterior wall of

the mastax and not to the fulcrum. Two huge, vacuolate salivary

glands, each nearly as large as the mastax itself, are attached to it

by a short, narrow neck.

The oesophagus is very long and slender. The gastric glands

are large and filled with highly refractive globules. The stomach

is not separated from the intestine. The ovary of the free-swim¬

ming female is normal.

The ganglion is very large and saccate. A retrocerebral sac

appears to be present, but no duct has been observed. The eyespot

is at the posterior end of the ganglion.

Total length 140-175/x; toes ll-ld^a; trophi 18 wide, 12fji long.

Proales werneckii is parasitic in galls on various species of

Vaucheria. The free-swimming young, to which the description

428 Wisconsin Academy of Sciences, Arts, and Letters.

exclusively refers, probably enters the Taitc/ierm-filament through

the point, and the alga forms a gall around it. The rotifer feeds

on the protoplasm within reach, and when mature begins to lay

eggs ; as many as 50-60 may be laid by a single individual. Simul¬

taneously the body swells enormously, becoming almost spherical;

this is caused principally by the enlargement of the stomach, which

is probably the result of the accumulation of waste materials. Ap¬

parently no discharge of faecal matter takes place in the mature

animal.

The male has been described by Rousselet; it is remarkable in

possessing a functional mastax.

Rothert has made a very complete study of Proales werneckii,

describing the rotifer in Zoologische Jahrbiicher (see synonomy),

and the formation of the galls in Jahrbiicher f. wissenschaftliche

Botanik. We give a short summary of his conclusions.

The parasite appears to enter the thallus by way of the growing

point, where the cell-wall is thin and readily pierced by the trophi ;

the galls are formed bnly where the parasite is present. When the

point of the growing thallus is injured, it starts a new growth at

the base of the gall, giving to this the appearance of a lateral

branch. The “cap’^ of the gall is structurally different from the

walls. The rotifer finally eats up the entire contents, both proto¬

plasm and chromatophores, and the gall dies ; at the same time the

cap falls off and the young animals find their way out. After the

death of the gall a new supply of protoplasm will restore the in¬

jured section to normal life. The female is unable to complete its

development outside of the gall.

We have not had an opportunity to study the living animal; the

description is based on preserved material contributed by the late

C. P. Rousselet, and no doubt will need correction in some details.

FBOALES BREVIPES Harring and Myers, new species.

Plate XIX, figures 1, 2.

The body of this species is elongate, spindle-shaped and very

slender ; its greatest width is less than one fifth of the total length.

The integument is very flexible and the outline constantly chang¬

ing. The body is very transparent.

The length of the head segment is considerably greater than its

width; the anterior portion is separated from the head proper by

a slight transverse fold and is the equivalent of the rostrum of the

Earring Myers — Rotifer Fauna of Wisconsin — II. 429

forcipate Notommatids. The neck segment is fairly long and

nearly as wide as the body at its widest point. The anterior trans¬

verse folds are well marked. The abdomen is very nearly parallel¬

sided and ends in a slightly projecting tad, under which the cloaca

opens. The foot is very stout, but little smaller in diameter than

the abdomen; the anterior joint is twice as long as the nearly

hemispherical posterior joint. The toes are minute, slender and

conical, set wide apart, and freely movable, so that the-ir tips may

be brought into actual contact in the manner of a pair of forceps.

The dorsal antenna is a small setigerous papilla in the normal

position ; the lateral antennae have not been found.

The corona is obliquely ventral and weakly ciliated with the ex¬

ception of the two frontal, auricle-like areas, which are furnished

with strong cilia adapted to swimming. The mouth is at the

posterior margin of the corona. The rostrum is outside of the

corona, as the circumapical band has disappeared.

The mastax is very nearly identical wdh that of Proales

decipiens^ and consequently no figure is given. The incus is al¬

most straight; the fulcrum is slightly tapering from the base to¬

wards the ventral end. The rami are triangular and have a large

basal apophysis ; the inner edges are obscurely dentate. The manu-

bria are elongate, rod-shaped, and expanded anteriorly into broad

plates; the unci have each five well-developed teeth. An epi-

pharynx has not been found, but may be present ; on account of the

very small size it is difficult to make out the true form of the vari¬

ous elements of the mastax.

The oesophagus is very long and slender. The gastric glands,

ovary and bladder are normal. The stomach and intestine are not

separated by a constriction. The foot glands are pyriform and

rather small.

The ganglion is large and saccate. A retrocerebral sac is pres¬

ent, but there are no sub cerebral glands ; the sac is partly fused to

the ganglion and apparently ductless. No eyespot has been found.

Total length 90-120ja; toes 5-7 jn, distance apart 7~8/x; trophi 12/x.

A few specimens of this species have been found in sphagnum

growing on the banks of ditches at G-len Burnie, Maryland; Mr.

David Bryce has found it in sphagum collected in Otsego county.

New York, by Mrs. A. C. Clarke, and sent to him. It is Mosely re¬

lated to Proales decipiens, but is easily distinguished by its peculiar

toes, smaller size and more slender body.

430 Wisconsin Academy of Sciences, Arts, and Letters

PEOALES DAPHNICOLA Thompson.

Plate XVIII, figures 1-5.

Proales daphnicola Thompson, Science Gossip, vol. 28, 1892, p. 220, fig.

125. — Murray, Trans. Eoyal Soc. Edinburgh, vol. 45, 1906, p. 179, pi. 6,

fig. 26.

f Pleurotrocha sigmoidea Skorikov, Trav. Soc. Nat. Kharkov, vol. 30, 1896,

p. 284, pi. 7, fig. 8.

Pleurotrocha daphnicola Harring, Bull. 81 U. S. Nat. Mus., 1913, p. 84. —

Myers, Proc. U. S. Nat. Mus., vol. 52, 1917, p. 478, pi. 41, figs. 4-9.

The body is spindle-shaped, short and stout; its greatest width

is about one third of the total length. The integument is soft and

flexible, but the outline is fairly constant. It is a moderately trans¬

parent species.

The head is short, broad and truncate anteriorly ; it is separated

from the abdomen by a well-marked constriction. Its width is

about two thirds of the greatest width of the body and the length

considerably less. The abdomen is somewhat pyriform, ending in

a broad, but not very prominent tail. The foot is short and very

stout; it has two joints, the basal somewhat longer and broader

than the terminal, which is obliquely truncate posteriorly. The

two toes are short, stout, and bluntly conical, ending in a minute

tubule, through which the mucus glands discharge their contents.

The dorsal antenna is a small, setigerous papilla in the normal

position ; the lateral antennae are near the middle of the body.

The corona is very slightly oblique; the marginal ciliation is

relatively weak, with the exception of two lateral auricle-like areas

with very strong cilia adapted to swimming. The apical plate is

large and unciliated ; the buccal field is evenly covered with short,

close-set cilia. The mouth is near the ventral edge of the corona.

The mastax is very robust and furnished with powerful trophi,

closely resembling the malleate type. The fulcrum is short and

very broad. The rami are of an unusual form. The basal apophy¬

sis is very large and almost as long as the ramus itself, from which

it is separated by a very deep sinus; the dorsal end curves down¬

wards. The main portion of the rami is a broad, nearly rectangu¬

lar plate ; the inner margins are not dentate, but' form small pro¬

jecting cones at their junction with the dorsal margin. The right

uncus has three clubbed teeth, gradually decreasing in size from

the ventral margin, followed by three linear teeth ; the basal plate

is nearly square. The left uncus has three clubbed and two linear

teeth. The manubria are roughly equilateral triangles ; the ventral

Harring & Myers — Rotifer Fauna of Wisconsin — II. 431

angle curves downwards and the posterior angle towards the dorsal

side; the unci are hinged near the dorsal angle of the anterior

Inargin. There is no epipharynx and apparently no piston.

The oesophagus is moderately long and slender. There is

no constriction between the stomach and intestine. The gastric

glands are large and strongly compressed laterally. The ovary is

normal. The foot glands are very large, pyriform and somewhat

compressed; they open into large mucus reservoirs, which extend

almost to the tips of the toes. There is no bladder.

The ganglion is very large and saccate. There is no eyespot and

no trace of a retrocerebral organ.

Total length 300-400/a ; toes 25-35/a ; trophi 36/a wide, 30/a long.

Proales daphnicola is not very common; since first found by

Thompson it has been recorded only by Murray from Scotland and

by Myers from California; Mr. C. F. Rousselet sent us specimens

collected at Totteridge, Herts, England. It is commensal or, more

correctly, synoecious on Daphnids and, according to Murray, on

oligochaete worms; it seems to obtain nothing but transportation

from the host.

PEOALES EEINHARDTI (Ehrenberg) .

Plate XVI, figures 6-10.

? Vortcella succolata Muller, Anim. Inf us., 1786, p. 287, pi. 40, figs. 8-12.

? Furcularia succolata Lamarck, Hist. Nat. Anim. sans Vert., vol. 2, 1816,

p. 38.

Furcularia reinhardti Ehrenberg, Abh. Akad. Wiss. Berlin (for 1833), 1834,

p. 208; Infusionsthierchen, 1838, p. 420, pi. 18, fig. 4. — Dujardin, Hist.

Nat. Zooph., Inf., 1841, p. 651. — Eichwald, Bull. Soc. Imp. Nat., Moscou,

vol. 22, pt. 1, 1849, p. 529. — Daday, Ertek. Termesz. Korebol, vol. 19, No.

17, p. 11, pi. 1, figs. 4, 13, 19-21, 27, 31. — Levander, Acta Soc. Fauna et

Flora Fennica, vol. 12, No. 3, 1895, p. 33, pi. 2, fig. 15. — Voigt, Forsch-

ungsber. Biol. Stat. Plon, vol. 11, 1904, p. 44; Siisswasserfauna Deutsch-

lands, pt. 14, 1912, p. 103, fig. 191. — Lauterborn, Mitt. Pollichia, vol. 60,

No. 19, 1904, p. 116. — Lie-Pettersen, Bergens Mus. Aarbog, 1905, No.

10, p. 31, pi. 2, fig. 8. — Murray, Trans. Eoyal Soc. Edinburgh, vol. 45, 1906,

p. 180. — Sachse, Arch. Hydrobiol., vol. 9, 1914, p. 498, fig. 1. — Hauer,

Mitt. Bad. Landesver. Naturk., Freiburg i. Br., new ser., vol. 1, 1921, p. 184.

7 Distemma marinum Ehrenberg, Infusionsthierchen, 1838, p. 450, pi. 56,

fig. 4. — Hudson and Gosse, Rotifera, Suppl., 1889, p. 32, pi. 33, fig. 16.

f Endesma (marinum) Ehrenberg, Infusionsthierchen, 1838, p. 450.

? Furcularia gammari Plate, Zeitschr. Wiss. Zool., vol. 43, 1886, p. 236, pi.

7, fig. 42; Hudson and Gosse, Rotifera, Suppl., 1889, p. 61, pi. 34, fig. 8.

432 Wisconsin Academy of Sciences, Arts, and Letters,

Mytilia tavina Gosse, in Hudson and Gosse, Eotifera, 1886, vol. 2, p. 110,

pi. 26, fig. 8.

Mytilia teresa Gosse, Journ. Eoyal Micr. Soe., 1887, p. 3, pi. 1, fig. 7;

Hudson and Gosse, Eotifera, SuppL, 1889, p. 49, pi. 31, fig. 8.

Notommata theodora Gosse, Journ. Eoyal Micr. Soc., 1887, p. 862, pi. 14,

fig. 2; Hudson and Gosse, Eotifera, Suppl., 1889, p. 21, pi. 31, fig. 8.

Mytilia poecilops Gosse, Journ. Eoyal Micr. Soc., 1887, p. 869, pi. 15, fig 21;

Hudson and Gosse, Eotifera, Suppl., 1889, p. 49, pi. 31, fig. 51.

Mytilia producta Gosse, Journ. Eoyal Micr. Soc., 1887, p. 870, pi. 15, fig.

22; Hudson and Gosse, Eotifera, Suppl., 1889, p. 49, pi. 31, fig. 53.

Notommata reinhardti Hudson and Gosse, Eotifera, Suppl., 1889, p. 22.

Biops marina Bergendal, Acta XJniv. Lundensis, vol. 28, 1892, sect. 2, No. 4, p.

83, pis. 4, 5, fig. 27.

PleurotrocJia reinhardti Von Hopsten, Zool. Bidr. Uppsala, vol. 1, 1912,

p. 187, fig. 1; Harking, Bull. 81 U. S. Nat. Mus., 1913, p. 85; Kozar, Zool.

Anz., vol. 44, 1914, p. 416.

The body is elongate, slender and spindle-shaped; its greatest

width is less than one fourth of the total length. The integument

is very flexible and the outline is constantly changing. The entire

body is very transparent.

There is a well marked transverse fold separating the head and

abdomen. The head segment is subsquare; its length is slightly

greater than the width. The abdomen increases rather rapidly in

width for about one-third of its length; from this point it tapers

gradually to the tail, which is prominent and rounded posteriorly.

The foot is two-jointed and very long; the basal joint is short and

stout, about one-third of the length of the terminal joint, which

is very slender; the length of the foot is one fourth of the total

length. It is very contractile and may be completely telescoped

within the body. The toes are long and slender and have a char¬

acteristic lanceolate form; their length is one twelfth of the total

length.

The dorsal antenna is in the normal position ; the lateral anten¬

nae are near the base of the tail.

The corona is slightly oblique. The marginal wreath has lat¬

erally two strongly ciliated, auricle-like areas; the apical plate is

unciliated and fairly large; the buccal field is covered with short,

close-set cilia. The mouth is near the ventral edge of the corona.

The mastax is of the typical form of the genus. The fulcrum is

short, slender and tapering, its extreme end curving slightly for¬

ward. The rami are broadly triangular with a large basal apophy¬

sis and the inner margins have blunt, knoblike, interlocking teeth.

The left uncus has a large ventral tooth, clubbed at the tip; this

liarring Myers — Rotifer Fauna of Wisconsin — II, 433

is followed by three linear teeth of nearly equal length; the basal

plate is subsquare. The right uncus has a large ventral, clubbed

tooth, followed by a much more slender, minutely clubbed tooth

and three linear teeth; the basal plate is sub -triangular. The

manubria are very long and strongly curved posteriorly, so that

the ends are directed inwards and towards the dorsal side of the

mastax ; the two lateral cells are small, so that three fourths of the

entire length of the manubrium belongs to the central cell only.

The epipharynx consists of two thin, curved plates, imbedded in

the walls of the mastax at the sides of the mouth. There are indi¬

cations of the presence of a rudimentary piston, attached to the

ventral wall of the mastax, but not to the fulcrum.

The oesophagus is moderately long and slender. The stomach

and intestine are not separated by a constriction. The gastric

glands, ovary and bladder are normal. The foot glands are ex¬

cessively long, the gland itself being within the body and the duct

as long as the foot.

The ganglion is moderately large and saccate. The eyespot is

near the anterior margin ; it is double, composed of two triangular

pigment cells. No retrocerebral organ is present.

Total length 250-300/x; toes 20-25/x; trophi 32/x.

Proales reinhardti is not rare in brackish tidepools; we have

collected it near Atlantic City, New Jersey. It seems to occur also

in fresh water, according to Voigt, Lauterborn, Murray, Yon

Hofsten, Sachse and Hauer.

It may be questioned whether Furcularia gammari Plate should

be considered a synonym of this species ; with the exception of the

length of the foot there is complete agreement in everything else,

and, as the foot of Proales reinhardti is highly contractile, it is not

unlikely that Plate may have had a specimen of this species before

him.

According to Yon Hofsten Distyla weissei Eichwald is ‘‘un¬

doubtedly’’ a synonym of P. reinhardti; we are unable to see a

single character in Eichwald ’s description or figures that belongs

unmistakably to this species and the extrapolation required is far

too great to make it advisable to displace the generic name Proales

by Distyla. If a guessing contest is to be admitted, Endesma

Ehrenberg has a far better claim to consideration, but many nat¬

uralists will question the wisdom of assigning to any animal char-

434 Wisconsin Academy of Sciences, Arts, and Letters.

acteristics in direct opposition to those claimed for it by the orig¬

inal discoverer.

PEOALES SIMILIS De Beauchamp.

• Plate XVI, figures 1-5.

Proales similis De Beauchamp, Bull. Soc. Zool. Prance, vol. 32, 1908, p.

153, fig. 2.

Pleurotrocha similis Von Hofsten, Zool. Bidr. Uppsala, vol. 1, 1912, p. 186;

Harking, Bull. 81 U. S. Nat. Mus., 1913, p. 85.

The body is elongate, slender and fusiform; its greatest width

is one-fifth of the total length. The integument is very soft and

flexible, and the outline is somewhat variable on account of the

contractility of the animal. It is a very transparent species.

The length of the head segment is very slightly greater than its

width and a little less than the greatest width of the body. The

abdomen is separated from the head by a slight transverse fold and

increases slightly in width for one half its length ; from this point

it tapers gradually to the foot, ending in a minute tail, projecting

but very slightly beyond the general outline of the body. The foot

is relatively long, about one sixth the total length, slightly com¬

pressed dorso-ventrally and tapering towards the posterior end;

it is without any joints, but is frequently wrinkled. The toes are

moderately long, robust and conical; near mid-length they taper

a little more rapidly and end in acute points ; their length is about

one twentieth of the total length.

The dorsal antenna is a small setigerous papilla in the normal

position; the lateral antennae are unusually far forward and only

a short distance beyond mid-length of the body.

The corona is slightly oblique and has laterally two strongly

ciliated, auricle-like areas. The apical plate is unciliated and

rather small; the buccal field is covered with short, closely set

cilia. The mouth is near the ventral edge of the corona.

The mastax is intermediate between the virgate and the malleate

type. The fulcrum is short, broad at the base and tapers grad¬

ually towards the slightly fan-shaped ventral end. The rami are

roughly triangular and have a large basal apophysis; the alulae

are two acutely pointed cones at the external angles. No denticula-

tions are present on the inner edges of the rami. The unci have

each six teeth ; the last two on the dorsal margin are partly fused.

The manubria are broad and lamellar at the base, ending in a

Harring & Myers — Rotifer Fauna of Wisconsin — II. 435

slender posterior section. The epipharynx consists of two long,

slender, slightly curved rods imbedded in the anterior walls of the

mastax at the sides of the mouth. The piston is small and attached

to the ventral wall of the mastax.

The oesophagus is moderately long and slender. The gastric

glands are large, somewhat triangular and strongly compressed.

There is no distinct separation between the stomach and intestine.

The ovary and bladder are normal. The foot glands are rather

small and pyriform ; at the base of the toes there is a minute mucus

reservoir.

The ganglion is moderately large and saccate. The retrocerebral

sac is small and ductless. The large eyespot is at the posterior end

of the ganglion.

Total length 140-180/x toes 15-20/x ; trophi 24ju wide, 15/i, long.

Proales similis was described by De Beauchamp from material

collected in brackish tidepools at Saint-Jean-de-Luz, Basses-

Pyrenees, France. We find it common in similar places near At¬

lantic City, New J ersey ; it does not occur in freshwater ponds, as

far as we know.

PROALES MINIMA (Montet).

Plate XX, figures 1-4.

Pleurotrocha minima Montet, Eev. Suisse Zool., vol. 23, 1915, p. 323, pi.

13, fig. 33.

Proales minima Weber and Montet, Cat. Invert. Suisse, pt. 11, 1918, p. 103.

The body is short, saccate and very stout; its greatest width is

nearly equal to half the length of the body proper. The integu¬

ment is very delicate and flexible, but the outline remains vir¬

tually unchanged. The entire body is very hyaline.

The head is short, broad, and truncate anteriorly; its length is

about one half the greatest width of the body. It is separated

from the abdomen by a well defined constriction immediately be¬

hind the mastax. The abdomen is ovate in outline and ends poste¬

riorly in a short' tail. The foot is two- jointed, long and slender;

the basal joint is only half the length of the terminal joint and

somewhat larger in diameter. The toes are long and very slender ;

they are nearly cylindric for one half their length, and from there

taper gradually to long, needle-like points ; their length is one fifth

of the total length.

436 Wisconsin Academy of Sciences, Arts, and Letters,

The dorsal and lateral antennae are minute setigerous papillae

in the normal positions.

The corona is slightly oblique ; the marginal wreath has laterally

two auricle-like tufts of strong cilia for propulsion through the

water. The apical plate is small and unciliated ; the buccal field is

covered with closely set, short cilia. The mouth is near the ventral

edge of the corona.

The mastax is closely related to the malleate type, but appears

to have a weak piston, attached to the ventral wall and not to the

fulcrum. The incus is nearly straight; the rami are broadly tri¬

angular and crenate on their inner margins; the basal apophysis

is very large and projects above the general surface of the rami.

The fulcrum is very short and all but rudimentary. The right

uncus has five, and the left four teeth; the ventral tooth is large

and slightly clubbed at the tip, while the remaining teeth are much

smaller and more slender. The manubrium is unusual in form, as

only the central cell, or stem, is developed; there is no trace of

the lateral, usually lamellar cells; it tapers from the base to near

mid-length and ends in a slender, rod-like distal portion, slightly

incurved at the tip. The epipharynx consists of two fairly large,

triangular plates, imbedded in the anterior walls of the mastax,

above the basal apophysis of the rami and immediately in front

of the unci.

The oesophagus is relatively short and slender. The gastric

glands are small and rounded. There is no distinct separation be¬

tween the stomach and intestine. The ovary and bladder are nor¬

mal. The foot glands are very minute and probably not functional.

The ganglion is fairly large end saccate. A rudimentary sac is

fused to the posterior end of the ganglion ; the duct is present, but

does not reach the anterior surface of the head. There is no eye-

spot.

Total length 80-1 00/a ; toes 12-1 8/x ; trophi 12/a long, 10/a wide.

Proales minima seems to be rare ; it was found by Montet in moss

that had been kept for months; we have found it under similar

circumstances. The sphagnum in which it occurred had been col¬

lected in Long Pond on the Pocono plateau by Paul Lukenbach,

of Bethlehem, Pennsylvania.

Harring & Myers — Rotifer Fauna of Wisconsin — II. 437

PROALES DOLIARIS (Eousselet).

Plate XIX, figures 3-7.

Microcodides doliaris Eousselet, Journ. Quekett Micr. Club, ser. 2, vol. 6,

1895, p. 120, pi. 7, fig. 6. Voigt, Forschungsber. Biol. Stat. Plon, vo. 11,

1904, p. 20, pi. 2, fig. 9. — Voronkov, Trudy Hidrobiol. Slants. Glubokom

Oz., vol. 2, 1907, p. 86. — Stevens, Trans. Devonshire Ass. Sci., 1912, p. 686.

Collin, Siisswasserfauna Deutschlands, pt. 14, 1912, p. 60, fig. 95. —

Jakubski, Eozpr. Wiad. Muz. Dzieduszyckich, vol. 1, No. 3-4, 1915, p. 11.

Mikrocodides doliaris Harking, Bull. 81 U. S. Nat. Mus., 1913, p. 70.

The body is short, extremely stout and gibbous; its greatest

width is nearly equal to one half the total length. The integument

is very flexible, but the outline is nevertheless quite constant. The

entire body is hyaline.

The head is short, broad and obliquely truncate, joining the

abdomen without any constriction ; its width is about one half of

the greatest width of the body. The abdomen is ovoid or nearly

spherical; it terminates in a short, sleeve-like tail surrounding the

base of the foot. The dorsal surface is marked with flve or six in¬

distinct, transverse folds. The foot is short and two-jointed; the

length of the terminal joint is barely equal to its width, of the

basal joint nearly twice the width. The toe is single, acutely

pointed and fusiform ; its length is about one tweKth of the total

length. As two normal foot glands are present, it is evident that

the single toe originated by the fusion of two separate toes and is

not to be considered as an unpaired toe; it would therefore be

erroneous to attach any special signiflcance to this distinctive fea¬

ture.

The dorsal antenna is a small setigerous papilla at the junction

of the head and abdomen; the lateral antennae are somewhat far¬

ther back than usual.

The corona is strongly oblique and consists of a circumapical

band of relatively short cilia with two lateral, auricle-like tufts of

long cilia adapted to swimming. On the unciliated apical plate

are two minute papillae with a few sensory setae ; the buccal field

is evenly ciliated. The mouth is near the ventral edge of the

corona.

The mastax represents a somewhat unusual modification of the

intermediate type common to this genus. The fulcrum is a nearly

parallel-sided, thin lamella. The medial portion of the rami form

a roughly lyrate forceps; this is extended laterally by very thin

lamellae, thus giving the incus a triangular outline. The basal

438 Wisconsin Academy of Sciences, Arts, and Letters.

apophyses take the form of long, conical, divergent, hornlike

prongs. The right ramus has near the base a lamellar projection,

curving towards the left and with five or six marginal denticles;

this is followed by four widely spaced, short, conical teeth on the

inner edge, the last one terminal. The left ramus has, opposite the

lamellar projection of the right ramus, two short teeth, very close

together, followed by three widely spaced teeth. The right uncus

has seven, and the left six, well developed teeth, clubbed at the

tips and united by a thin basal plate ; each ramus has an additional,

imperfect tooth at the ventral margin. The manubria are moder¬

ately long and have a broad basal plate. The piston is rudi¬

mentary.

The oesophagus is long and slender. The stomach and intestine

are separated by a slight constriction. The gastric glands, ovary

and bladder are normal. The two pyriform foot glands are within

the body and discharge through long, very slender ducts through

the single toe.

The ganglion is very large and saccate; the eyespot is on the

ventral side, close to the mastax and apparently affected by its

movements. No retrocerebral organ is present.

Total length 250-300/x ; toes 20-25ju- ; trophi 25/a.

Proales doliaris is not rare in regions with very soft water; we

have found it in ponds and pools in Oneida and Vilas Counties,

Wisconsin, and around Atlantic City, New Jersey.

The propriety of placing this species in the genus Proales may

be open to question; there can be none as to its removal from

Mikrocodides, which is related to Cyrtonia and not at all to the

Notommatids. However, if it is not to be made the type of a new

genus, on account of the somewhat aberrant structure of the mas-

tax, the assignment to Proales seems unobjectionable.

Genus PROALINOPSIS Weber.

Notommatid rotifers with fusiform, illoricate body, with a well-

marked constriction separating head and abdomen; there is a

gradual reduction of the diameter of the body towards the tail;

the foot is short and two- jointed, with two fairly long toes ; on the

tail or the basal foot joint is a knoblike papilla with a tuft of

setae or a spine.

The corona is an oblique disc with short marginal cilia and two

lateral, auricle-like areas with very long cilia ; the apical plate is

Harring & Myers — Rotifer Fauna of Wisconsin — II. 439

small and enclosed by the marginal wreath; the buccal field is

evenly ciliated and the mouth is at or near the ventral margin.

The mastax is intermediate between the malleate and the vir-

gate types ; the fulcrum may be either short or long ; the rami are

symmetrical, large and triangular, with a well developed basal

apophysis; the unci have about eight slender teeth, clubbed at the

tips ; the manubria are long. No epipharynx is present ; the piston

is partly or wholly attached to the fulcrum.

The retrocerebral organ is rudimentary or absent; the eyespot,

when present, is cervical.

Type of the genus. — Proalinopsis caudatus (Collins) —Notom-

mata caudata Collins.

PROALINOPSIS STAURHS Harring and Myers, new species.

Plate XX, figures 5-9.

The body is fairly slender and spindle-shaped ; its greatest width

is about one fourth of the entire length. The integument is very

flexible and the outline constantly changing in response to the

contractions of the animal. The entire body is as hyaline as

P. caudatus.

The head and abdomen are separated by a deep constriction.

The head segment is somewhat longer than wide and convex ante¬

riorly; its width is about two thirds of the greatest width of the

body. The abdomen is spindle-shaped and widest near the middle ;

from there it tapers gradually to the tail, a small, knoblike pajpilla

bearing a stiff spine, one fifth as long as the body. The foot is

stout and fairly long; it has two joints, the terminal somewhat

longer than the basal. The toes are rather stout at the base and

end in very slender, acute points; their length is a little less than

one fifth of the total length.

The dorsal antenna is a large, knoblike elevation in the normal

position ; it has a funnel-shaped central depression with a small

tuft of sensory setae. The lateral antennae have not been observed.

The corona is an elongate oval area covering the oblique anterior

surface of the head and terminating a short distance below the

mouth on the ventral side. The marginal cilia are rather short,

except on two lateral, auricle-like areas, which are provided with

long and powerful cilia for swimming. The unciliated apical plate

is small; the buccal field is evenly ciliated.

440 Wisconsin Academy of Sciences, Arts, and Letters.

The mastax is of a type intermediate between the malleate and

virgate; the primary function of the unci is evidently to crush

the food, but there is also a piston, in this instance attached to

the ventral floor of the mastax and perhaps incidentally to the ful¬

crum. The rami are triangular and apparently without teeth on

the inner edges; the basal apophysis is large and immediately be¬

hind it there is a shallow groove across the rami. The fulcrum is

short and tapers to the posterior end, which is slightly expanded.

The unci have eight or nine long, slender teeth, clubbed at the

tips and decreasing in size towards the posterior margin. The

manubrium is very long and well developed; the dorsal cell con¬

tinues almost to the posterior end as a thin lamella.

The oesophagus is short and slender. There is no distinct con¬

striction between the stomach and intestine. The ovary is of nor¬

mal form; the nuclei are large and as hyaline as the plasma, but

of higher refractive index. The posterior portion of the cloaca

appears to function as a bladder. The gastric glands are small and

nearly spherical. The foot glands are large and pyriform.

The ganglion is large and saccate. There is no trace of a retro-

cerebral organ or eyespot.

Total length lOO^i ; toes 18ju, ; fepine 22/x ; trophi 15[x.

Proalinopsis staurus is not rare among floating and submerged

sphagnum in soft water lakes and ponds. We have collected it at

Mamie Lake, Eagle River and Lac Vieux Desert, Vilas County,

Wisconsin, in lakes and ponds near Atlantic City, New Jersey, and

in sphagnum ditches in Polk County, Florida. While closely re¬

lated to P. caudatus, it is readily distinguished by the long, slender

toes, lack of an eyespot and the conspicuous posterior spine. It is

difficult to narcotize and usually rolls itself up into a ball,

armadillo-fashion.

SubfamUy NOTOMMATINAE.

Genus NOTOMMATA Ehrenberg.

Notommatid rotifers with spindle-shaped, illoricate body, having

a distin<jt constriction or neck behind the mastax, separating the

head and the abdomen; posteriorly the body is abruptly reduced

to a very short, usually two- jointed foot with two short, pointed

toes; the cloaca opens dorsally at the base of the foot, under a

projecting fold of the integument, or ^^tail.^^

Earring (& Myers — Rotifer Fauna of Wisconsin — II. 441

The corona is an elongate oval area covering the oblique ante¬

rior surface of the head and continuing beyond the mouth on the

ventral surface as a projecting chin; the marginal cilia are rela¬

tively short, except on two latero-frontal areas, provided with long

and powerful cilia adapted to swimming, in the majority of species

seated on auricles, short tubular, retractile evaginations of the in¬

tegument. The apical plate is nearly always enclosed by the mar¬

ginal ciliation and has frequently a projecting skinfold or rostrum,

at the base of which are the openings of the ducts of the retro-

cerebral organ; the buccal field and chin are covered with very

short, dense cilia ; the mouth is approximately in the center.

The mastax is virgate and usually somewhat asymmetric; the

fulcrum is very large and nearly at a right angle to the roughly^

hemispherical rami, which are occasionally faintly denticulate on

the inner edges; the manubria are long and expanded anteriorly

into broad plates ; the unci have at least one strongly developed

ventral tooth and usually some additional, more or less rudimen¬

tary teeth. The piston is a powerful, muscular organ, filling the

entire cavity of the mastax and attached to the fulcrum. Two

rod-shaped transverse supports are imbedded in the walls of the

mastax below the posterior margin of the rami; some very small,

accessory teeth are frequently attached to the ventral margin of

the unci ; an epipharynx is rarely present.

The retrocerebral organ is highly developed; the sac is always

present and the subcerebral glands are found in all but a very

few species. The eyespot is at the posterior end of the ganglion.

Type of the genus. — Notommata aurita (Muller) = Vorticella

aurita Muller.

A large number of species have been described which we have

not been able to identify satisfactorily with any of the species

known to us. The majority are probably now to be considered as

hopeless ; in some cases, at least, it is clear that the same name has

been used for a number of actually quite different species. Notom¬

mata brachyota is apparently a valid species, if we may judge

from the number of records, but we have not found an animal that

could be made to agree with Ehrenberg’s description.

Notommata hrachyota Ehrenberg, Abh. Akad. Wiss. Berlin (for 1831), 1832,

pp. 51, 132, pi. 4, fig. 8; Infusionsthierchen, 1838, p. 435, pi. 51, fig. 3. —

Hudson and Gosse, Rotifera, 1886, vol. 2, p. 24, pi. 17, fig. 1. — Wierzejski,

Rozpr. Akad. Umiej., Wydz. Mat.-Przyr., Krakow, ser. 2, vol. 6, 1893, p. 228.

442 Wisconsin Academy of Sciences , Arts, and Letters,

— Weber, Rev. Suisse Zool., vol. 5, 1898, p. 446, pi. 18, figs. 1, 3. — Voigt,.

Forscimngsber. Biol. Stat. Plon, vol. 11, 1904, p. 39; Siisswasserfauna

Deutschlands, pt. 14, 1912, p. 101, fig. 186. — Weber and Montet, Cat.

Invert. Suisse, pt. 11, 1918, p. 114.

Notommata celer Bergendal, Acta Univ. Lundensis, vol. 28, 1892, sect. 2, No.

4, p. 60.

Notommata constrict a (Muller).

Vorticella constricta Muller, Animalcula Infusoria, 1786, p. 293, pi. 42,,

figs. 6, 7.

Furcularia constricta Lamarck, Hist. Nat. Anim. sans Vert., vol. 2, 1816,

p. 38.

Notommata constricta Ehrenberg, Isis (Oken), vol. 26, 1833, col. 247.

Notommata cylindriformis Glasscott, Proc. Royal Dublin Soc., new ser., vol.

8, 1893, p. 47, pi. 3, fig. 5.

Notommata granularis Ehrenberg, Abh. Akad. Wiss, Berlin (for 1831), 1832,.

p. 133; Infusionsthierchen, 1838, p. 427, pi. 50, fig. 2.

Notommata gravitata Lie-Pettersen, Bergens Mus. Aarb., 1905, No. 10, p. 29,

pi. 2, figs. 3-5. (See Lindia tecusa).

Notommata gronlandica Bergendal, Acta Univ. Lundensis, vol. 28, 1892, sect.

2, No. 4, p. 56, pis. 2, 3, fig. 21. — Lucks, Rotatorienfauna Westpreussens,

1912, p. 52.

Notommata larviformis Glasscott, Proc. Royal Dublin Soc., new ser., vol. 8,

1893, p. 48, pi. 3, fig. 6.

Notommata Umax Gosse, Journ. Royal Micr. Soc., 1887, p. 862, pi. 14, fig. 3.

— Hudson and Gosse, Rotifera, Suppl. 1889, p. 20, pi. 31, fig. 6.

Notommata longipes Bergendal, Acta Univ. Lundensis, vol. 28, 1892, sect. 2,

No. 4, p. 66, pi. 2, fig. 20.

Notommata lucens Glasscott, Proc. Royal Dublin Soc., new ser., vol. 8, 1893, p..

79, pi. 6, fig. 6.

Notommata megaladena Schmarda, Neue wirbellose Thiere, 1859, vol. 1, p. 54,

pi. 13, fig. 111.

Notommata melanoglena Schmarda, Neue wirbellose Thiere, 1859, vol. 1, p.

53, pi. 12, fig. 109.

Notommata pentophtJialma Hilgendorf, Trans. New Zealand Inst., vol. 31,,

1899, p. 116, pi. 9, fig. 4.

Notommata pleurotroclia Ehrenberg, Ber. Verb. Akad. Wiss. Berlin, 1840,,

p. 218.

Notommata rapax Gosse, Journ. Royal Micr. Soc., 1887, p. 865.

Notommata sulcata Schmarda, Neue wirbellose Thiere, 1859, vol. 1, p. 53, pL

13, fig. 110.

Notommata symMotica Kozar, Kosmos (Lwow), vol. 36, 1911, pp. 401, 407,.

figs. 14, 15. [= Dicranophorus auritus (Ehrenberg)?].

Notommata tarda Bergendal, Acta Univ. Lundensis, vol. 28, 1892, sect. 2^

No. 4, p. 55, pi. 2, fig. 16.

Notommata volitans Glasscott, Proc. Royal Dublin Soc., new ser., vol. 8„

1893, p. 47, pi. 3, fig. 4.

Harring & Myers — Rotifer Fauna of Wisconsin — II. 443

NOTOMMATA EPAXIA Harring and Myers, new species.

Plate XXI, figures 1-5.

The body is very slender and almost cylindric ; its greatest width

is less than one fifth of the total length. The integument is very

fiexible, but the outline is quite constant. It is a very transparent

species.

The head and neck segments are separated by a slight trans¬

verse fold and a second and somewhat deeper fold divides the

neck from the abdomen ; the length of these two segments is nearly

equal to their width. The abdomen is cylindric for nearly its en¬

tire length ; posteriorly it is rapidly reduced to the base of the foot.

The tail has a single, moderately large, rounded lobe. The foot

has two joints, the posterior very short, the anterior somewhat

broader and longer. The toes are short, straight cones; their

length is about one twentieth of the total length. The abdomen is

longitudinally fluted dorsally and laterally; the ventral surface is

smooth.

The dorsal antenna is a small setigerous pit in the normal posi¬

tion; the lateral antennae have not been observed.

The corona extends down on the ventral side about one fourth

of the length of the body; the post-oral portion projects slightly

from the body, forming a chin. The rostrum or cuticular fold on

the apical plate is well marked. The auricles are rather small

and provided with strong tufts of cilia, continuous with the

corona.

The mastax is of the virgate or pumping type, but differs from

the normal in having two large salivary glands on the ventral

side, in the angles between the mallei and fulcrum. The trophi

are robust and slightly asymmetric. The fulcrum is long, slender

and tapering towards the posterior end, which is slightly expanded

and incurved. The ventral portion of the rami is triangular; at

the apex there is a strong, blunt tooth, and the inner margin of

the dorsal portion is denticulate. There is a narrow, oval opening

between the rami on the ventral side, below the anterior tooth;

the edge of the right ramus opposite this opening is smooth, while

the left ramus is denticulate and striate. The unci have a sub¬

square, lamellar basal plate ; the ventral tooth is large and clubbed

at the tip. The right uncus has two small, slender teeth close to

the ventral tooth ; from their base a diagonal rib crosses the uncus

to the dorsal margin, where it joins a very slender fourth tooth.

444 Wisconsin Academy of Sciences, Arts, and Letters. .

The right uncus has only one slender tooth following the ventral

tooth, but is otherwise identical with the left. At the point of the

right uncus there are three very small accessory teeth; the left

uncus has four similar teeth, gradually decreasing in size. The

manubria are nearly straight, with a broad, subsquare lamella at

their anterior ends. A pair of slightly curved rods are imbedded

in the walls of the mastax near the posterior edge of the rami and

parallel with the dorsal branch ; their function is to aid in keeping

the cavity open during the pumping action.

The oesophagus is long and slender. There is a slight constric¬

tion between the stomach and intestine. The gastric glands, ovary

and bladder are normal. The foot glands are small and club-

shaped ; they discharge into a minute, spherical mucus reservoir at

the base of the toes.

The ganglion is large and saccate. The retrocerebral sac is

nearly spherical and the spaces between its vacuoles are crowded

with bacteroids, giving it the appearance of being filled with small,

black globules. The subcerebral glands are large and vacuolate,

but ductless and fused with the ganglion. The eyespot is at the

posterior end of the ganglion.

Total length 225-250/^; toes 12-14/a; trophi 38/a.

Notommata epaxia was collected at Oceanville, near Atlantic

City, New Jersey, where it occasionally occurs in small numbers.

It is closely related to N. aurita, but is readily distinguished by its

much more slender form, the striation and the much smaller retro-

cerebral sac, as well as the large mastax with its salivary glands.

NOTOMMATA CODONELLA Harring and Myers, new species.

Plate XXI, figures 6-10.

The body is elongate, slender and spindle-shaped; its greatest

width is less than one fourth of the total length. The integument

is very flexible, but the outline remains quite constant. The en¬

tire body is very transparent.

The transverse folds limiting the head and neck segments are

well marked. The head segment is short and broad ; the neck seg¬

ment increases very slightly in width towards the posterior end;

its length is somewhat greater than the width. The abdomen con¬

tinues the outline of the neck segment and increases in width for

about three fourths of its length; it is rounded posteriorly and

ends in a very broad tail with a large, truncate median lobe and

Earring (& Myers — Rotifer Fauna of Wisconsin — II, 445

two minute lateral lobes. The foot is two- join ted and very short;

the basal joint does not project beyond the tail. The two toes are

short, slender and conical; their inner margins are nearly straight

and the outer slightly curved; the length is about one sixteenth

of the total length.

The corona extends down on the ventral side about one fourth

the length of the body; the post-oral portion projects from the

body as a fairly prominent chin. The auricles are short and stout,

with robust tufts of cilia, continuous with the corona.

The dorsal and lateral antennae are small setigerous papillae

in the normal positions.

The mastax is of the normal virgate type and the trophi some¬

what asymmetric. The fulcrum is long and slender and tapers

towards the posterior end, which is slightly expanded and in¬

curved. The rami have fairly deep transverse grooves below

the accessory teeth of the unci; the right ramus has in front of

this groove a broad, subsquare, lamellar tooth, projecting diagon¬

ally towards the left ; the anterior margin is coarsely dentate. The

dorsal section of the rami immediately below the unci is strongly

curved and the inner, opposing edges crenate. The right uncus

has a linear ventral tooth, followed by a strong, clubshaped tooth ;

behind this are two smaller, clubshaped teeth, very close together,

and from their base a diagonal rib crosses the uncus to unite with

a linear, rudimentary tooth just inside the dorsal margain of the

basal plate uniting all the teeth. The left uncus has a linear

ventral tooth and a stout, clubshaped tooth, larger than its mate

on the opposite side; close to this there is a second, somewhat

smaller tooth. From the base of this a somewhat curved, rudi¬

mentary fourth tooth crosses the basal plate diagonally; nearly

parallel to this and separated from it by a considerable space is

a rudimentary fifth tooth.

Short accessory teeth are attached to the ventral edges of the

unci at their tips; the right uncus has four very slender teeth,

and the left three broad, obtuse teeth, successively decreasing in

size. The central section of the manubrium is very stout and

curves towards the ventral side at the tip ; the basal plate is large

and irregular in outline. A pair of curved rods, attached at their

ventral ends to the inner surfaces of the rami, pass under the

manubria and terminate below the dorsal tips of the rami; they

are imbedded in the walls of the mastax and assist in supporting

them during the pumping action.

446 Wisconsin Academy of Sciences, Arts, and Letters.

The oesophagus is long and slender. The gastric glands are

small and rounded. There is no constriction between the stomach

and intestine. The ovary and bladder are normal. The foot glands

are short and pyriform.

The retrocerebral sac is elongate conical and truncate poste¬

riorly ; including the duct its length is fully one third of the length

of the entire body. The subcerebral glands are elongate and

fusiform ; their length is one half the length of the sac. Bacteroids

are abundant in the posterior end of the sac and in the glands be¬

yond the level of the eyespot. The ganglion is relatively small

and saccate ; the eyespot is at its posterior end.

Total length 300-350/;t ; toes 18-21/x ; trophi 38/x long, 45ja wide.

N otommata codonella is locally fairly common, but seems to be

found only in neutral or slightly acid waters. We have collected

it in a shallow bay of Range Line Lake' near Three Lakes, Oneida

County, Wisconsin, and in ponds in Atlantic County, New Jersey.

It is closely related to N. cerherns, N. galena, N. collaris, etc. ; but

is readily recognized by the retrocerebral sac and the broad, trun¬

cate tail.

NOTOMMATA THOPICA Harring and Myers, new species.

Plate XXII, figures 5-9.

The body is elongate, slender and spindle-shaped; its greatest

width is less than one fourth of the total length. The integument

is very flexible, but the outline remains fairly constant. The body

is transparent, but somewhat milky.

The transverse folds limiting the head and neck segments are

well marked. The head segment is short and broad; the neck

.segment is almost parallel-sided and its width nearly equal to its

length. The abdomen is deeply striate longitudinally and in¬

creases very slightly in width for about two thirds of its length;

it is rounded posteriorly and ends in a short tail with a very broad,

rounded median lobe and two minute lateral lobes. The foot has

two very short joints; the basal joint projects by about one half

of its length beyond the tail. The two toes are short, slender and

conical, ending in fairly acute points; their length is about one

sixteenth of the total length.

The corona extends down on the ventral side about one fourth

of the length of the body; the post-oral portion projects from the

Harring & Myers — Rotifer Fauna of Wisconsin — II. 447

body as a fairly prominent chin. The auricles are moderately

large and the ciliation continuous with the corona.

The dorsal antenna is a large, circular papilla with a shallow

central depression bearing a minute tuft of setae; the lateral an¬

tennae are minute tubules with a few long setae in the normal

positions.

The mastax is virgate and slightly asymmetric. The fulcrum

is long and moderately stout, tapering slightly towards the poste¬

rior end, which is somewhat enlarged and incurved. The ventral

portion of the rami is roughly semicircular; at the apex there is

on the left ramus a single, blunt tooth, interlocking with two sim¬

ilar teeth on the right ramus. The dorsal portion of the rami is

not denticulate. There is a narrow opening between the ventral

edges of the rami, beginning above the inconspicuous basal

apophysis of the right ramus and limited anteriorly by the apical

teeth ; the edge of the right ramus opposite this opening is smooth,

while the left has about 8 or 9 minute, close-set teeth or denticles.

The alula of the left ramus, which is somewhat longer than the

right, ends in a needle-like, slightly curved spur, which is char¬

acteristic for this species. The unci are asymmetric and have only

one well developed tooth. The left uncus has a subsquare basal

plate with a linear rudimentary tooth crossing diagonally from

the base of the ventral tooth to a point close to the two parallel,

rudimentary teeth at the dorsal margin. Close to the ventral tooth

are two very small teeth, which are reduced to little more than

the clubshaped points; between these and the diagonal tooth there

is an additional, blunt, marginal tooth. The right uncus has a

strongly curved, rudimentary tooth limiting the basal plate on

the dorsal side; the diagonal tooth does not project beyond the

margin of the basal plate. Following the ventral tooth are three

rudimentary teeth, of which only the middle one can be traced be¬

yond the margin of the basal plate. Four accessory teeth, gradu¬

ally decreasing in size, are attached to the point of the left uncus ;

the right has three similar, but more slender teeth. The manubria

are nearly straight, with a broad, subsquare basal plate ; the poste¬

rior ends are slightly incurved and decurved posteriorly. A pair

of supporting rods are imbedded in the walls of the mastax be¬

hind the posterior edges of the dorsal branch of the rami; their

dorsal ends are slightly expanded and upcurved The piston is

large and muscular, filling the entire cavity of the mastax. There

448

Wisconsin Academy of Sciences y ArtSy and Letters.

are two large salivary glands in the ventral angles of the mastax,

between the fulcrum and manubria.

The oesophagus is very long and slender. There is a slight

reduction in diameter marking the separation of the stomach and

intestine, but no distinct constriction. The gastric glands are large,

thin and disc-shaped. The ovary and bladder are normal. The

foot glands are small and pyriform; they discharge into a mucus

reservoir at the base of the toe, nearly half as large as the gland

itself.

The ganglion is moderately large and saccate. The retrocerebral

sac is very long and pyriform ; bacteroids are occasionally present,

scattered through the posterior half of the sac. The subcerebral

glands are saccate and almost as long as the sac ; they always con¬

tain bacteroids, collected into very dense, globular masses, thus

simulating an additional pair of eyespots. The true eyespot is at

the posterior end of the ganglion, just below the dorsal antenna.

Total length 250-300/x ; toes 15-18ja ; trophi 36/x.

Notommata thopica was first found by Mr. L. M. Dorsey, of the

Philadelphia Academy of Natural Sciences, in a pond in Fairmount

Park, Philadelphia, Pennsylvania. It is common at Bargaintown,

near Atlantic City, New Jersey. It is closely related to N. epaxia,

N. codonella and other species of the central group of the genus,

but readily distinguished by the two globular aggregations of

bacteroids in the subcerebral glands, as well as by the salivary

glands. It is possible that this may be Ehrenberg’s Triophthalmus

dorsualisy but as he insists that the two additional eyespots are not

to be confused with the black, granular masses of other Notom-

matids and Otoglena, it seems best not to use his name for this

species, especially as it is not known whether it occurs in Germany

at all. If his drawing really looks like the animal he saw, it is

probably an iJosp/iora-species.

NOTOMMATA DONETA Harring and Myers, new species.

Plate XXII, figures 1-4.

The body is slender and spindle-shaped; its greatest width is

about one fifth of the total length. The integument is very fiexible,

but the outline is quite constant. It is a very transparent species.

The head and neck segments are of nearly equal length, a little

less than their width. The anterior transverse folds are well

marked. The abdomen increases very gradually in width for about

Earring & Myers — Rotifer Fauna of Wisconsin — II. 449

three fourths of its length and is rounded posteriorly ; it is faintly

plicate longitudinally. The tail is prominent and has a single,

median, rounded lobe. The foot has two very short joints, the

terminal smaller than the basal joint. The toes are long and very

slender; the basal portion, about one third of the total length, is

straight, and from this point they curve somewhat abruptly out¬

wards and downwards, ending in very acute points. Their length

is about one sixth of the total length.

The dorsal and lateral antennae are small setigerous papillae in

the normal positions.

The corona extends down on the ventral side somewhat more

than one fourth of the length of the body; the post-oral portion

projects from the body as a small chin. The auricles are moder¬

ately large, and the ciliation is continuous with the corona.

The mastax is virgate and the trophi very nearly symmetrical.

The rami are broadly triangular in ventral view and curve gradu¬

ally from the base to the dorsal points. The basal apophysis is

very prominent and is separated from the main portion of the

ramus by a deep sinus. The inner margins of the rami are armed

with about twelve minute teeth, beginning a short distance behind

the basal sinus and continuing to the point of attachment of the

unci, gradually increasing in length. The fulcrum is long and

slender, tapering gradually to the posterior end, which is slightly

incurved. The unci have a single, well developed ventral tooth

and a rudimentary second tooth, crossing the basal plate diagonally

to its dorsal angle. Seven small accessory teeth are attached to

the point of the left uncus and six to the right uncus. The

manubria are long and slender, with a small basal plate. A pair

of slightly curved, slender rods are imbedded in the walls of the

mastax below the posterior edge of the rami and assist in its sup¬

port during the pumping action. The epipharynx consists of two

very slender, tapering and decurved rods immediately in front

of the unci.

The oesophagus is moderately long and slender. The gastric

glands, ovary and bladder are normal. There is no distinct sep¬

aration between stomach and intestine. The foot glands are small

and pyriform.

The retrocerebral sac is large and rounded; bacteroids are not

abundant. The subcerebral glands are very large, vacuolate and

ductless; they are fused to the posterior end of the large, saccate

450 Wisconsin Academy of Sciences, Arts, and Letters.

ganglion at the level of the eyespot without any distinct junc¬

tion line.

Total length 275-300/>i; toes 45-50ja; trophi 40, u.

Notommata doneta is rare ; we have found only a few specimens

in Starvation Lake, about 4 miles south of Eagle Kiver, Vilas

County, Wisconsin, and at Oceanville, near Atlantic City, New

Jersey. Its closest relatives are N. aurita, N. cyrtopus and N. tel-

mata; the unusually long and peculiar toes are sufficient to dis¬

tinguish it from these species.

NOTOMMATA TITHASA Harring and Myers, new species.

Plate XXIII, figures 1-5.

The body is elongate, spindle-shaped and slender; its greatest

width is about one fifth of the total length. The integument is

very fiexible and the outline is constantly changing in response

to the contractions of the animal. The entire body is transparent,

but has a very faint yellowish tinge.

The neck is very slightly constricted, but no anterior transverse

folds are present. The head is rather small and triangular in*

dorsal view on account of a large, slightly obtuse rostrum. The

abdomen increases very gradually in width for about three fourths

of its length and then tapers somewhat more rapidly to a broad

tail projecting very slightly from the body. The foot is stout,

fairly long and obscurely wrinkled. The toes are very long and

stout, incurved and decurved; the ventral edge does not curve

evenly, but is faintly indented near the tip and a short distance

from the base. Their length is one eighth of the total length.

The dorsal antenna is a small setigerous papilla in the normal

position; the lateral antennae are somewhat farther back than

usual.

The corona has two large, strongly ciliated areas corresponding

to the auricles of other Notommatids, but not evertile. The buccal

field is evenly ciliated and continues down on the ventral side of

the body for about one fourth of its length, forming posteriorly

a slight chin. The dorsal arc of the eircumapical band has dis¬

appeared.

The mastax is of a very simple virgate type and the trophi

very small. The fulcrum is rodlike and very slender, slightly

curved posteriorly^ The rami are triangular and without denticu-

lations on the inner margins, which do not quite meet, but enclose

Harring & Myers — Rotifer Fauna of Wisconsin — II. 451

a narrow, elongate, ventral opening; the alulae are very large

and curved. The unci are small, triangular lamellae with a very

faint median tooth. The manubria show no indication of cellular

division; the basal plate is triangular and the posterior portion

rodlike, ending in a small crutch. Instead of being approximately

parallel to the longitudinal axis of the mastax, as in other species

of the genus, the manubria are nearly at right angles to it. The

epipharynx consists of two small, very thin, triangular plates, im¬

bedded in the walls of the mastax at the sides of the mouth. The

piston is large, but very weak.

The oesophagus is moderately long and slender. There is no

constriction between stomach and intestine. The gastric glands are

small and rounded. The ovary and bladder are normal. The foot

glands are pyriform and nearly as long as the foot.

The ganglion is an elongate, pyriform sac; at its posterior end

is a rounded, moderately large, ductless retrocerebral sac, which

does not contain bacteroids. No eyespot is present.

Total length 175-200/x; toes 25-28/x; trophi 15/x.

Notommata tithasa is not common; we have found it at Three

Lakes, Oneida County, Wisconsin, and in ponds around Atlantic

City, New Jersey; a few specimens occurred in some collections

made by Dr. H. S. Jennings in the Huron Kiver at Ann Arbor,

Michigan. This species is readily recognized by the unusually

long and robust toes.

Genus TAPHROCAMPA Gosse.

Notommatid rotifers with cylindric or spindle-shaped, illoricate

or semiloricate body, marked with permanent or evanescent trans¬

verse folds; the foot is rudimentary and the cloaca opens dorsally

at the base of the toes, under a projecting fold of the integument

or tail.

The corona is an elongate oval area covering the oblique anterior

surface of the head and continuing beyond the mouth on the ven¬

tral surface; the marginal cilia are relatively short, except on the

auricles, which have long and powerful cilia adapted to swimming.

The apical plate is enclosed by the marginal ciliation; a rostrum

may be present ; the buccal field is covered with short, dense cilia ;

the mouth is approximately in the center of the corona.

The mastax is virgate with strongly asymmetric trophi ; the

fulcrum is long and slender and nearly at a right angle to the

452 Wisconsin Academy of Sciences, Arts, and Letters.

roughly hemispherical rami; the manubria are long and slender

with a rudimentary basal plate; the unci have a strongly devel¬

oped ventral tooth and one or two additional, rudimentary teeth.

The piston is very large and fills the entire cavity of the mastax.

The rodshaped transverse supports, which are found imbedded in

the walls of the mastax below the posterior margin of the rami

in many Notommatids, are absent in this genus.

The retrocerebral sac is present in all of the species, but there

are no subcerebral glands. The eyespot is at the posterior end

of the ganglion.

Type of the genus. — Taphrocampa annulosa Gosse.

TAPHROCAMPA ANNULOSA Gosse.

Plate XXIII, figures 6-10,

Taphrocampa annulosa Gosse, Ann. Mag. Nat. Hist., ser. 2, vol. 8, 1851,

p. 199. — Hudson and Gosse, Eotifera, 1886, vol. 2, p. 16, pi. 17, fig. 12. —

Petr, Sitzungsber. Bohm. Ges. Wiss. (for 1890), 1891, p. 220. — Ternetz,

Rotat. Umg. Basels, 1892, p. 11. — Glasscott, Proc. Royal Dublin Soc.,

new ser., vol. 8, 1893, p. 43. — Wierzejski, Rozpr. Akad. Umiej., Wydz.

Mat.-Pryzr., Krakow, ser. 2, vol. 6, 1893, p. 227 — Levander, Acta Soc.

Pauna et Plora Pennica, vol. 12, No. 3, 1895, p. 26. — Weber, Rev. Suisse

ZooL, vol. 5, 1898, p. 433, pi. 17, figs. 11-13. — Stenroos, Acta Soc. Pauna

et Plora Pennica, vol. 17, No. 1, 1898, p. 123. Hempel, Bull. Illinois

State Lab. Nat. Hist., vol. 5, 1898, p. 369. — Jennings, Bull. U. S. Pish

Comm., vol. 19 (for 1899), 1900, p. 84, pi. 16, figs. 13, 14. — Voigt, Por-

schungsber. Biol. Stat. Plon, vol. 11, 1904, p. 38; Siisswasserfauna Deutsch-

lands, pt. 14, 1912, p. 92, fig. 165. — Voronkov, Trudy Hidrobiol. Slants.

Glubokom Oz., vol. 2, 1907, p. 197. — Kofoid, Bull. Illinois State Lab. Nat.

Hist., vol. 8, No. 1, 1908, p. 217. — Lie-Pettersen, Bergens Mus. Aarb.

(for 1909), 1910, No. 15, p. 38. — Mola, Ann. Biol. Lac., vol. 6, 1913, p.

240. — ^Weber and Montet, Cat. Invert. Suisse, pt. 11, 1918, p. 105.

The body is elongate, slender and very nearly cylindric, taper¬

ing slightly to the foot; its greatest width is about one fourth of

the total length. The integument is leathery and almost semi-

loricate; its dorsal surface is viscous and usually covered with

adhering particles of fioccose material.

There is no distinct separation between the head and abdomen;

both are transversely plicate, about ten ridges in the entire length,

of which three may be taken as belonging to the head. The plica¬

tions do not form rings around the body; the lateral ridges alter¬

nate with those of the dorsal surface, forming a sort of ‘ * bellows ’

fold. The ventral surface is very faintly plicate. The tail is as

Harring & Myers — Botifer Fauna of Wisconsin — II. 453

wide as the body and rounded posteriorly; it is separated from

the abdomen by a deep transverse groove. The foot is short and

very broad; it has no joints. The toes are rather short, slender,

conical and slightly decurved ; their length is about one fifteenth

of the length of the body, and they are separated by an interspace

of nearly half their length.

The dorsal antenna is a minute setigerous pit in the normal po¬

sition ; the lateral antennae have not been observed.

The corona is strongly oblique and extends down on the ventral

side about one fourth of the length of the body ; the post-oral por¬

tion projects very slightly from the body, but does not form a

chin. The auricles are small and rounded; the ciliation is con¬

tinuous with the corona. The mouth is nearly in the center.

The mastax is virgate with very slender and slightly asymmetric

trophi of a simple type. The fulcrum is long and slender, tapering

gradually to the posterior end, which is expanded into an oval

plate for the attachment of the muscles of the piston. The rami

are approximately triangular in ventral view and have well de¬

veloped alulae, ending in acute points; the right ramus has below

the apex a small, blunt tooth, fitting into a recess on the left

ramus. On the external edge of the rami there is a strongly curved

dorsal extension, which forms a nearly right angle with the rami;

a deep sinus separates it anteriorly from the points of the rami.

The unci have two teeth, of which the ventral one is more robust

than the second tooth ; the basal plate is very narrow and lamellar,

with a rudimentary tooth at its dorsal edge. The manubria are

slightly curved near the base ; the posterior portion is straight and

very slender, ending in a spatulate expansion, which curves slightly

inwards. The piston is very large and fills the entire cavity of

the mastax.

The oesophagus is moderately long and begins high up on the

mastax. The gastric glands, ovary and bladder are normal. There

is a slight constriction between the stomach and intestine. The

foot glands are small and pyriform.

The ganglion is large and saccate. The retrocerebral organ

consists of a small, pyriform sac, usually filled with bacteroids.

The eyespot is at the posterior end of the ganglion.

Total length 175-200/x; toes 12-15ja; trophi 26/x.

Taphrocampa annulosa is common everywhere in ponds and

among wet mosses. It is closely related to Taphrocampa selenura,

454 Wisconsin Academy of Sciences , ArtSy and Letters.

but is readily distinguished by its smaller size and short toes, as

well as by the differences in the trophi.

TAPHBOCAMPA SELENUBA Gosse.

Plate XXIV, figures 5-9.

Taphrocampa selenura Gosse, Journ. Eoyal Micr. Soc., 1887, p. 1, pi. 1, fig.

1. — Hudson and Gosse, Rotifera, Suppl., 1889, p. 20, pi. 31, fig. 5. — ^Wier-

ZEJSKI, Eozpr. Akad. Umiej., Wydz. Mat.-Przyr., Krakow, ser. 2, vol. 6,

1893, p. 227. — Weber, Eev. Suisse Zool., vol. 5, 1898, p. 436, pi. 17, figs.

14, 15. — Voigt, Porschungsber. Biol. Stat. Plon, vol. 11, 1904, p. 38;

Susswasserfauna Deutschlands, pt. 14, 1912, p. 92, fig. 166. — ^Voronkov,

Trudy Hidrobiol. Slants. Glubokom Oz., vol. 2, 1907, p. 95. — Montet, Eev.

SuissQ Zool. vol. 23, 1915, p. 325. — Weber and Montet, Cat. Invert. Suisse,

pt. 11, 1918, p. 106.

Taphrocampa viscosa Levander, Acta Soc. Fauna et Flora Fennica, vol. 12,

No. 3, 1895, p. 26, pi. 2, fig. 14. — Stenroos, Acta Soc. Fauna et Flora

Fennica, vol. 17, No. 1, 1898, p. 123.

The body is elongate, slender and very nearly cylindric, taper¬

ing slightly to the foot; its greatest width is about one fifth of

the total length. The integument is leathery and almost semi-

loricate; its surface is viscous and usually covered with adhering

particles of floccose material.

There is no distinct separation between the head and abdomen;

both are transversely plicate, about eleven ridges in the entire

length, of which three may be considered as belonging to the head.

The plications do not form rings around the body; the lateral

ridges alternate with those on the dorsal surface and meet them

in a “ bellows ’’-fold. The ventral surface is very faintly plicate.

The tail is slightly narrower than the body and rounded poste¬

riorly; it is separated from the abdomen by a deep transverse

groove. The foot is rudimentary, the anus opening just above the

toes; these are long, slender, tapering and decurved; they are far

apart at the base and form nearly a semicircle, when seen from

the dorsal or ventral side ; their length is about one ninth of the

total length.

The dorsal antenna is a minute setigerous pit in the normal po¬

sition; the lateral antennae have not been observed.

The corona is strongly oblique and extends down on the ventral

side for nearly one third of the length of the body; there is no

chin. The auricles are small and rounded; the ciliation is con¬

tinuous with the corona. The mouth is a little below the center.

Barring & Myers — Rotifer Fauna of Wisconsin — 11. 455

The mastax is virgate and the trophi slender and strongly asym¬

metric, the left side being more strongly developed than the right.

The fulcrum is long and slender and tapers towards the posterior

end, which is slightly enlarged and bent inwards, providing attach¬

ment for the muscles of the piston. The rami appear triangular in

ventral view and are bent nearly at a right angle at the extreme

anterior point. The left ramus has near the base a prominent,

blunt tooth on its inner edge, and immediately behind this a broad,

lamellar and finely striated tooth. Near the anterior angle there

are two large teeth, separated by a slight interspace. The right

ramus is coarsely denticulate; about eight rudimentary teeth are

present. The left uncus has a large ventral tooth, followed by a

much smaller and shorter second tooth; the right uncus has a

similar ventral tooth, followed by two smaller teeth; the basal

plates are somewhat triangular in outline and bordered by a mar¬

ginal rib. The manubria have a subsquare basal plate, with a

straight posterior branch ending in a triangular, plate-like ex¬

pansion. The piston is large and fills the entire cavity of the

mastax.

The oesophagus is moderately long and begins high up on the

mastax. The gastric glands, ovary and bladder are normal. There

is no distinct separation between the stomach and intestine. The

foot glands are small and pyriform.

The ganglion is moderately large and saccate. The retrocerebral

organ consists of a rather small, pyriform sac, usually opaque with

bacteroids. The eyespot is at the posterior end of the ganglion.

Total length 225-275/x; toes 25~30/>i; trophi 36/>t.

Taphrocampa selenura is widely distributed, but always in small

numbers. It is closely related to Taphrocampa annulosa, but is

readily distinguished by its larger size, as well as by the peculiar

form of the toes and the more robust trophi.

TAPHROCAMPA CLAVIGEEA Stokes.

Plate XXIV, figures 1-4.

Taphrocampa clavigera Stokes, Ann. Mag. Nat. Hist., ser. 6, vol. 18, 1896,

p. 18, pi. 7, fig. 2.

The body is elongate, fusiform and very slender; its greatest

width is about one sixth of the total length. The integument is

soft and very fiexible and the outline is constantly changing.

There is no distinct separation between the head and abdomen;

the contracted animal is marked with evanescent annulations,

456 Wisconsin Academy of Sciences, Arts, and Letters.

which disappear when the body is fully extended in swimming.

There is no true foot and the tail is a small fold above the toes;

between the toes is a minute, but quite distinct, rounded papilla.

The toes are short, slightly decurved and emarginate on their

inner edges, which gives them the appearance of being clawed;

their length is about one twentieth of the length of the body.

The dorsal antenna is a minute setigerous papilla in the normal

position ; the lateral antennse have not been observed.

The corona extends down on the ventral side somewhat more

than one fourth of the length of the body. The auricles are small

and rounded; a fairly prominent rostrum is present and curves

down over the corona. The mouth is a little below the center.

The mastax is virgate with strongly asymmetric trophi of a

simple type. The fulcrum is very long and slender, tapering

gradually towards the posterior end, which is expanded into an

oval plate for the attachment of the muscles of the piston. The

rami are roughly triangular in ventral view; the dorsal branch

forms an acute angle with the ventral portion. The alula of the

left ramus is nearly as long as the ventral part of the ramus itself

and forms a very acute angle with the fulcrum. The right alula

is smaller and somewhat more divergent; it is excavate dorsally

and the sinus is reenforced by a strong rib, easily seen in the ven¬

tral view. The unci have a strong ventral tooth, followed by a

much smaller second tooth. The manubria are rodlike and formed

entirely by the median cell, only a slight ventral hump indicating

the normal posterior limit of the ventral cell. The right manu¬

brium is only two thirds as long as the left, and both are slightly

expanded at the posterior end. The piston is very large and fills

the entire cavity of the mastax. Two salivary glands are present;

the left gland is very large and curves under the mastax, while

the right gland is rudimentary and apparently not functional.

The oesophagus is long and slender. The gastric glands are

very small and rounded. There is no distinct separation between

stomach and intestine. The bladder is formed by an expansion of

the cloaca. The foot glands are large and slightly club-shaped.

The ovary is normal.

The ganglion is large and elongate saccate. The retrocerebral

organ consists of a small, pyriform sac, usually opaque with bac-

teroids; no subcerebral glands are present. The eyespot is a

granular mass of red pigment at the posterior end of the ganglion.

Total length 150-200/>t toes 8-10/x trophi 28/u,.

Harring (& Myers — Rotifer Fauna of Wisconsin — II. 457

Taphrocampa clavigera' was collected by Stokes at Trenton, New

Jersey; our material is from Bargaintown, near Atlantic City,

where it occurs occasionally in small numbers. While without the

pronounced, permanent annulations of T. annulosa and selenura,

it is evidently closely related to these two species; the mastax

differs only in details from that of T. annulosa.

Genus DBILOPHAGA Vejdovsky.

Notommatid rotifers with elongate, slender, spindle-shaped,

illoricate body, usually without distinct separation of head and

body and with very flexible integument; the head is cylindric and

elongate, with the mouth at a considerable distance from the front ;

the tail is rudimentary and the foot very short and apparently

unjointed; the toes are minute and conical.

The corona is reduced to a simple, circumapieal ring of cilia.

The mastax is virgate and the trophi of very simple form, usually

protrusile and adapted as organs of attachment to the body of the

host; rami large and strongly curved, without denticulation on

the inner margin; mallei slender and rodshaped, unci reduced

to small, oval plates; salivary glands very large.

Neither retrocerebral organ nor eyespot are present.

At least two of the species are ectoparasitic on oligochaete

worms; the third species appears to be free-living.

Type of the genus. — Brilophaga Bucephalus Vejdovsky.

This genus includes, in addition to D. judayi, described in vol¬

ume twenty, two other species, listed below. Neither has been

studied by anybody but the original discoverer.

Brilophaga bucephalus Vejdovsky, Sitzungsber. Bohm. Ges. Wiss. Prag (for

1882), 1883, p. 390, pi. 1, figs. 1-8. — De Beauchamp, Bull. Soc. Zool. France,

vol. 29, 1904, p. 157, figs. A, B.

Brilophaga delagei De Beauchamp, Bull. Soc. Zool. France, vol. 29, 1904, p.

159, fig. C; vol. 30, 1905, p. 121, fig. 3.

Genus PLEUROTROCHA Ehrenberg.

Notommatid rotifers with short, stout, illoricate, ovoid or globose

body, with a distinct neck separating the head and abdomen; the

foot is long and cylindric or slightly tapering; the toes are very

short and conical and may be either separate or fused.

458 Wisconsin Academy of Sciences, Arts, and Letters.

The corona is slightly oblique and consists of a marginal wreath

of cilia with lateral, auricle-like tufts of long cilia adapted to

swimming ; the apical plate is unciliated and the buccal field evenly

covered with short, close-set cilia; the mouth is near the ventral

edge of the corona.

The mastax is virgate and the trophi very simple; the fulcrum

is long and rodlike, the rami triangular, curved and not denticulate ;

the manubria are very long and the basal plate much reduced;

the unci are feeble and have only a single distinct tooth; the

piston is very large.

The eyespot is single and at the posterior end or on the lower

surface of the ganglion; there is no trace of the retrocerebral

organ.

Type of the genns. — Pleurotrocha petromyzon Ehrenberg.

This genus was created by Ehrenberg for P. petromyzon, which

he supposed to be without an eyespot. When the error was dis¬

covered, he transferred this species to his ‘ ‘ Noah ’s Ark, ’ ’ the genus

Notommata, into which he thrust all illoricate, freeswimming roti¬

fers with cervical eyespot. He then proceeded, in accordance with

his usual custom, to use the generic name for other species with¬

out eyespots and subsequent authors have followed him in this

change of concept. As P. petromyzon does not belong to Notom¬

mata as now understood, the original generic name must be used

for it, and the obscure species later referred to Pleurotrocha, listed

below, must be placed on a firm foundation or dropped as insuffi¬

ciently described.

Pleurotrocha aurita Bergendal, Acta Univ. Lundensis, vol. 28, 1892, 'sect. 2,

No. 4, p. 49, pi. 2, fig. 15.

Pleurotrocha constricta Ehrenberg, Abh. Akad. Wiss. Berlin (for 1831),

1832, p. 129; Infunsionsthierchen, 1838, p. 419, pi. 48, fig. 1. — Hudson and

Gosse, Kotifera, 1886, vol. 2, p. 19, pi. 18, fig. 3. — Von Hofsten, Arkiv

Zool., Stockholm, vol. 6, No. 1, 1909, p. 12.

Pleurotrocha gibba (Ehrenberg).

Eydatina gibba Ehrenberg, Abh. Akad. Wiss. Berlin (for 1831), 1832,

p. 127.

Pleurotrocha gibba Ehrenberg, Infusionsthierchen, 1838, p. 418, pi. 47, fig.

4. — Hudson and Gosse, Rotifera, 1886, vol. 2, p. 20, pi. 18, fig. 5.

Theora gibba Eyferth, Mikroskopische Siisswasserbewohner, 1877, p. 51;

Einfachsten Lebensformen, 1878, p. 83; 1885, p. 108.

Pleurotrocha renalis Ehrenberg, Monatsber. Akad. Wiss. Berlin, 1840, p. 218.

Earring & Myers — Rotifer Fauna of Wisconsin — II, 459

Pleurotrocha truncata Gosse.

Pleurotrocha truncata Gosse, Ann. Mag. Nat. Hist., ser. 2, vol. 8, 1851,

p. 199.

Theora truncata Eckstein, Zeitschr. Wiss. Zool., vol. 39, 1883, p. 372.

FLEUEOTBOCHA PETBOMYZON Ebreuberg.

Plate XXV, figures 1-4.

Pleurotrocha petromyBon Ehrenberg, Abh. Akad. Wiss. Berlin, 1830, p. 46. —

Von Hofsten, Arkiv Zool., Stockholm, vol. 6, No. 1, 1909, p. 12.

^ Notommata gihha Ehrenberg, Abh. Akad. Wiss. Berlin (for 1831), 1832,

p. 132, pi. 4, fig. 15; Infusionsthierchen, 1838, p. 430, pi. 52, fig. 4.

Notommata petromyzon Ehrenberg, Infusionsthierchen, 1838, p. 427, pi. 50,

fig. 7. — Gosse, Phil. Trans. Eoyal Soc. London, vol. 146, 1856, p. 432, pi. 17,

figs. 27-31.

Proales petromyzon Hudson and Gosse, Eotifera, 1886, vol. 2, p. 37, pi. 18,

fig. 9. — Kertesz, Budapest Eotat. Eaun., 1894, p. 30. — Hood, Proc. Eoyal

Irish Acad., ser. 3, vol. 3, 1895, p. 680. — Skorikov, Trav. Soc. Nat. Khar-

kow, vol. 30, 1896, p. 291. — Weber, Eev. Suisse Zool., vol. 5, 1898, p. 469,

pi. 18, figs. 21-23. — Voigt, Eorschungsber. Biol. Stat. Plon, vol. 11, 1904,

p. 42; Susswasserfauna Deutschlands, pt. 14, 1912, p. 90, fig. 160. —

Be Beauchamp, Arch. Zool. Exper., ser. 4, vol. 6, 1907, p. 9, fig. 5. — ^Lie-

Pettersen, Bergens Mus. Aarbog (for 1909), 1910, No. 15, p. 43. — Mola,

Ann. Biol. Lac., vol. 6, 1913, p. 243. — Montet, Eev. Suisse Zool., vol. 23,

1915, p. 322. — Jakubski, Eozpr. Wiad. Muz. Dzieduszyckich, vol. 1, No.

3-4, 1915, p. 16. — ^Weber and Montet, Cat. Invert. Suisse, pt. 11, 1918,

p. 99.

Notops laurentinus Jennings, Bull. Michigan Fish Comm., No. 3, 1894, p. 12,

figs. 3, 4.

Proales laurentinus Jennings, Bull. Michigan Pish Comm., No. 6, 1896, p. 91.

Pleurotrocha laurentina Harring, Bull. 81 U. S. Nat. Mus., 1913, p. 85.

The body is short, stout and gibbous ; its greatest width is about

one third of the entire length. The integument is soft and very

flexible, but the general outline is nevertheless quite constant. The

body is very transparent.

The head and abdomen are separated by a rather shallow con¬

striction. The head is small and short and somewhat oblique an¬

teriorly. The abdomen is pyriform and tapers posteriorly to the

base of the foot; there is no tail. The foot is long, cylindric and

two-jointed ; the length of the basal joint is equal to its width and

the terminal joint is twice the length of the basal. The toes are

very short, conical, acutely pointed and very slightly recurved at

the tips; their length is about one twentieth of the length of the

body.

The antennae are minute setigerous papillae, the dorsal in the

normal position, the lateral just beyond mid-length.

460 Wisconsin Academy of Sciences, Arts, and Letters.

The corona is frontal and consists of a circumapical band of

cilia with two strongly developed lateral, auricle-like tufts of cilia

adapted to swimming; the buccal field is evenly ciliated and the

mouth is near the ventral edge of the corona.

The mastax is virgate and the trophi of very simple form. The

rami are approximately triangular, strongly curved longitudinally

and have large, rounded alulae. The fulcrum is a very long, slen¬

der, tapering rod, incurved and slightly expanded at the posterior

end. The unci are triangular plates with one weak ventral and

a rudimentary second tooth; additional teeth are represented by

three faint striae. The manubria are long and double-curved,

tapering gradually from the broad base to the posterior end; near

mid-length there is a small projecting lobe on the ventral edge;

their longitudinal direction is nearly at right angles to the fulcrum.

Two small curved rods are imbedded in the anterior walls of the

mastax and serve to support the edges of the mouth during the

pumping action.

The oesophagus is short and slender. The gastric glands are

elongate oval and strongly compressed laterally. There is no con¬

striction separating the stomach and intestine. The ovary is

large and of a somewhat irregularly oval outline. A small bladder

is present. The foot glands are very long and nearly cylindric;

they discharge into a large mucus reservoir at the base of the toes.

The ganglion is large and saccate, reaching nearly to the pos¬

terior end of the mastax. No retrocerebral organ is present. The

eyespot is at the extreme end of the ganglion and consists of a

very small sphere of minute pigment granules.

Total length 225-250yLt; toes 12/x; trophi 32/x.

Pleurotrocha petromyzon is cosmopolitan in its distribution, but

does not usually occur in large numbers. It has been reported as

attaching itself to other living organisms (Infusoria, Entomos-

traca) ; we have not observed it in this condition and do not know

under what circumstances it occurs. It is evidently not a case of

parasitism, in the proper sense of this word, but rather one of

synoecia or “ Raumparasitismus ” ; the rotifer obtains free trans¬

portation and probably nothing more. The highly developed foot

glands point to the possibility of an adaptation of this kind ; they

are found in all rotifers that are known to be synoecious or pseudo-

parasitic.

Earring & Myers — Rotifer Fauna of Wisconsin — II. 461

PLEUROTEOCHA KOBUSTA (Glasscott) .

Plate XXV, figures 5-8.

Microcodon robustus Glasscott, Proc. Eoyal Dublin Soc., new ser., vol. 8,

1893, p. 40, pi. 3, fig. 2.

Microcodides robustus Eousselet, Journ. Quekett Micr. Club, ser. 2, vol. 6,

1895, p. 121, pi. 6, fig. 1.

? Microcodides abbreviatus Stenroos, Acta Soc. Fauna et Flora Fennica, vol.

17, No. 1, 1898, p. 113, pi. 1, fig. 20.

Mikrocodides robustus Harking, Bull. 81 IT. S. Nat. Mus., 1913, p. 71.

The body is short, stout and gibbous ; its greatest width is about

two fifths of the entire length. The integument is leathery and

the outline remains quite constant. The entire body is very trans¬

parent.

The head and abdomen are separated by a deep constriction.

The head segment is short and very broad ; its width is about two

thirds of the greatest width of the body. The abdomen is globose

and ends in a short, sleeve-like tail surrounding the base of the

foot. On the posterior half of the abdomen are five or six well

marked longitudinal folds; they are continuous from side to side

and not interrupted posteriorly. The foot is long, stout and

slightly tapering; it has three joints of approximately equal

length. The toe is single and abruptly reduced near the middle;

its length is a little less than one tenth of the total length of the

animal. As there are two well developed foot glands it is evident

that the single toe is a result of the fusion of two originally sep¬

arate toes.

The dorsal antenna is a large, knoblike elevation on the pos¬

terior part of the head, immediately in front of the neck; it has a

funnel-shaped central depression with a small tuft of sensory

setae. The lateral antennae are minute tubules with a few very

short setae in the normal position.

The corona is oblique and consists of a circumapical band of

cilia with two lateral, auricle-like tufts of strong cilia adapted to

swimming ; the buccal field is evenly ciliated. The mouth is in the

normal position, near the ventral margin of the corona, and not

in the center, as stated by Eousselet.

The mastax is virgate and of very simple form. The fulcrum

is very long, slender and tapering; the posterior end is slightly

expanded for the attachment of the muscles of the piston. The

rami are roughly triangular and without teeth or denticulations ;

a somewhat abrupt bend divides them into a rather short ventral

462

Wisconsin Academy of Sciences, Arts, and Letters.

and a long dorsal section. The alulae are very prominent. The

unci have only one tooth, slender and clubbed at the tip ; there is

no basal plate. The manubria are very long and their general

direction is nearly at right angles to the fulcrum, so that they

almost reach the dorsal side ; the basal plate is small and its edges

are parallel to the strongly curved anterior end of the principal

rib; the posterior branch is very slender and slightly recurved.

The piston is bulky, but not very powerful. The epipharynx con¬

sists of two slender, strongly curved rods, imbedded in the walls

of the mastax at the sides of the mouth.

The oesophagus is short and slender. There is no constriction

between stomach and intestine. The gastric glands are small and

nearly spherical. The ovary is irregularly elongate and reaches

from the bladder nearly to the mastax; the nuclei are large and

irregularly polygonal. A fairly large bladder is present. The

two foot glands are tapering and fully as long as the foot.

The ganglion is large and saccate. There is no trace of the

retrocerebral organ. The eyespot is well towards the front of the

head and seems to be seated on the mastax instead of the lower

surface of the ganglion, as it follows the movements of the mastax.

Total length 180-200/i; foot and toes 55-60/a; toes 16-18/*;

trophi 22/*.

Pleurotrocha rohusta is rather rare ; we have found it in Oneida

and Vilas Counties, Wisconsin, and in ponds and ditches around

Atlantic City, New Jersey. As this species has nothing in common

with Mikrocodides chlaena except the fused toes, it has been trans¬

ferred to Pleurotrocha, with which it seems to agree fairly well.

No good reasons ever existed for referring it to Mikrocodides; the

fused toes can not be considered a generic character to the exclu¬

sion of everything else. The supposed identity of the corona in

the two species is an error ; neither M. chlaena nor P. rohusta has

the mouth at the center of the corona ; as pointed out by De Beau¬

champ, M. chlaena has a corona of the type of Cyrtonia, and the

mastax is malleate.

Genus CEPHALODELLA Bory de St. Vincent.

Notommatid rotifers with prismatic or spindle-shaped, illoricate

or partly loricate body, having a slight constriction or neck sep¬

arating the head and abdomen and passing without definite limit to

the rudimentary foot, which is not jointed and has two slender toes.

Harring Myers — Rotifer Fauna of Wisconsin — 11. 463

The corona is an obliquely frontal disc with long marginal cilia

and two lateral tufts of densely set long cilia, especially adapted

to swimming ; the apical plate is enclosed by the marginal ciliation ;

the buccal field is sparsely ciliated. The mouth is slightly below

the center of the corona and the lips occasionally project as a

^‘beak.’’

The mastax is virgate ; the fulcrum is long and straight, nearly

always slightly expanded at the posterior end to provide a greater

surface for the attachment of the muscles of the piston ; the rami

are imperfectly developed, the uncus having only a single, slender

tooth and the manubrium usually rodshaped, with or without a

terminal crutch ; the piston is a large, powerful muscle attached to

the fulcrum and filling the entire cavity of the mastax.

The retrocerebral organ is absent in nearly all the species and

rudimentary when present, being limited to a small sac with par¬

tially atrophied duct, which does not reach the surface of the

corona. The eyespot may be cervical, frontal (single or double)

or absent.

Type of the genus. — Cephalodella catellina ( Muller )=06rcarm

catellina Muller.

The definition of this genus, which includes all the species of

Diaschiza Gosse, as revised by Dixon-Nuttall and Freeman, has

been broadened sufficiently to admit some evidently closely related

species. We have based it on the peculiar form of the body, a

head segment separated from the abdomen by a slight constric¬

tion, no distinct separation between the abdomen and foot, corona

slightly oblique with mouth near the center, and the specialized

type of the virgate mastax, which varies only in minute details

throughout the genus as here constituted. We do not consider

the divided lorica as a generic character ; in many species it is only

a polite fiction. Neither can the small tuft of setae at the base

of the toes be accepted as such; it is so difficult to find that we

have made no use of it ; although the genus includes a great many

species, they are readily distinguished by easily ascertainable

differences.

The ‘ ‘ beak ’ ’ must be observed in the living animal ; in preserved

material the lips nearly always project somewhat. We have not

figured the trophi of all the species included, as the differences

are slight, but enough have been given to show the range of varia¬

tion within the genus.

464 Wisconsin Academy of Sciences, Arts, and Letters.

The change of the familiar name Biaschiza is regrettable, but is

seemingly made unavoidable by the inclusion of Biglena catellina,

of Ehrenberg and Gosse. De Beauchamp* objects to identifying

Ehrenberg’s animal with Muller’s Cercaria catellina, reproducing

the figures of Muller and Weber. We are quite willing to admit

that Muller’s figure is, to say the least, poor, but Weber’s is not

a great deal better, especially if consideration is given to the im¬

provements made in optical instruments in the interval between

1786 and 1888. The usual custom in similar doubtful cases is to

abide by the choice of the “first reviser”, a distinction clearly

belonging to Ehrenberg. As he claims to have recognized Cercaria

catellina Muller, the simplest and most consistent procedure is to

accept the identification as correct. Denying it would not dispose

of the generic name Biglena; although this was originally defined

by a synonymic citation only, there is no doubt about the identity

of the animal for which Ehrenberg created the genus. His figure

in the Infusionsthierchen of 1838 is unmistakable and this species

must remain the type of Biglena, whether it retains the specific

name catellina (Muller) or the next available, probably granulans

Weisse, used for the male; for the reasons given above it seems

preferable to use catellina (Muller). However, this had already

been made the type of a genus Cephalodella by Bory de St. Vincent

in his compilation of 1826, which under the circumstances may be

considered fortunate, as it obviates the displacement of the name

Biaschiza by Biglena Ehrenberg, which has long been used for

the forcipate Notommatids. Such transpositions are very confus¬

ing, even when absolutely unavoidable.

Biaschiza Gosse is not tenable under any circumstances; Du jar-

din created a genus Plagiognatha in 1841, designating as type P.

felis, in his opinion identical with Muller’s Vorticella felis, but his

figure shows beyond reasonable doubt that the animal he actually

studied was Ehrenberg’s Furcularia gihha, more familiar as

Biaschiza gihba. Consequently, as Biaschiza can not possibly be

retained, the least objectionable solution appears to be the resur¬

rection of Cephalodella Bory de St. Vincent, a course already sug¬

gested by Eyferth (Einfachsten Lebensformen, 1878, p. 83,

Biglena) :

“ . . . D. catelliria, die gemeinste von alien, ist jedenfalls abzutrennen.

In der Form des Kauers und der Derbheit der Cutieula steht sie (wie Notom-

Soc. Zool. France, vol. 38, 1914, p. 291 ff.

Earring & Myers — Rotifer Fauna of Wisconsin — 11. 465

mata lacinulata) der Furcularia naher, von der sie aber durch den sehr knrzen

Fuss zu stark abweicht. Am besten wiirde sie unter dem alten Namen Ceph-

alodella Bory als besondere Gattung abzusondern sein. ’ ’

The three species listed below evidently belong to this genus.

They have not occurred in our collections and all appear to be

rare; C. crassipes and C. leptodactyla were described from single

specimens.

CEFHALODELLA CEASSIPES (Lord)

Diaschiza crassipes Lord, Trans. Manchester Micr. Soc. (for 1903), 1904,

p. 78, pi. 3, fig. 3.

CEFHALODELLA DERBYI (Dixon-Nuttall and Freeman).

Diaschwa derbyi Dixon-Nuttall and Freeman, Journ. Eoyal Micr. Soc.,

1903, p. 131, pi. 4, fig. 13.

CEFHALODELLA LEPTODACTYLA (Hauer).

Furcularia leptodactyla Hauer, Mitt. Bad. Landesver. Naturk., Freiburg i.

Br., new ser., vol. 1, 1921, p. 183; Arch. Hydrobiol., vol. 13, 1922, p. 693,

pi. 9, fig. 1.

CEFHALODELLA CATELLINA (Muller).

Plate XXVII, figures 3-5.

Cercaria catellina Muller, Anim. Infus., 1786, p. 130, pi. 20, figs. 12, 13.

Vorticella larva Muller, Anim. Infus., 1786, p. 286, pi. 40, figs. 1-3.

Furcocerca catellina Lamarck, Hist. Nat. Anim. sans Vert., vol. 1, 1815,

p. 448.

Furcularia larva Lamarck, Hist. Nat. Anim. sans Vert., vol. 2, 1816, p. 37.

CepJialodella catellina Bory de St. Vincent, Class. Anim. Micr., 1826, p. 43.

— Harring, Bull. 81 U. S. Nat. Mus., 1913, p. 24. — Kozar, Zool. Anz.,

vol. 44, 1914, p. 416.

Dicranophorus catellinus Nitzsch, Enc. Wiss. u. Kiinste, sect. 1, vol. 16,

1827, p. 68.

Diglena catellina Ehrenberg, Abh. Akad. Wiss. Berlin (for 1829), 1830,

p. 8; (for 1831), 1832, p. 137, pi. 4, fig. 17; Infusionsthierchen, 1838,

p. 444, pi. 55, fig. 3. — Bartsch, Jahresh. Ver. Naturk. Wiirttemberg,

vol. 26, 1870, p. 39. — Daday, Erdelyi Muz.-Egyl. Evkon., new ser., vol.

2, pp. 198, 217, pi. 9, figs. 1-3. — Eckstein, Zeitschr. Wiss. Zool., vol. 39,

1883, p. 371, pi. 26, figs. 40, 41. — Hudson and Gosse, Eotifera, 1886, vol.

2, p. 53, pi. 19, fig. 10. — ^Weber, Arch. Biol., Liege, vol. 8, 1888, p. 46,

pi. 34, figs. 1-6; Eev. Suisse Zool., vol. 5, 1898, p. 492, pi. 19, figs. 12-14.

— Wierzejski, Eozpr. Akad. Umiej., Wydz. Mat.-Przyr., Krakow, ser. 2,

vol. 6, 1893, p. 232. — Kertbsz, Budapest Eotat. Faun., 1894, p. 31. —

Voigt, Forschungsber. Biol. Stat. Plon, vol. 11, 1904, p. 48. — De Beau¬

champ, Arch. Zool. Exper., ser. 4, vol. 10, 1909, p. 280; Bull. Soc. Zool.

France, vol. 38, 1914, p. 297, fig. 1. — Lucks, Eotatorienfauna Westpreus-

sens, 1912, p 56. — Sachse, Siisswasserfauna Deutschlands, pt. 14, 1912,

466 Wisconsin Academy of Sciences^ Arts, and Letters.

p. 106, fig. 199. — Mola, Ann. Biol. Lae., vol. 6, 1913, p. 246. — ^Montet,

Eev. Suisse Zool., vol. 23, 1915, p. 329. — Jakubski, Eozpr. Wind. Muz.

Dzieduszyckicli, vol. 1, No, 3-4, 1915, p. 19. — ^Whitney, Journ. Exper.

Zool., vol. 20, 1916, p. 274, figs. 4, 5.

? Leiodina capitata Moreen, Ann. Sci. Nat., vol. 21, 1830, p, 124, pi, 3,

fig. 2; Bijdr. Natuurk. Wetensch., vol. 5, 1830, p. 211, fig.

Furcularia catellina Blainville, Diet. Sei. Nat., vol. 60, 1830, p. 152.

Typhlina furca Ehrenberg, in Hemprieh and Ehrenberg, Symb. Phys. Anim,

Evert., 1831 (1832?), pi. 1, figs. 17 b, 2, 3; not Cercaria furca Muller.

Flagiognatha catellina Dujardin, Hist. Nat. Zooph., Inf., 1841, p. 652.

Flagiognatha hyptopus Dujardin, Hist. Nat. Zooph., Inf., 1841, p. 653, pi.

21, fig. 8; not Notommata hyptopus Ehrenberg.

Diglena granularis Weisse, Bull. Phys.-Math. Aead. St. Petersburg, vol. 8,

1849, eol. 300.

f Heterognathus diglenus Schmarda, Neue wirbellose Thiere, vol. 1, 1859,

p. 52, pi. 12, fig. 107.

F Notops forcipita Glasscott, Proe. Eoyal Dublin Soe., new ser., vol. 8,

1893, p. 79, pi. 6, fig. 5.

? Froales algicola Kellicott, Trans. Amer. Mier. Soe., vol. 19, 1897, p. 48.

F Biaschiza tenuior Murray, Brit. Antaretie Exped. 1907-9. Eep. Sei. Inv.,

vol. 1, 1910, p. 57, pi. 13, fig. 16.

F Diglena volvocicola Zavadovski, IJehen. Zap. Moskovsk. Gor. Univ. Shan-

iavskago, vol. 1, 1916, p. 246, pi. 3, text figs. 1-19.

F Diglena catellina parasitica Zavadovski, Uchen. Zap. Moskovsk. Gor. TJniv.

Shaniavskago, vol. 1, pt. 2, 1916, App., p. 4.

Diaschiza catellina Weber and Montet, Cat. Invert. Suisse, pt. 11, 1918,

p. 143.

The body is short, stout and strongly gibbous dorsally. The head

is very large and oblique anteriorly. The neck is not very strongly

marked. The abdomen is strongly compressed laterally ; the ventral

edge is nearly straight, the dorsal gently curved for about two

thirds of its length and the posterior third rounded. The lorica is

rather flexible, and the plates somewhat indistinct; the lateral

clefts are wide and parallel-sided. The small, short foot is wholly

ventral and the posterior portion of the abdomen projects over

and beyond it as a huge tail. The toes are short, almost straight

and taper gradually to acute points ; when closely appressed to the

body, they project very little beyond the tail; their length is about

one fifth of the total length.

The corona is very strongly oblique and distinctly convex with¬

out projecting lips.

The mastax is very large and the trophi of the normal type;

the fulcrum is very long and slightly expanded at the posterior

end ; the manubria are rodshaped and decurved, ending in a semi¬

circular, dorsal expansion.

Harring (S; Myers — Rotifer Fauna of Wisconsin — II. 467

The ganglion is very large and saccate. The retrocerebral organ

is absent and the eyespot frontal and double, consisting of two

closely approximated spheres of red pigment.

Total length 105 — 110/x; toes 18 — 20/x; trophi 45ja.

CepJialodella catellina is apparently of worldwide, but some¬

what erratic distribution ; this is probably to be attributed to un¬

usually narrow limits of the conditions necessary for its existence ;

when these are satisfied, it will be found in great profusion.

Diglena volvocicola Zavadovski is at least very closely related

to C. catellina and probably identical with it ; the figures given by

the author do not show any differences that might be considered

of specific value. Some physiological dissimilarities are described,

the most striking one being the parasitism of the animal in Volvox

colonies. The author suggests the alternative name catellina para¬

sitica for this form, if it should prove to be only a variety.

CEPHALODULLA ANGUSTA Myers, new species.

Plate XXVII, figure 2.

The body is small, stout and gibbous dorsally. The head is very

large, slightly deflexed, and strongly oblique anteriorly. The neck

is indistinctly marked. The abdomen increases slightly in width

for about two thirds of its length; the posterior third is rounded

dorsally. The lorica is very flexible, but the plates well marked;

the lateral clefts are fairly wide anteriorly and the edges diverge

slightly and gradually towards the posterior end. The foot is small

and conical with a minute tail somewhat beyond mid-length. The

toes are very short, slender and slightly recurved, tapering grad¬

ually to acute points ; their length is one sixth of the total length.

The corona is strongly oblique and convex without projecting lips.

The mastax is large and of the normal type; the fulcrum is

slightly ' expanded posteriorly, the manubria rodlike with ends

strongly decurved, but not crutched. The gastric glands are small.

The ganglion is very long and pyriform. The eyespot is frontal

and double, the two halves very close together; the retrocerebral

organ is absent.

Total length 90-95/x; toes 15-18jit.

Cephalodella angusta is not common; we have collected it only

in a large pond at Oceanville, New Jersey, among Biccia and float¬

ing sphagnum in soft, acid water.

468 Wisconsin Academy of Sciences, Arts, and Letters,

CEFHALODELLA EPITEDIA Myers, new species.

Plate XXVII, figure 7.

The body is fairly slender, laterally compressed and slightly

gibbous dorsally. The head is long and convex anteriorly. The

neck is somewhat indistinct. The abdomen increases gradually

in width for about two thirds of its length; the posterior third is

gently rounded. The integument is very flexible and the plates

ill-deflned; the lateral clefts are very obscure, but apparently

parallel-sided and rather narrow. The foot is conical and rather

narrow at the base ; the small tail is near the posterior end. The

toes are short, straight and slender, nearly parallel-sided for about

three fourths of their length and somewhat abruptly reduced to

acute, slightly recurved, clawlike points; their length is less than

one fifth of the total length. The foot glands are very small and

pyriform.

The corona is moderately oblique and strongly convex without

projecting lips.

The mastax is large and of the typical form; the fulcrum is

slender and slightly expanded at the extreme posterior end, the

manubria very slender, rodlike and not crutched. The gastric

glands are small and rounded.

The ganglion is very long and saccate; the retrocerebral organ

is absent and the eyespot frontal and double, the two small pig¬

ment spheres fairly wide apart.

Total length 135-140/>t; toes 24-26/x.

Cephalodella epitedia is found among algae and detritus in

brackish and saltwater ditches near Atlantic City, New Jersey. It

resembles C. angusta and gracilis in general form, but differs in

the shape of the toes and in never being found in fresh water, to

which these two species appear to be confined.

. CEPHALODELLA PAXILLA Myers, new species.

Plate XXVI, figure 6.

The body is elongate, slender, cylindric and very nearly parallel¬

sided. The head is large and slightly oblique anteriorly. The neck

is not very strongly marked. The abdomen is cylindric for nearly

its entire length, abruptly rounded at the extreme posterior end;

the integument is thin and flexible and the plates indistinct; the

lateral clefts are fairly wide anteriorly and the edges diverge very

Harring & Myers — Rotifer Fauna of Wisconsin — II. 469

slightly and gradually towards the extreme posterior end, which

is oblique and flaring. The foot is relatively small, conical and

longer on the ventral side than on the dorsal ; the tail is small and

near mid-length. The toes are short and nearly straight, the an¬

terior half cylindric and the posterior tapering, ending in acute

points; the dorsal edge is very slightly decurved, the ventral

straight for half its length, converging gradually towards the dorsal

margin ; their length is less than one fifth of the total length. The

foot glands are small and pyriform.

The corona is only slightly oblique, convex and without project¬

ing lips.

The mastax is large and of the typical form, but the trophi are

delicate; the fulcrum is a long, straight, very slender rod, not

expanded posteriorly, the manubria not crutched. The gastric

glands are small and rounded.

The ganglion is moderately long and saccate; the retrocerebral

organ is absent. The eyespot is frontal and double, the two spheres

fairly wide apart.

Total length 210-220/>i; toes 36-40jU,.

Cephalodella paxilla is rare ; we have collected it among floating

sphagnum in a soft, acid-water pond at Gravelly Run, near Mays

Landing, New Jersey.

CEPHALODELLA MARINA Myers, new species.

Plate XXVI, figure 7.

The body is moderately elongate, spindle-shaped, very slightly

compressed laterally and faintly gibbous dorsally. The head is

large, slightly deflexed and oblique anteriorly. The neck is some¬

what indistinct. The abdomen is slightly convex dorsally and

deepest near mid-length ; the lorica is very thin and flexible and the

plates ill-defined; the lateral clefts are narrow anteriorly and in¬

crease gradually in width towards the posterior end. The foot is

fairly long, stout and conical ; the small tail is a little beyond mid¬

length. The toes are short, very slender, slightly decurved and

taper gradually to very acute points ; their length is somewhat less

than one fifth of the total length. The foot glands are rather small

and elongate ovate.

The corona is decidedly oblique and somewhat convex without

projecting lips.

470 Wisconsin Academy of Sciences, Arts, and Letters.

The mastax is fairly large and of the normal type ; the fulcrum

is sKghtly expanded posteriorly, the manubria slender, rodlike

and not crutched. The gastric glands are small.

The ganglion is elongate and saccate ; the eyespot is frontal and

double, the two spheres rather wide apart. No retrocerebral organ

is present. The dorsal antenna is unusually far back on the head.

Total length 160-165/a; toes 28-30/a.

Cephalodella marina is common among algae and vegetable

detritus in shallow tidepools near Atlantic City, New Jersey. It

is related to C. mineri, but is readily distinguished by the very

slender toes and the uncrutched manubria, the more oblique corona

and larger head.

CEPHALODELLA INNESI Myers, new species.

Plate XXVI, figures 3-5.

The body is elongate, slightly compressed laterally and strongly

gibbous dorsally. The head is small, distinctly deflexed and

strongly oblique anteriorly. The neck is well marked. The ab¬

domen increases rapidly in width for about two thirds of its length ;

at this point the dorso-ventral depth is almost twice the depth of

the anterior margin; the posterior third of the dorsal edge curves

slightly downward. The lorica is flexible, but the plates are fairly

distinct; the lateral clefts are narrow anteriorly and increase

slightly in width towards the posterior end. The foot is long,

conical and extremely broad at the base ; the tail is small and some¬

what beyond mid-length. The toes are relatively short and robust,

very slightly enlarged at the base and taper gradually to slender,

acute points; their length is about one fifth of the total length.

The foot glands are very large and pyriform.

The corona is strongly oblique and convex without projecting lips.

The mastax is fairly large and of the typical form ; the fulcrum

is relatively short, stout and parallel-sided without any expansion

at the posterior end; the manubria are long and crutched. The

gastric glands are small.

The ganglion is moderately long and saccate; the eyespot is

frontal and double, the two halves fairly wide apart. No retro-

cerebral organ is present. The dorsal antenna is far back on the

head, a short distance in front of the neck.

Total length 200-210/a ; toes 40-44/a ; trophi 40/a.

Earring & Myers — Rotifer Fauna of Wisconsin — II. 471

Cephalodella innesi has been collected around Atlantic City,

New Jersey, in weedy ponds with soft, acid water. It is readily

distinguished by the small head and gibbous body and also by its

rapid, restless mode of swimming.

CEPHALODELLA MINERI Myers, new species.

Plate XXVI, figure 1.

The body is rather short and nearly cylindric, slightly gibbous

dorsally. The head is moderately large and somewhat deflexed.

The neck is not very distinctly marked. The abdomen is nearly

parallel-sided, slightly convex dorsally ; the lorica is very thin and

flexible, but the plates are fairly well defined; the lateral clefts

are very narrow and parallel-sided, slightly flaring at the extreme

posterior end. The foot is relatively short, conical and robust;

the very small tail is a little beyond mid-length. The toes are

short, strongly decurved, extremely broad at the base and taper

gradually and regularly to acute points ; their length is about one

fifth of the total length. The foot glands are large and ovate.

The corona is slightly oblique and convex without projecting lips.

The mastax is fairly large and of the typical form ; the fulcrum

is rather stout and slightly expanded posteriorly, the manubria

long, very slender and crutched. The gastric glands are small.

The ganglion is elongate and saccate; the eyespot is double

and frontal, the two pigment spheres fairly wide apart, and the

retrocerebral organ absent.

Total length 125/a; toes 27/a.

Cephalodella mineri is very abundant among Fontinalis nova-

angliae in a brackish stream near Tuckahoe, Atlantic County,

New Jersey.

CEPHALODELLA ELONGATA Myers, new species.

Plate XXVI, figure 2.

The body is elongate, cylindric and extremely slender. The head

is unusually long and convex anteriorly. The neck is indistinct.

The abdomen is cylindric throughout its length ; the lorica is very

thin and flexible and the plates ill-defined; the lateral clefts are

very obscure, but apparently very narrow and parallel-sided. The

foot is short and conical; the small tail is near the posterior end.

The toes are short, slightly decurved, and taper gradually and

472 Wisco7isin Academy of Sciences, Arts, and Letters.

evenly to very acute points. The foot glands are very small and

pyriform.

The corona is only slightly oblique, but strongly convex and

without projecting lips.

The mastax is moderately large and of the typical form; the

fulcrum is slender and slightly recurved at the posterior end ; the

manubria are crutched. The gastric glands are elongate pyriform,

the obtuse end forward.

The ganglion is very long and saccate; the retrocerebral organ

is absent and the eyespot frontal and double, the two pigment

spheres fairly wide apart.

Total length 105-115/a; toes 21-25/a.

Cephalodella elongata is not common; we find it occasionally in

weedy ponds and sphagnum bogs around Atlantic City, New Jersey.

It is readily recognized by the very slender body, as well as the

pyriform gastric glands.

CEPHALODELLA GIBBA (Ehrenljerg) .

Plate XXX, figures 4-6.

Furcularia gib'ba Ehrenberg, Abh. Akad. Wiss. Berlin (for 1831), 1832,

p. 130, pi. 4, fig. 16; Infusionsthierchen, 1838, p. 420, pi. 48, fig. 3. —

Gosse, Phil. Trans. Eoyal Soe. London, vol. 146, 1856, p. 433, pi. 17, figs.

35-37. — Eckstein, Zeitschr. Wiss. Zool., vol. 39, 1883, p. 374. — Hudson

and Gosse, Eotifera, 1886, vol. 2, p. 43, pi. 19, fig. 13. — Tessin, Arch.

Naturg. Mecklenburg, vol. 43, 1890, p. 150. — ^Wierzejski, Eozpr. Akad.

Umiej., Wydz. Mat.-Przyr., Krakow, ser. 2, vol. 6, 1893, p. 230. —

Kertesz, Budapest Eotat. Faun., 1894, p. 31. — Skorikov, Trav. Soc. Nat.

Kharkow, vol. 30, 1896, p. 294. — Stenroos, Acta Soc. Fauna et Flora

Fennica, vol. 17, No. 1, 1898, p. 133.

Plagiognatha felis Dujardin, Hist. Nat. Zooph., Inf., 1841, p. 652, pi. 18,

fig. 3; not Vorticella felis Muller.

BiascMza semiaperta Gosse, in Hudson and Gosse, Eotifera, 1886, vol. 2,

p. 80, pi. 22, fig. 10. — Bilfinger, Jahresh. Ver. Naturk. Wiirttemberg,

vol. 48, 1892, p. 116; ibid., vol. 50, 1894, p. 54. — Wierzejski, Eozpr.

Akad. Umiej., Wydz. Mat.-Przyr., Krakow, ser. 2, vol. 6, 1893, p. 237. —

Kertesz, Budapest Eotat. Faun., 1894, p. 35. — Levander, Acta Soc.

Fauna et Flora Fennica, vol. 12, No, 3, 1895, p. 44. — Stenroos, Acta Soc.

Fauna et Flora Fennica, 17, No. 1, 1898, p. 155. — Weber, Eev. Suisse

Zool., vol. 5, 1898, p. 551, pi. 21, figs. 1-3. — EunnstrOM, Zool. Anz., vol.

34, 1909, p. 270. — ^Lie-Pettersen, Bergens Mus. Aarbog (for 1909),

1910, No. 15, p. 59. — Mola, Ann. Biol. Lac., vol. 6, 1913, p. 255.

BiascMza gihba Dixon-Nuttall and Freeman, Journ. Royal Micr. Soc.,

1903, p. 6, pi. 1, fig. 1. — Voigt, Forschungsber. Biol. Stat. Plon, vol. 11,

1904, p. 63. — Voronkov, Trudy Hidrobiol. Slants. Glubokom Oz., vol. 2,

Harring & Myers — Rotifer Fauna of Wisconsin — II. 473

1907, p. 105. — De Beauchamp, Arch. Zool. Exper., ser. 4, vol. 10, 1909,

p. 203, fig. XXI A. — Von Hopsten, Arkiv Zool., Stockholm, vol. 6, No. 1,

1909, p. 50. — Lucks, Rotatorienfauna Westpreusseiis, 1912, p. 92. — ■

Sachse, Siisswasserfauna Deutschlands, pt. 14, 1912, p. 223, fig. 227. —

Montet, Rev. Suisse Zool., vol. 23, 1915, p. 330. — Jakubski, Rozpr. Wiad.

Muz. Dzieduszyckich, vol. 1, No. 3-4, 1915, p. 21. — ^Weber and Montet,

Cat. Invert Suisse, pt. 11, 1918, p. 135.

The body is moderately elongate, slightly compressed laterally

and gibbous dorsally. The head is large and oblique anteriorly.

The neck is well marked. The abdomen increases slightly in width

for about two thirds of its length and is rounded posteriorly; the

lorica is firm and the plates very distinct; the lateral clefts are

rather narrow anteriorly and increase gradually in width towards

the posterior end. The foot is small and conical; the small tail

is near mid-length. The toes are very long, straight or recurved

and very slender; the basal portion is broad and tapers rapidly

to the nearly parallel-sided, very slightly tapering main portion;

the extreme tips are conical. The length of the toes is about one

third of the total length.

The corona is oblique and strongly convex without projecting lips.

The mastax is very large and the trophi of the normal type.

The inner ventral edges of the rami are near the apex provided

with comblike, denticulate lamellae; the fulcrum is broadly ex¬

panded at the posterior end an'd the manubria are strongly

crutched; the unci have only a single tooth. The gastric glands

are rather small.

The ganglion is very long and saccate ; no retrocerebral organ

is present. The eyespot is frontal and consists of a spherical cap¬

sule, the anterior half transparent, the posterior filled with pigment

granules.

Total length 250-300/a; toes 70-80/a; trophi 60/a.

Cephalodella gihha is abundant in weedy ponds everywhere. It

is somewhat variable, especially in the curvature of the toes ; speci¬

mens with virtually straight toes are not rare and in the early

spring months they predominate over the curved toed variety.

CEPHALODELLA GRACILIS (Ehrenberg) .

Plate XXVII, figure 1.

Furcularia gracilis Ehrenberg, Abh. Akad. Wiss. Berlin (for 1831), 1832,

p. 130; Infusionsthiercken, 1838, p. 421, pi. 48, fig. 6. — Eckstein,

Zeitschr. Wiss. Zool., vol. 39, 1883, p. 374, pi. 26, fig. 43. — Hudson and

474 Wisconsin Academy of Sciences, Arts, and Letters.

Gosse, Eotifera, 1886, vol. 2, p. 42, pi. 19, fig. 14. — Wierzejski, Eozpr.

Akad. Umiej., Wydz. Mat.-Przyr., Krakow, ser. 2, vol. 6, 1893, p. 229. —

Skorikov, Trav. Soc. Nat. Kharkow, vol. 30, 1896, p. 294. — Stenroos,

Acta Soc. Fauna et Flora Fennica, vol. 17, No. 1, 1898, p. 133.

Biaschiza taurocepJialus tenua Hilgendorf, Trans. New Zealand Inst.,

vol. 31, 1899, p. 124, pi. 10, fig. 9c-d.

Diaschiza gracilis Dixon-Nuttall and Freeman, Journ. Eoyal Micr. Soc.,

^1903, p. 10, pi. 1, fig. 4. — Voigt, Forschungsber. Biol. Stat. Plon, vol. 11,

1904, p. 64. — Voronkov, Trudy Hidrobiol. Stants. Glubokom Oz., vol. 2,

1907, p. 105. — Von Hofsten, Arkiv Zool., Stockholm, vol. 6, No. 1, 1909,

p. 50. — Sachse, Siisswasserfauna Deutschlands, pt. 14, 1912, p. 119, fig.

228. — Barring, Proc. U. S. Nat. Mus., vol. 47, 1913, p. 528. — Montet,

Eev. Suisse Zool., vol. 23, 1915, p. 330. — Jakubski, Eozpr. Wiad. Muz.

Dzieduszyckich, vol. 1, No. 3-4, 1915, p. 21. — ^Weber and Montet, Cat.

Invert. Suisse, pt. 11, 1918, p. 137.

The body is rather short, laterally compressed and slightly

gibbous dorsally. The head is relatively short, broad and convex

anteriorly. The neck is well marked. The abdomen increases

gradually in width for about two thirds of its length ; the posterior

third is gently rounded; the lorica is thin and flexible, but the

plates are fairly distinct; the lateral clefts are rather narrow an¬

teriorly and increase slightly in width towards the posterior end.

The foot is conical and rather short; the very small tail is some¬

what beyond mid-length. The toes are short, fairly slender, very

slightly recurved and taper gradually and evenly to acute points;

their length is about one fifth of the total length. The foot glands

are moderately large and pyriform.

The corona is oblique and strongly convex without projecting lips.

The mastax is fairly large and of the normal type ; the fulcrum

is very slender and slightly recurved posteriorly, but not expanded ;

the manubria are slender, rodlike, decurved at the ends and not

crutched. The gastric glands are small.

The ganglion is elongate and saccate; the retrocerebral organ

is absent and the eyespot frontal.

Total length 125-130/;t ; toes 22-25/r.

Cephalodella gracilis is common everywhere in weedy ponds,

it has a certain resemblance to C. sterea, but is readily distinguished

by the form of the toes and its much smaller size and stouter body.

CEPHALODELLA STEREA (Gosse).

Plate XXVII, figure 6.

b'urcularia sterea Gosse, Journ. Eoyal Micr. Soc., 1887, p. 864, pi. 14, fig. 8.

— Hudson and Gosse, Eotifera, Suppl., 1889, p. 25, pi. 31, fig. 15. —

Harring & Myers — Rotifer Fauna of Wisconsin — II. 475

WiEiiZEjSKi, Eozpr. Akad. Umiej., Wydz. Mat.-Przyr., Krakow, ser. 2,

vol. 6, 1893, p. 230.

Diaschim sterea Dixon-Nuttall and Feeeman, Journ. Eoyal Micr. Soc.,

1903, p. 8, pi. 1, fig. 3. — Voronkov, Trudy Hidrobiol. Stants. Glubokom

Oz., vol. 2, 1907, p. 105. — ^Whitney, Journ. Exper. ZooL, vol. 20, 1916,

p. 273, figs. 3, 4. — Weber and Montet, Cat. Invert. Suisse, pt. 11, 1918,

p. 136, fig. 37.

The body xS moderately stout and slightly gibbous dorsally. The

head is large and strongly oblique anteriorly. The neck is well

marked. The abdomen increases gradually and regularly in width

for about three fourths of its length and diminishes rapidly from

this point to the base of the foot; the lorica is relatively firm and

the plates well marked; the lateral clefts are somewhat wider

than usual and nearly parallel-sided. The foot is large and robust ;

the tail is very prominent and extends very slightly beyond the end

of the foot. The toes are rather short, stout, minutely recurved

posteriorly and taper gradually to very acute points ; the dorsal and

ventral edges are very slightly undulate; their length is less than

one fourth of the total length. The foot glands are large and

pyriform.

The corona is strongly oblique and convex without projecting lips.

The mastax is fairly large and of the normal type; the fulcrum

is slightly expanded at the posterior end and the manubria strongly

crutched. The gastric glands are rather small.

The ganglion is very large and saccate; some specimens appear

to possess a rudiment of the retrocerebral sac at the posterior end

of the ganglion. The eyespot is frontal and consists of two hemi¬

spherical pigment masses within a single capsule.

Total length 180-190/a ; toes 40-45/a.

Cephalodella sterea is somewhat sporadically distributed; when

found, it is usually very abundant. It is readily distiniguished

by the large tail.

CEPHALODELLA GLOBATA (Gosse).

Plate XXXI, figure 3.

Diaschisa glolata Gosse, Journ. Eojal Micr. Soc., 1887, p. 362, pi. 8, fig. 4.

— Hudson and Gosse, Rotifera, Suppl., 1889, p. 37, pi. 31, fig. 30. —

Dixon-Nuttal and Freeman, Journ. Royal Micr. Soc., 1903, p. 7, pi. 3,

fig. 9.

Furcularia sphaerica Gosse, Journ, Royal Micr. Soc., 1887, p. 864, pi. 14,

fig. 7. — Hudson and Gosse, Rotifera, Suppl., 1889, p. 26, pi. 31, fig. 16.

476 Wisconsin Academy of Sciences, Arts, and Letters.

The body is short, stout and gibbous dorsally. The head is rather

short, very broad, slightly deflexed and oblique anteriorly. The

neck is not very strongly marked. The abdomen is short and very

broad ; the dorsal edge is strongly convex. The lorica is relatively

firm and the plates well marked ; the lateral clefts are narrow an¬

teriorly and increase very slightly in width towards the posterior

end. The foot is very short and broad; the small tail reaches

nearly to the posterior end. The toes are short, rather slender,

slightly decurved and taper gradually to acute points ; their length

is about one fifth of the total length. The foot glands are small

and pyriform.

The corona is oblique and somewhat convex without projecting

lips.

The mastax is large and of the normal type; the fulcrum is

slightly expanded at the posterior end and the manubria long and

strongly crutched. The gastric glands are small.

The ganglion is very long and saccate ; no retrocerebral organ is

present. The eyespot is frontal and consists of a somewhat diffuse

cluster of very pale pigment granules.

Total length 125-130/x ; toes 22-25/x.

Cephalodella glohata is very widely, but somewhat sporadically

distributed and is not often found in large numbers. It resembles

C. physalis, but the toes are different, the head is smaller and the

eye is frontal ; physalis has a cervical eyespot, is considerably larger

and has relatively longer toes.

CEPHALODELLA PORFICULA (Ehrenberg) .

Plate XXXIV, figures 1-3.

? Bistemma forficula Ehrenberg, Abh. Akad. Wiss, Berlin (for 1831), 1832,

p. 139; Infusionsthierehen, 1838, p. 449, pi. 56, fig. 2. — Hudson and

Gosse, Kotifera, 1886, vol. 2, p. 41; Suppl., 1889, p. 31, pi. 33, fig. 19.

Furcularia forficula Ehrenberg, Infusionsthierehen, 1838, p. 421, pi. 48,

fig. 5. — Bartsch, Jahresh. Ver. Naturk. Wiirrtemberg, vol. 26, 1870, p.

41. — Eckstein, Zeitsehr. Wiss. Zool., vol. 39, 1883, p. 375, pi. 26, fig. 44.

— Hudson and Gosse, Rotifera, 1886, vol. 2, p. 41, pi. 20, fig. 1. — Kelli-

COTT, Proc. Amer. Soc. Micr., vol. 10, 1888, p. 94. — Wierzejski, Eozpr.

Akad. Umiej., Wydz. Mat.-Przyr., Krakow, ser. 2, vol. 6, 1893, p. 229. —

Jennings, Bull. Michigan Pish Comm., No. 3, 1894, p. 17. — Kertbsz,

Budapest Rotat. Faun., 1894, p. 31. — Skorikov, Trav. Soc. Nat. Kharkow,

vol. 30, 1896, p. 293. — Weber, Rev. Suisse Zool., vol. 5, 1898, p. 473, pi.

19, figs. 1, 2. — Stenroos, Acta Soc. Fauna et Flora Fennica, vol. 17, No.

1, 1898, p. 132, pi. 1, fig. 29.- — Voigt, Forschungsber. Biol. Stat. Plon,

Earring (& Myers — Rotifer Fauna of Wisconsin — 11. 477

vol. 11, 1904, p. 44, pi. 3, fig. 21; Siisswasserfauna Deutschlands, pt. 14,

1912, p. 103, fig. 192. — De Beauchamp, Arch. Zool. Exper.,- ser. 4, vol. 6,

1907, p. 8, fig. 4. — Von Hofsten, Arkiv ZooL, Stockholm, vol. 6, No. 1,

1909, p. 16, fig. 1. — Mola, Ann. Biol. Lac., vol. 6, 1913, p. 244, —

Jakubski, Eozpr. Wiad. Muz. Dzieduszyckich, vol. 1, No. 3-4, 1915, p. 18.

— Weber and Montet, Cat. Invert. Suisse, pt. 11, 1918, p. 117.

Bistemma laeve Eichwald, Bull. Soc. Imp. Nat. Moscou, vol. 20, 1847, pt.

2, p. 343, pi. 9, fig. 4.

Furcularia tuhiformis King, Journ. Quekett Micr. Club, ser. 2, vol. 5, 1893,

p. 139, pi. 8, figs. 1-5.

Furcularia triliamata Stenroos, Acta Soc. Fauna et Flora Fennica, vol. 17,

No. 1, 1898, p. 133, pi. 2, fig. 14.

Notops falcipes Linder, Kev. Suisse Zool., vol. 12, 1904, p. 238, pi. 4, fig. 6.

Cephalodella forficula Barring, Bull. 81 U. S. Nat. Mus., 1913, p. 25.

The body is spindle-shaped, elongate and fairly slender. The

head is relatively large and slightly oblique anteriorly. The neck

is marked by a shallow constriction. The anterior half of the ab¬

domen is very nearly cylindric, the posterior slightly tapering ; the

integument is very flexible and the characteristic, discontinuous

lorica seems to be totally lacking. The foot is ill-defined, but a

slight basal constriction is usually present; the tail is small, but

fairly prominent, and belongs to the abdomen rather than to the

foot. The toes are short, stout, recurved and acutely pointed ; the

bases are somewhat swollen and near mid-length there is on the

dorsal side a prominent, toothlike spine, preceded by a transverse

row of small, acute spicules, from two to four in number. The

length of the toes is about one fifth of the total length. The foot

glands are very long and slightly clubshaped.

The corona is slightly oblique and convex without projecting lips.

The mastax is relatively large and of the normal type ; the inner

edges of the rami are serrate at the apex; the fulcrum is very

slightly expanded at the posterior end. The manubria are some¬

what unusual on account of the presence of a well developed, oval

basal plate ; the posterior ends are very slightly enlarged, but not

crutched. The gastric glands are small. There is a very faint con¬

striction between the stomach and intestine.

The ganglion is elongate and saccate; no retrocerebral organ is

present. The eyespot is frontal and of somewhat unusual struc¬

ture; its anterior half is transparent and the posterior pigment

mass is divided into two segments by a median, clear space.

Total length 160-1 65/x; toes 30-35ja; trophi 45/x.

478 Wisconsin Academy of Sciences y Arts, and Letters.

Cephalodella forficula is common in weedy ponds everywhere.

The propriety of including this species in the genus Cephalodella,

which is virtually equivalent to Gosse’s Diaschiza, as emended by

Dixon-Nuttall and Freeman, will probably not be questioned by

anybody, as the presence or absence of a lorica is now generally

conceded to be of small importance.

CEPHALODELLA PANAEISTA Myers, new species.

Plate XXXI, figures 5-7.

The body is very large, elongate and slender. The head is fairly

large and oblique anteriorly. The neck is marked by a slight con¬

striction. The abdomen increases gradually and very slightly in

width for about three fourths of its length ; the dorsal edge curves

downwards posteriorly to the base of the foot. The integument is

very flexible and the plates indistinct; the lateral clefts are very

obscure, but apparently fairly wide and parallel-sided. The foot

is short, stout and conical; the very small tail is near mid-length.

The toes are very long, stout and recurved, tapering gradually to

acute points ; their length is a little less than one third of the total

length. On the dorsal edge there is occasionally a single, toothlike

spine ; its distance from the base of the toe is about one third of the

length. The foot glands are extremely long and slightly club-

shaped.

The corona is strongly oblique, convex and without projecting

lips.

The mastax is large and the trophi robust. On the inner ventral

edges of the rami there are near the apex two denticulate, comblike

lamellae, the left one much larger than the right. The fulcrum

is long and straight, slightly expanded posteriorly; the manubria

are short, recurved posteriorly, but not crutched, and have a large

basal plate. The unci have the typical single tooth. The gastric

glands are small.

The ganglion is elongate and pyriform ; no retrocerebral organ is

present. The eyespot is frontal and the anterior part of the

capsule is without pigment, simulating a ‘‘lens’^

Total length 360-375/>t; toes 102-105/x; trophi 65/x..

Cephalodella panarista is rare ; we have found a few specimens at

Four Mile Run, near Washington, District of Columbia, and in

ponds at Tuckerton, Ocean County, New Jersej^ as well as in col-

Harring (& Myers — Rotifer Fauna of Wisconsin — II. 479

lections made by Dr. Birge at the Fish Hatchery, Bass Island,

Lake Erie, during the Great Lakes Biological Investigations in

1901.

CEPHALODBLLA AURICULATA (Muller) .

Plate XXVIII, figure 6.

Vorticella auriculata Muller, Verm. Terr. Fluv., vol. 1, pt. 1, 1773, p. 111.

Vorticella lacinulata Muller, Anim. Infus. 1786, p. 292, pi. 42, figs. 1-5=

Vorticella auriculata renamed.

Ecclissa lacinulata Schrank, Fauna Boica, vol. 3, pt. 2, 1803, p. 107.

Ecclissa hermanni Schrank, Fauna Boica, vol. 3, pt. 2, 1803, p. 109.

Furcularia lacinulata Lamarck, Hist. Nat. Anim. sans Vert., vol. 2, 1816,

p. 38.

Furcularia lohata Bory de St. Vincent, Enc. Meth., Zooph. (pt. 2), 1827,

p. 4:25=Vorticella lacinulata renamed.

Notommata lacinulata Ehrenberg, Abh. Akad. Wiss. Berlin, 1830, p. 46;

(for 1831), 1832, p. 134; Infusionsthierehen, 1838, p. 428, pi. 48, fig. 1+,

pi. 51, fig. 4. — Leydig, Zeitschr. Wiss. Zool., vol. 6, 1854, p. 38. — Gosse,

Phil. Trans. Royal Soc. London, vol. 146, 1856, p. 432, pi. 17, figs. 32-34.

— Bartsch, Jahresh. Ver. Naturk. Wiirrtemberg, vol. 26, 1870, p. 338. —

Eckstein, Zeitschr. Wiss. Zool., vol. 39, 1883, p. 364, pi. 24, fig. 22. —

Plate, Jenaische Zeitschr. Naturw., vol. 19, 1885, p. 23, pi. 1, fig. 6. —

Hudson and Gosse, Rotifera, 1886, vol. 2, p. 26, pi. 17, fig. 9. —

WiERZEJSKi, Rozpr. Akad. Umiej., Wydz. Mat.-Przyr., Krakow, ser. 2,

vol. 6, 1893, p. 228.

i EeTcinia calopodaria Morren, Ann. Sci. Nat., vol. 21, 1830, p. 139, pi. 3,

fig. 4; Bijdr. Natuurk. Wetensch., vol. 5, 1830, p. 225, fig.

P DeTcinia minutula Morren, Ann. Sci. Nat., vol. 21, 1830, p. 144, pi. 3, fig.

5; Bijdr. Natuurk. Wetensch., vol. 5, 1830, p. 230, fig.

? EeTcinia compta Morren, Ann. Sci. Nat., vol. 21, 1830, p. 146, pi. 3, fig. 7;

Bijdr. Natuurk. Wetensch., vol. 5, 1830, p. 231, fig.

Proales gibha Gosse, in Hudson and Gosse, Rotifera, 1886, vol. 2, p. 37,

pi. 18, fig. 8. — Skorikov, Trav. Soc. Nat. Kharkow, vol. 30, 1896, p. 290.

Notommata ovulum Gosse, Journ. Royal Micr. Soc., 1887, p. 2, pi. 1, fig. 3;

Hudson and Gosse, Rotifera, Suppl., 1889, p. 20, pi. 31, fig. 7.

Plagiognatha lacinulata Tessin, Arch. Naturg. Mecklenburg, vol. 43, 1890,

p. 149, pi. 1, fig. 9.

Notommata cuneata Thorpe, Journ. Royal Micr. Soc., 1891, p. 305, pi. 7,

fig. 5.

? Notostemma maTcrocephala Bergendal, Acta Univ. Lundensis, vol. 28,

1892, sect. 2, No. 4, pi. 69, pis. 2, 3, fig. 19.

? Notostemma bicarinata Bergendal, Acta Univ. Lundensis, vol. 28, 1892,

sect. 2, No. 4, p. 70, pi. 2, fig. 18.

Diaschisa lacinulata Levander, Acat. Soc. Fauna et Flora Fennica, vol. 12,

No. 3, 1895, p. 43. — -Weber, Rev. Suisse Zool., vol. 5, 1898, p. 545, pi. 18,

figs. 5-7. — Stenroos, Acta Soc. Fauna et Flora Fennica, vol. 17, No. 1,

1898, p. 156. — Dixon-Nuttall and Freeman, Journ. Royal Micr. Soc.,

1903, p. 11, pi. 2, fig. 6. — Voigt, Forschungsber. Biol. Stat. Plon, vol. 11,

480 Wisconsin Academy of Sciences, Arts, and Letters.

1904, p. 64. — Voronkov, Trudy Hidrobiol. Stants. Glubokom Oz., vol. 2,

1907, p. 106. — EunnstrOm, Zool. Anz., vol. 34, 1909, p. 270. — Lie-Pet-

TERSEN, Bergens Mus. Aarbog (for 1909), 1910, No. 15, p. 58. —

Sachse, Siisswasserfauna Deutsclilands, pt. 14, 1912, p. 119, figs. 229,

230. — Mola, Ann. Biol. Lac., vol. 6, 1913, p. 256. — Montet, Kev. Suisse

Zool., vol. 23, 1915, p. 330. — Jakubski, Eozpr. Wiad. Muz. Dzieduszy-

ckich, vol. 1, No. 3-4, 1915, p. 21. — Weber and Montet, Cat. Invert.

Suisse, pt. 11, 1918, p. 138.

Flagiognatcha lacinulata Skorikov, Trav. Soc. Nat. Kharkow, vol. 30,

1896, p. 292, pi. 7, fig. 9.

^ BiascMza taurocephalus Hilgendorf, Trans. New Zealand Inst., vol.

31, 1899, p. 123, pi. 10, fig. 9.

Diaschiza auriculata Barring, Bull. 81 U. S. Nat. Mus., 1913, p. 33;

Proc. U. S. Nat. Mus., vol. 47, 1913, p. 528.

The body is short, stout and somewhat prismatic; its greatest

depth is just behind the neck. The head is very large, oblique an¬

teriorly and slightly deflexed. The neck is well marked. The ab¬

domen is nearly parallel-sided, slightly narrower at the posterior

end. The lorica is fairly rigid and the plates distinct ; the lateral

clefts are narrow anteriorly and widen slightly and gradually to¬

wards the posterior end. The foot is short, stout and broadly

conical; the fairly large tail is somewhat beyond mid-length. The

toes are short, moderately stout and decurved, tapering gradually

to acute points; their length is about one fifth of the total length.

The foot glands are fairly large and pyriform.

The corona is oblique and strongly convex with prominent, beak¬

like lips.

The mastax is very large and of the normal type ; the fulcrum is

long and expanded posteriorly; the manubria very slender and

strongly recurved. The gastric glands are small and usually tinted

brownish-red in mature animals.

The ganglion is very large and saccate ; the eyespot is at the

posterior end. The retrocerebral organ is absent.

Total length 120-130/x ; toes 22-25/i,.

Cephalodella auriculata is common everywhere in weedy ponds.

In spite of its small size it is readily recognized by the peculiar,

jerky manner of swimming.

This species was first described by Muller under the name of

V orticella auriculata ; in his Animal cula Infusoria the specific name

was changed to lacinulata. seemingly under the impression that it

was fashioned somewhat like the flowers of the “BluebeH”, hollow

Earring & Myers — Rotifer Fauna of Wisconsin — 11. 481

and the margin cut into lappets or ‘ ‘ lacinulae ^ \ As auriculata is

the oldest name, it will have to take the place of lacinulata; both

are obviously misnomers.

CEPHALODELLA EXIGUA (Gosse).

Plate XXVIII, figure 2.

DiascMsa exigua Gosse, in Hudson and Gosse, Eotifera, 1886, vol. 2,

p. 78, pi. 22, fig. 15. — Dixon-Nuttall and Freeman, Journ. Eoyal

Micr. Soc., 1903, p. 133, pi. 3, fig. 10.— Von Hofsten, Arkiv Zool.,

Stockholm, vol. 6, No. 1, 1909, p. 52. — Sachse, Siisswasserfauna Deutseh-

lands, pt. 14, 1912, p. 121, figs. 233-234.— -Montet, Eev. Suisse Zool., vol.

23, 1915, p. 330. — Jakubski, Eozpr, Wiad. Muz. Dzieduszyckieh, vol. 1,

No. 3-4, 1915, p. 20. — ^Weber and Montet, Cat. Invert. Suisse, pt. 11,

1918, p. 140.

The body is rather short, stout and oblique anteriorly. The head

is very large and deflexed. The neck is well marked. The abdomen

is laterally compressed at the posterior end ; the dorsal and ventral

edges are almost parallel, the dorsal slightly arched; the lorica is

flexible, but the plates are fairly distinct ; the lateral clefts are wide

and parallel-sided anteriorly, the posterior ends slightly flaring.

The foot is very small, almost tubular, and gives the abdomen the

appearance of being squarely truncate posteriorly ; the tail is very

small and not far from the end of the foot. The toes are short, slen¬

der, slightly decurved and taper gradually to acute points; their

length is a little more than one fifth of the total length. The foot

glands are small and pyriform.

The corona is strongly oblique and distinctly convex without

projecting lips.

The mastax is large and of the normal type ; the fulcrum is very

slightly expanded at the posterior end; the manubria are very

slender and strongly recurved posteriorly, but not crutched. The

gastric glands are moderately large.

The ganglion is very elongate and saccate ; the retrocerebral organ

is absent. The large eyespot is at the posterior end of the ganglion.

Total length 90-95/x; toes 20-22/x,

C ephalodella exigua is present in weedy ponds everywhere, usu¬

ally in large numbers.

482 Wisconsin Academy of Sciences, Arts, and Letters,

CEPHALODELLA HOODII (Qosse).

Plate XXVIII, jSgure 1.

? Plagiognatlia lacinulata Dujardin, Hist. Nat. Zooph., Inf., 1841, pi. 18,

fig. 6; not Forticella lacinulata Muller.

Diaschi^a lioodii Gosse, in Hudson and Gosse, Eotifera, 1886, vol. 2, p. 79,

pi. 22, fig. 15. — Dixon-Nuttall and Preeman, Journ. Eoyal Mier. Soc.,

1903, p. 129, pi. 2, fig. 5. — Sachse, Siisswasserfauna Deutschlands, pt. 14,

1912, p. 120, figs. 231-232. — Jakubski, Eozpr. Wiad. Muz. Dzieduszyckicli,

vol. 1, No. 3-4, 1915, p. 21. — Weber and Montet, Cat. Invert. Suisse, pt.

11, 1918, p. 139.

DiaschiBa rJiamphigera Gosse, Journ. Eoyal Micr. Soc., 1887, p. 6, pi. 2,

fig. 20.— Hudson and Gosse, Eotifera, Suppl., 1889, p. 38, pi. 31, fig. 32.

Plagiognatlia gracilis Tessin, Arch. Naturg. Mecklenburg, vol. 43, 1890, p.

150, pi. 1, fig. 10.

Diaschiza valga Bilpinger, Jahresh. Ver. Naturk. Wurrtemberg, vol. 50,

1894, p. 53. — Weber, Eev. Suisse Zook, vol. 5, 1898, p. 549, pi. 20, figs.

26-28. — Lie-Pettersen, Bergens Mus. Aarbog (for 1909), 1910, No. 15,

p. 58. — Mola, Ann. Biol. Lac., vol. 6, 1913, p. 256.

The body is fairly slender and somewhat gibbous dorsally. The

head is fairly large, oblique anteriorly and slightly deflexed. The

neck is well marked. The abdomen is unusually elongate and the

dorsal line arched and rounded posteriorly; its greatest depth is

at mid-length; the lorica is firm and the plates well marked. The

lateral clefts are narrow and parallel-sided in the anterior half of

their length and widen gradually towards the posterior end. The

foot is rather small and conical; the tail is fairly prominent and

near mid-length. The toes are rather short, fairly stout and de-

curved, tapering gradually to acute points; their length is a little

less than one fourth of the total length. The foot glands are rather

small and pyriform.

The corona is moderately oblique and strongly convex with

prominent, beaklike lips.

The mastax is fairly large and of the normal type ; the fulcrum is

very slightly expanded posteriorly, the manubria short, very slen¬

der, rodlike and slightly recurved at the ends, but' not crutched.

The gastric glands are small and ovate.

The ganglion is very large and saccate ; the eyespot is fairly large

and at the posterior end of the ganglion. The retrocerebral organ

is absent.

Total length 140~145/x ; toes 32-35/x.

Cephalodella hoodii is not common; we have collected it at Madi¬

son, Wisconsin, in Fairmount Park, Philadelphia, Pennsylvania,

and near Los Angeles, California.

Earring & Myers — Botifer Fauna of Wisconsin — II. 483

CEPHALODELLA PLICATA Myers, new species.

Plate XXVIII, figures, 3-4.

The body is rather short and strongly gibbous dorsally. The

head is fairly large and somewhat deflexed. The neck is well

marked. The abdomen is strongly arched dorsally, with its greatest

depth near mid-length; the lorica is very firm and the plates well

marked; posteriorly it projects over the foot nearly to the base of

the toes. The dorsal and lateral clefts are abnormally deep and

their edges project as very distinct ridges, as shown in the optical

section of the body in figure 4; the dorsal cleft is straight-sided

and acute-angled, the lateral clefts rounded at the bottom. Sym¬

biotic zoochlorellae are invariably present and arranged in a

fairly regular and quite constant pattern. The foot is rather small ;

the minute tail is near the posterior end. The toes are rather short,

fairly stout, tapering, and slightly deeurved ; their length is about

one fourth of the total length. The foot glands are rather small and

pyriform.

The corona is moderately oblique and convex without projecting

lips.

The mastax is fairly large and of the normal type ; the fulcrum

is very slightly expanded at the posterior end; the manubria are

short, slender and without terminal crutch. No gastric glands

are present.

The ganglion is moderately elongate and saccate; no retrocere-

bral organ is present. The eyespot is at the posterior end of the

ganglion.

Total length 105-110/a ; toes 24-27/a.

Cephalodella plicata is not rare in soft, acid water ponds among

Fontinalis and submerged sphagnum ; we have collected it in Loon

Lake, Vilas County, and Starvation Lake, Oneida County, Wis¬

consin, and also at Bargaintown, near Atlantic City, New Jersey.

It has a superficial resemblance to C. hoodii and C. ventripes, but

the projecting beak is absent; the exceptionally deep dorsal and

lateral cleft's and the symbiotic zoochlorellae are sufficient to dis¬

tinguish it from these species.

484 Wisconsin Academy of Sciences, Arts, and Letters.

CEPHALODELLA VENTRIPES (Dixon-Nuttall) .

Plate XXVIII, figure 5,

Diaschiza ventripes Dixon-Nuttall, Journ. Quekett Mier. Club, ser. 2, vol.

8, 1901, p. 25, pi. 2, figs. 1-3. — ^Dixon-Nuttall and Freeman, Journ.

Eoyal Micr. Soc., 1903, p. 14, pi. 2, fig. 7. — Voronkov, Trudy Hidrobiol.

Stants. Glubokom Oz., vol. 2, 1907, p. 105.

The body is moderately stout and gibbous dorsally. The head

is large, strongly oblique anteriorly and slightly deflexed. The neck

is well marked. The abdomen is arched dorsally and the ventral

line straight; the lorica is very firm and the plates distinct, over¬

hanging the foot. The lateral clefts are rather narrow anteriorly

and increase gradually in width towards the posterior end. The

foot is small and broadly conical; the tail is prominent and near

the end of the foot. The toes are rather short, stout and decurved,

tapering gradually to acute points ; their length is a little less than

one fifth of the entire length. The foot glands are very small and

pyriform.

The corona is strongly oblique and convex with prominent, beak¬

like lips.

The mastax is large and of the normal type; the fulcrum is

slightly expanded posteriorly, the manubria are short, very slender

and the ends strongly recurved, but not crutched. The gastric

glands are fairly large and elongate ovate.

The ganglion is large and saccate ; the large, lens-shaped eyespot

is at the posterior end. No retrocerebral organ is present.

Total length 135-140ju ; toes 25-28jtx.

Cephalodella ventripes is widely distributed in weedy ponds, but

is not usually found in large numbers. It is readily distinguished

by the overhanging lorica and its large size.

CEPHALODELLA PHYSALIS Myers, new species.

Plate XXIX, figures 3-5.

The body is very short, stout and gibbous dorsally. The head

is very large and obliquely truncate anteriorly ; its length is. less

than the dorso-ventral depth. The neck is well marked. The ab¬

domen is but little longer than wide, somewhat prismatic and

deepest near mid-length. The lorica is quite firm and the lateral

cleft is wide. The foot is very short and stout; the tail is small

and knoblike and near the posterior end of the foot. The toes

Harring & Myers — Rotifer Fauna of Wisconsin — II. 485

are blade-shaped, decurved and acutely pointed; the dorsal edge

is evenly curved and the ventral edge has a blunt cusp about one

third of its length from the base; the length of the toes is one

fourth of the total length.

The corona is strongly oblique and the lips project as a small

beak.

The mastax is large and the trophi of normal type ; the fulcrum

is slightly expanded posteriorly and the manubria rodlike, not

crutched.

The ganglion is very large and saccate; the eyespot is at the

posterior end. There is no retrocerebral organ.

Total length 150-160ju, ; toes 35-40/i ; trophi 45ja.

Cephalodella physalis is not rare among sphagnum and sub¬

merged plants in soft-water ponds around Atlantic City, New

Jersey. It resembles C. Upara, but the body is not as stout; the

much longer toes and the presence of an eyespot are sufficient to

separate it from this species.

CEPHALODELLA STEIGOSA Myers, new species.

Plate XXIX, figure 7.

The body is moderately elongate, nearly eylindric and slightly

gibbous dorsally. The head is relatively long and strongly oblique

anteriorly. The neck is marked by a shallow constriction. The

abdomen increases very slightly and gradually in width for about

three fourths of its length ; from this point the dorsal edge curves

downward to the base of the foot. The integument is very flexible

and the plates ill-defined; the lateral clefts are somewhat obscure,

apparently narrow and parallel-sided throughout their length.

The foot is short and broadly conical ; the tail is a small, rounded

boss near mid-length. The toes are very long, slender, slightly de¬

curved and taper gradually to extremely acute, needle-like points ;

their length is about one fourth of the total length. The foot

glands are rather small and pyriform.

The corona is strongly oblique, decidedly convex and without

projecting lips.

The mastax is fairly large and of the normal type ; the fulcrum

is slightly expanded posteriorly, the manubria very slender and

not crutched. The gastric glands are small and rounded.

486 Wisconsin Academy of Sciences , Arts, and Letters.

The ganglion is elongate and saccate; no retrocerebral organ

is present. The eyespot is at the extreme posterior end of the

ganglion.

Total length 125/*; toes 33/>t.

Cephalodella strigosa is not common ; we have found it in weedy

ponds and among submerged sphagnum in Vilas County, Wiscon¬

sin, and around Atlantic City, New Jersey. There is a slide of

this species in the U. S. National Museum, mounted by C. F.

Rousselet and according to the label collected in Epping Forest;

it is erroneously determined as Diaschiza derhyi Dixon-Nuttall

and Freeman. This is probably the nearest relative of C. strigosa,

but differs in having recurved toes.

CEPHALODELLA TANTILLA Myers, new species.

Plate XXX, figure 2.

The body is moderately elongate, laterally compressed and

strongly gibbous dorsally. The head is large and slightly deflexed.

The neck is not very strongly marked. The abdomen increases

gradually in width for two thirds of its length and is somewhat

abruptly rounded posteriorly; the lorica is firm and the plates

well marked. The lateral clefts are rather narrow anteriorly and

increase gradually in width towards the posterior ends, which are

somewhat flaring. The foot is short and broad ; the fairly promi¬

nent tail is a little beyond mid-length. The toes are wide apart

at the base, long, slender and recurved, tapering gradually from

the base to acute points ; their length is about one third of the total

length. The foot glands are small and pyriform.

The corona is oblique and strongly convex without projecting

lips.

The mastax is relatively large and of the normal type; the

fulcrum is very stout and slightly expanded at the extreme end,

the manubria very slender, rodshaped and recurved, but not

crutched. The gastric glands are rather small.

The ganglion is long and saccate ; no retrocerebral organ is pres¬

ent. The eyespot is at the posterior end of the ganglion.

Total length 115~120/x; toes 38-40/a.

Cephalodella tantilla is common in weedy ponds with soft,

acid water; we have collected it in Vilas and Oneida Counties,

Wisconsin, around Atlantic City, New Jersey, and in Polk County,

Earring & Myers — Rotifer Fauna of Wisconsin — 11, 487

Florida. It bears a striking resemblance to C. gibha, from which

it differs in the much smaller size, cervical eyespot, acutely pointed

toes and the form of the trophi.

CEPHALODELLA COMPEESSA Myers, new species.

Plate XXX, figure 1.

The body is moderately elongate, slightly gibbous dorsally and

strongly compressed laterally. The head is rather small and dis¬

tinctly deflexed. The neck is well marked. The abdomen increases

gradually in width towards the posterior end. The lorica is fairly

rigid and the plates distinct; the lateral clefts are narrow an¬

teriorly and increase gradually in width towards the posterior end.

The foot is broad at the base and fairly stout. The toes are long,

slender and cylindric for about two thirds of their length; the

posterior third tapers gradually to very acute points; the dorsal

edge is straight throughout and the ventral bends upwards to meet

it. The length of the toes is about one third of the total length.

The corona is oblique and distinctly convex without projecting

lips.

The mastax is large and of the normal type ; the fulcrum is long,

stout and slightly expanded at the posterior end; the manubria

are very slender, rodlike and strongly recurved posteriorly. The

gastric glands are small and irregularly ovate.

The ganglion is long and saccate ; the eyespot is at the posterior

end, well towards the ventral side. The retrocerebral organ is

absent.

Total length 140-145/>t; toes 45-48/a.

Cephalodella compressa is common in sphagnum and among sub¬

merged plants in soft, acid water ponds and bogs in Vilas and

Oneida Counties, Wisconsin, and around Atlantic City, New Jer¬

sey. It is readily recognized by the strongly compressed body.

CEPHALODELLA DORSEYI Myers, new species.

Plate XXX, figure 7.

The body is relatively short, stout and slightly gibbous dorsally.

The head is large, slightly deflexed and strongly oblique ante¬

riorly. The neck is well marked. The abdomen increases grad¬

ually in width for about two thirds of its length and is gently

rounded posteriorly ; the lorica is firm and the plates distinct ; the

488 Wisconsin Academy of Sciences, Arts, and Letters.

lateral clefts are fairly wide at the ends and slightly narrower at

mid-length. The foot is short and stout; the tail is fairly promi¬

nent and near the posterior end. The toes are very long and

wide apart at the base, fairly stout and tapering for about

one fifth of their length; from this point they are very nearly

cylindric and end in acute, conical tips, the dorsal edge straight

and the ventral bending abruptly upwards to meet it. The length

of the toes is about one third of the total length. The foot glands

are small and pyriform.

The corona is strongly oblique and distinctly convex without

projecting lips.

The mastax is large and of the normal type; the fulcrum is

moderately long and slightly expanded at the posterior end; the

manubria are slender, rodshaped, slightly clubbed and decurved

posteriorly, but not crutched. The gastric glands are small and

ovate.

The ganglion is long and saccate; the eyespot is very lafge and

saucer-shaped and at the posterior end of the ganglion. The retro-

cerebral organ is absent.

Total length 145-150/x; toes 50-54/a.

Cephalodella dorseyi is not very common. We have collected

it among submerged sphagnum and other plants in soft, acid water

ponds around Atlantic City, New Jersey. It is readily recog¬

nized by the long, very slender toes with their peculiar, clawlike

points.

CEPHALODELLA PITJLCA Myers, new species.

Plate XXX, figure 3.

The body is elongate, slender, slightly curved and laterally com¬

pressed. The head is relatively small and short and distinctly de-

flexed. The neck is not strongly marked. The abdomen is nearly

parallel-sided, very slightly gibbous dorsally and concave ven-

trally ; the lorica is firm and the plates well marked. The lateral

clefts are narrow anteriorly and increase slightly in width to¬

wards the posterior end. The foot is fairly long and stout; the

small tail is near the posterior end. The toes are long, slender

and slightly recurved, moderately stout at the base and taper grad¬

ually to extremely slender, acute points; their length is a little

less than one third of the total length. The foot glands are rather

small and pyriform.

Harring & Myers — Rotifer Fauna of Wisconsin — II. 489

The corona is oblique and strongly convex without projecting

lips.

The mastax is small and of the normal type; the fulcrum is

very slightly expanded posteriorly and the manubria distinctly

crutched. The gastric glands are small and somewhat triangular.

The ganglion is long and saccate; the small eyespot is at the

posterior end, on the dorsal side. No retrocerebral sac is present.

Total length 130-135/x ; toes 38-40)U.

Cephalodella hiulca is not rare in soft, acid water ponds among

Fontinalis, Riccia and sphagnum ; we have collected it at Bargain-

town and Oceanville, New Jersey. It resembles superficially

0. strepta, but is much larger and without the head sheath of this

species; an additional distinction is the presence of an eyespot in

C. hiulca.

CEPHALODELLA ELEGANS Myers, new species.

Plate XXXI, figure 8.

The body is elongate, very slender, somewhat curved and slightly

compressed dorso-ventrally. The head is relatively large and very

slightly defiexed. The neck is not very strongly marked. The

abdomen is slightly curved dorsally and nearly straight ventrally;

the lorica is firm and the plates distinct. The lateral clefts are

narrow anteriorly and increase gradually and considerably in

width towards the posterior end. The foot is small and terminates

in a short, cylindric section; the tail is very minute and only a

short distance beyond the lorica. The toes are very long and

slightly decurved, fairly stout at the base and tapering rapidly

to slender, nearly cylindric, acutely pointed rods; their length is

about one third of the total length. The foot glands are rather

small and pyriform.

The corona is slightly oblique and moderately convex without

projecting lips.

The mastax is small and of the normal type; the fulcrum is

slightly expanded at the posterior end and the manubria are dis¬

tinctly recurved and increase somewhat in width towards the ends.

The gastric glands are small and oval.

The ganglion is long and pyriform; the eyespot is fairly large

and at the posterior end of the ganglion; no retrocerebral organ

is present.

Total length 135-140/x; toes 48-50/x.

490 Wisconsin Academy of Sciences, Arts, and Letters.

Cephalodella elegans is not common ; we have collected it in small

numbers among sphagnum and other submerged plants in a swamp

near Oceanville, New Jersey. It is readily distinguished by the

very slender, depressed body and the long toes.

CEPHALODELLA GALBINA Myers, new species.

Plate XXXI, figure 1.

The body is very short, stout and gibbous dorsally. The head

is relatively short, very broad and slightly deflexed. The neck is

not strongly marked. The abdomen is short and very broad, with

its greatest width near mid-length. The lorica is very flexible,

but the plates are well marked; the lateral clefts are narrow an¬

teriorly and increase gradually in width towards the posterior end.

The foot is short and stout ; the tail is fairly prominent and some¬

what beyond mid-length. The toes are very long, slender, slightly

decurved, rather stout at the base and taper gradually to very

acute points; their length is about two fifths of the total length.

The foot glands are small and pyriform.

The corona is slightly oblique and strongly convex without pro¬

jecting lips.

The mastax is huge and of the normal type; the fulcrum is

nearly half the length of the entire body and the posterior end

very slightly expanded, the manubria short, slender, slightly re¬

curved and clubbed posteriorly, but not crutched. The gastric

glands are very small and ovate.

The ganglion is very long and saccate; no retrocerebral organ

is present. The eyespot is at the posterior end of the ganglion.

Total length 100-110/x ; toes 38-42/x.

Cephalodella galhina is rather rare ; we have collected it only

among submerged sphagnum in a soft, acid water pond at Gravelly

Kun, near Mays Landing, New Jersey. Its nearest relative is prob¬

ably C. dorseyi, from which it differs in the much shorter and

stouter body, huge mastax and long, decurved toes.

CEPHALODELLA BELONE Myers, new species.

Plate XXXI, figure 2.

The body is small and conical, tapering increasingly from the

head towards the foot. The head is very large, fully one third the

length of the entire body and somewhat wider than the abdomen.

Earring & Myers — Rotifer Fauna of Wisconsin — II. 491

The neck is well marked. The abdomen tapers rapidly and in¬

creasingly from the neck to the base of the foot; the lorica is

fairly rigid and the plates are well marked ; the lateral clefts are

very narrow and parallel-sided. The foot is moderately long, coni¬

cal and rather small at the base; the tail is small and a little

beyond mid-length. The toes are extremely long, straight and

slender, slightly enlarged at the base and tapering very gradually

to the minutely rounded tips; their length is about two fifths of

the entire length.

The corona is somewhat oblique and convex with prominent

beaklike lips.

The mastax is large and of the normal type; the fulcrum is

relatively stout and slightly expanded posteriorly; the manubria

slender, rodlike and slightly recurved, but not crutched. The

gastric glands are small.

The ganglion is long and pyriform; the eyespot is cervical, at

the posterior end of the ganglion ; no retrocerebral organ is present.

Total length 120-125|Lt; toes 45-50/a.

Cephalodella helone is not common; we have collected it among

Fontinalis in a decadent lake two miles east of Eagle River, Vilas

County, Wisconsin, and in a bog pool at Bargaintown, near At¬

lantic City, New Jersey. The appearance of the animal is at first

somewhat puzzling, whether alive or dead: when swimming the

toes are never separated and it looks very much like a diminutive

Trichocerca (=Battulus) ; when death occurs, the toes are thrown

out sidewise, thus resembling a small Monommata. It is probably

related to C. cuneata, but readily distinguished by the presence of

the eyespot, as well as by its peculiar behavior.

CEPHALODELLA NANA Myers, new species.

Plate XXIX, figure 1.

The body is relatively short and conical, tapering evenly and

gradually from the corona to the base of the toes ; the dorsal edge

is gently curved. The head is very large, nearly half the length of

the entire body and much wider than the abdomen. The neck is

well marked. The abdomen is very short, little more than one third

of the length of the body and tapers gradually towards the foot;

the lorica is moderately flexible and the plates distinct ; the lateral

clefts are narrowest near mid-length and wide at the posterior end.

The foot' is fairly large and conical ; the small tail is unusually far

492 Wisconsin Academy of Sciences, Arts, and Letters.

back on the foot. The toes are long, slender, wide apart at the base,

double-curved and taper gradually to acute, bristle-like points;

their length is about one third of the total length. The foot glands J

are small and pyriform. ^

The corona is strongly oblique and convex with prominent, beak- 1

like lips. I

The mastax is very large and of the normal type; the fulcrum is ^

slightly expanded at the posterior end; the manubria are slender, ;

slightly clubbed and recurved at the ends, but not crutched. The

gastric glands are small.

The ganglion is very long and somewhat pyriform; the eyespot

is at the posterior end. No retrocerebral organ is present.

Total length 105-110/a; toes 35-40/x.

Cephalodella nana has been collected only in sphagnum along the

marshy edges of Corduroy Creek, near Absecon, New Jersey. It

resembles C. cuneata, but the presence of an eyespot and the sig¬

moid curvature of the toes is sufficient to distinguish it.

CEPHALODELLA XENICA Myers, new species.

Plate XXIX, figure 6.

The body is moderately elongate and gibbous dorsally. The head

is large and oblique anteriorly. The neck is marked by a rather

shallow constriction. The abdomen is arched dorsally, its greatest

depth near mid-length ; the lorica is fairly rigid and the plates well

marked ; the lateral clefts are wide and parallel-sided. The foot is

large and robust ; the prominent tail is near the posterior end. The

toes are short, blade-shaped and very broad ; they increase slightly

in width for about half their length, then decrease rather suddenly

to very slender, conical tips, slightly blunted at the extreme ends;

their length is less than one fifth of the total length. The foot

glands are rather small and ovate.

The corona is oblique and strongly convex without projecting

lips.

The mastax is fairly large and of the normal type; the fulcrum

is very slightly expanded at the posterior end; the manubria are

long and have a very pronounced terminal crutch. The gastric

glands are small and oval.

The ganglion is elongate and pyriform; no retrocerebral organ

is present. The eyespot is at the posterior end of the ganglion.

Total length 128/1 ; toes 22/i.

Harring & Myers — Rotifer Fauna of Wisconsin — 11. 493

Cephalodella xenica was found in considerable numbers in ma¬

terial collected by Dr. H. S. Jennings in Huron River at Ann Ar¬

bor, Michigan, in 1901. Its nearest relative is C. eupoda, which

differs in its greater size and in the absence of the eyespot, as well

as the flexible integument.

CEPHALODELLA NELITIS Myers, new species.

Plate XXXII, figure 1.

The body is elongate, very slender and somewhat prismatic. The

head is fairly small and strongly oblique anteriorly. The neck is

not strongly marked. The abdomen increases slightly in width

for about two thirds of its length and from this point tapers slightly

to the base of the foot ; it is somewhat variable in width and speci¬

mens with parallel-sided body are occasionally met with. The lorica

is very thin and flexible, but the plates are fairly well marked ; the

lateral clefts are rather narrow and parallel-sided for nearly their

entire length ; the extreme posterior ends are slightly flaring. The

foot is short and conical, but broad at the base; the tail is very

small. The toes are short, strongly decurved and very slender;

from the slightly bulbous base they taper very gradually to acute

points ; their length is about one fifth of the total length. The foot

glands are small and pyriform.

The corona is oblique, strongly convex and without projecting

lips.

The mastax is rather small and of the normal type ; the fulcrum

is stout and slightly expanded posteriorly, the manubria rodlike,

very slender and not crutched. The bladder is large.

The ganglion is elongate and saccate; eyespot and retrocerebral

organ are absent.

Total length 135ju,; toes 22jii.

Cephalodella nelitis was collected in a pond with soft, acid water

among submerged sphagnum, at Gravelly Run, near Mays Landing,

New Jersey. Its nearest relative is C. melia; a comparison is given

under this species.

CEPHALODELLA MELIA Myers, new species.

Plate XXXII, figure 2.

The body is elongate, somewhat prismatic and gibbous dorsally.

The head is rather small, but relatively long in comparison with

494 Wisconsin Academy of Sciences^ Arts^ and Letters.

its width, and obliquely truncate anteriorly. The neck is well

marked. The abdomen increases gradually and considerably in

width for about three fourths of its length; the dorsal line curves

rapidly downward in the posterior fourth. The lorica is very thin

and flexible and the edges of the plates rather difficult to trace ; the

lateral clefts are very narrow anteriorly and increase gradually in

width towards the posterior end. The foot is long and conical and

the tail prominent. The toes are fairly long and slender, enlarged

at the base, parallel-sided for about four fifths of their length, de-

curved and end in small conical points, prolonged by a fairly long,

bristle-like nib, continuing the line of the dorsal edge ; their length

is about' one fourth of the total length. The foot glands are fairly

large and pyriform.

The corona is oblique, strongly convex and without projecting

lips.

The mastax is moderately large and of the normal type ; the ful¬

crum curves upwards at the posterior end; the manubria are not

crutched. Symbiotic zoochlorellae are present in abundance in the

walls oT the stomach. The bladder is very large.

The ganglion is long and saccate; eyespot and retrocerebral

organ are absent.

Total length 130-145ja ; toes 28-32/x.

Cephalodella melia was originally collected in soft, acid water

ponds and bogs in the neighborhood of Atlantic City, New Jersey;

it has not been seen for two years, but was until then fairly com¬

mon. Its nearest relative is probably C. neliiis, from which it is

readily distinguished by the form and length of the toes and the

shorter and strongly gibbous body, as well as by the presence of

zoochlorellae.

CEPHALODELLA MEGALOCEPHALA (Glasscott).

Plate XXXII, figures 5-7.

i Pleurotrocha leptura Ehrenberg, Abh. Akad. Wiss. Berlin (for 1831),

1832, p. 129, pi. 4, fig. 18; Infusionsthierchen, 1838, p. 419, pi. 48, fig. 2.

? Hudson and Gosse, Eotifera, 1886, vol. 2, p. 20, pi. 18, fig. 4. —

? Voigt, Siisswasserfauna Deutschlands, pt. 14, 1912, p. 86, fig. 149.

? Furcularia lactistes Gosse, Journ. Koyal Micr. Soc., 1887, p. 863, pi. 14,

fig. 5. — Hudson and Gosse, Eotifera, SuppL, 1889, p. 25, pi. 31, fig. 13.

Furcularia megalocephala Glasscott, Proc. Eoyal Dublin Soc., new ser., vol. 8,

1893, p. 56, pi. 4, fig. 3.

Harring & Myers- — Rotifer Fauna of Wisconsin — 11. 495

/ Diglena inflata Glasscott, Proe. Eoyal Dublin Soc., new ser., vol. 8, 1893,

p. 60, pi. 4, fig. 6.

Biaschiza megalocephala Eousselet, Journ. Quekett Micr. Club, ser. 2, vol. 6,

1895, p. 123, pi. 7, fig. 5. — Dixon-Nuttall and Freeman, Journ. Eoyal

Micr. Soc., 1903, p. 139, pi. 4, fig. 14. — Voigt, Forschungsber. Biol. Stat.

Plon, vol. 11, 1904, p. 65. — Sachse, Siisswasserfauna Deutschlands, pt. 14,

1912, p. 123, figs. 240-242. — Jakubski, Eozpr. Wiad. Muz. Dzieduszyckich,

vol. 1, No. 3-4, p. 21. — Hauer, Mitt. Bad. Landesver. Naturk., Freiburg i.

Br., new ser., vol. 1, 1921, p. 179.

The body is fairly stout and gibbous dorsally. The head is very

large and extremely oblique. The neck is not strongly marked.

The abdomen increases considerably in width for about two thirds

of its length and from this point the dorsal line curves rapidly to

the base of the foot. The lorica is very thin and flexible and the

edges of the plates ill-defined; the lateral clefts are narrow and

parallel-sided. The presence of a dorsal cleft is denied by Dixon-

Nuttall and Freeman; according to Hauer it is really present, but

the connecting portion of the integument is convex instead of

forming a deep groove, as in other species of this genus. The foot

is stout and conical, but its base is ill-defined; the knoblike tail is

near mid-length. The toes are short, stout and decurved, tapering

gradually to acute points; their length is about one sixth of the

total length. The foot glands are large and pyriform.

The corona is extremely oblique, very slightly convex and with¬

out projecting lips.

The mastax is large and differs very little from the typical form,

except in the feeble development of the mallei.

The ganglion is very elongate and saccate; eyespot and retro-

cerebral organ are absent.

Total length 195-210jr ; toes 34-38/x ; trophi 30jLt.

Cephalodella megalocephala is very widely distributed, if not cos¬

mopolitan; it is common in weedy ponds wherever we have col¬

lected. Dixon-Nuttall and Freeman included this species in the

genus Diaschiza under protest; this was not unreasonable under

their somewhat narrow and artificial definition of this genus. No

objection can be raised against its inclusion in Cephalodella^ if

the definition given above is accepted for this genus, basing it pri¬

marily on the form of the corona and the mastax, secondarily on

the form of the head, body and foot, and neglecting the degree of

development of the lorica, even its total absence, as well as the

496 Wisconsin Academy of Sciences , Arts, and Letters.

presence of the minute and all but invisible tuft of setae at the base

of the toes, which we consider trivial.

Ehrenberg’s figure and description of Pleurotrocha leptura prob¬

ably refer to this species; his figure of the mastax accompanying

the original description shows that it belongs to C ephalodella. As

his name has never been used by anybody else for the species under

consideration and the identification is not absolutely certain, its

resurrection at this time is probably unnecessary.

CEPHALODELLA PHELOMA Myers, new species.

Plate XXXII, figure 3.

The body is elongate, slender and nearly cylindric. The head is

unusually long and extremely oblique. The neck is slightly con¬

stricted dorso-ventrally and excessively compressed transversely,

its width being less than half the width of the head. The abdomen

is fusiform and increases slightly in width towards the base of the

foot; the integument is so flexible that it can hardly be called a

lorica, and the plates are very obscure ; the lateral clefts are narrow

anteriorly and increase gradually in width towards the posterior

end. The foot is large and robust; the tail is fairly prominent.

The toes are short, stout and decurved, with the ventral edges some¬

what undulate; the basal portion is nearly cylindric and the pos¬

terior tapers gradually to acute points; their length is about one

sixth of the total length. The foot glands are very long and tubu¬

lar.

The corona is excessively oblique and slightly convex without pro¬

jecting lips.

The mastax is large and of the normal type ; the fulcrum is broad

and very slightly expanded posteriorly, the manubria unusually

long, nearly straight, rodlike and stout. The bladder is very

large.

The ganglion is very long and pyriform ; eyespot and retrocere-

bral organ are absent.

Total length 200/x; toes 35/x.

C ephalodella pheloma is rare; we have found only a few speci¬

mens among Fontinalis and submerged sphagnum growing in soft,

acid water in a shallow pond at Estellville, near Atlantic City,

New Jersey. It appears to be closely related to C. megalocephala,

but is readily distinguished by the more slender body, long foot

Harring (& Myers — Rotifer Fauna of Wisconsin — 11. 497

glands, the form of the toes, as well as the remarkable compression

of the neck, so striking in a dorsal view of the animal; when the

species was first found, we assumed this to be caused by a patho¬

logical condition, but this was evidently an error, as it is always

present in perfectly healthy individuals.

CEPHALODELLA TENUIOR (Gosse).

Plate XXXIII, figure 3.

Diaschiza tenuior Gosse, in Hudson and Gosse, Rotifera, 1886, vol. 2, p. 81,

pi. 22, fig. 14. — Dixon-Nuttall and Freeman, Journ. Royal Micr. Soc.,

1903, p. 135, pi. 4, fig. 12. — Sachse, Siisswasserfauna Deutschlands, pt. 14,

1912, p. 123, fig. 239. — Jakubski, Rozpr. Wiad. Muz. Dzieduszyekich,

vol. 1, No. 3-4, 1915, p. 21. — Weber and Montet, Cat. Invert. Suisse, pt.

11, 1915, p. 141.

The body is elongate, slender and nearly cylindric. The head is

large, slightly deflexed and oblique anteriorly. The neck is not

strongly marked. The abdomen is very nearly parallel-sided,

slightly gibbous posteriorly; the lorica is very flexible and the

plates indistinct; the lateral clefts are narrow anteriorly and in¬

crease slightly and evenly in width for about two thirds of their

length; at this point the dorsal plates become rounded and the

cleft widens rapidly. The foot is relatively short and broadly coni¬

cal ; the tail is near the end of the foot and fairly prominent. The

toes are short, very nearly straight and parallel-sided for about

two thirds of their length; beginning here the ventral edge curves

upwards to meet the dorsal edge in an acute point; the length of

the toes is one flfth of the total length. The foot glands are very

small.

The corona is oblique, strongly convex and without projecting

lips.

The mastax is large and of the normal type; the fulcrum is

slightly expanded posteriorly and the manubria erutched. The

gastric glands are small and occasionally have a brownish tint.

The ganglion is very long and pyriform ; eyespot and retrocere-

bral organ are absent.

Total length, 120~125/>i; toes 22-24jLt.

Cephalodella tenuior is fairly common in weedy ponds every¬

where. It is closely related to C. forficata, from which it differs in

the form of the toes, the much smaller size, flexible lorica and the

form of the lateral clefts.

498 Wisconsin Academy of Sciences, Arts, and Letters.

CEPHALODELLA EETUSA Myers, new species.

Plate XXXIII, figure 6.

The body is rather short, fusiform and fairly stout. The head

is large and obliquely truncate anteriorly. The neck is not very

strongly marked. The abdomen is nearly parallel-sided and very

slightly gibbous dorsally ; the lorica is fairly rigid and the plates

well marked; the lateral clefts are narrow and parallel-sided for

about one half of their length, increasing slightly in width towards

the posterior end. The foot is moderately large and robust; the

tail is very small. The toes are short and stout; the ventral edge

is straight in its entire length, the dorsal for about three fourths

of its length, and from this point it curves rapidly towards the

ventral edge, meeting it in an acute point ; the length of the toes

is about one fourth of the entire length. The foot glands are small

and pyriform.

The corona is strongly oblique and convex without projecting

lips.

The mastax is fairly large and of the normal type ; the fulcrum

is expanded posteriorly and the manubria crutched.

The ganglion is very large and saccate ; eyespot and retrocerebral

organ are absent.

Total length 100/x ; toes 24ja.

C ephalodella retusa is not common; we have collected it among

sphagnum growing on the bottom in shallow water of Doughty’s

Mill Pond, about two miles west of Absecon, New Jersey. It is

related to C. forficata, but is much smaller and the form of the

toes is quite different.

CEPHALODELLA DIXON-NUTTALLI Myers, new species.^

Plate XXXIII, figures 4-5.

The body is elongate, fairly slender and laterally compressed.

The head is rather small, slightly deflexed, and obliquely truncate

anteriorly. The neck is not strongly marked. The abdomen in¬

creases gradually and very regularly in width from the neck to

the posterior end ; the dorsal and ventral edges are almost straight

lines. The lorica is very thin and flexible and the plates are diffi¬

cult to trace ; the lateral clefts are narrow anteriorly and the edges

diverge rapidly and regularly towards the posterior end, where

their distance apart is half the width of the abdomen. The foot

Harring & Myers — Rotifer Fauna of Wisconsin — II. 499

is nearly half as long as the abdomen and very stout ; the tail is

small and somewhat beyond mid-length. The toes are short and

taper very slightly for one half of their length, changing abruptly

to conical, acute points ; their length is a little less than one fifth of

the total length. The foot glands are very small and pyriform.

The corona is oblique, slightly convex and without projecting

lips.

The mastax is rather small and of the normal type ; the fulcrum

is very slightly expanded posteriorly, the manubria very slender

and not crutched. The gastric glands are very small.

The ganglion is moderately elongate and pyriform; eyespot and

rectrocerebral organ are absent.

Total length 160/x; toes 30/x.

Cephalodella dixon-nuttalli is known only from Lake Kathan,

Oneida County, Wisconsin; it was collected among submerged

sphagnum and other small aquatic plants. All the specimens seen

had the toes closely appressed, producing the appearance of a single

toe.

CEPHALODELLA FORFICATA (Ehrenberg) .

Plate XXXIII, figure 7.

Notommata forficata Ehrenberg, Abh. Akad. Wiss. Berlin (for 1831),

1832, p. 134.

Notommata forcipata Ehrenberg, Infusionsthierchen, 1838, p. 428, pi. 51,

fig. 5.

Furcularia caeca Gosse, Ann. Mag. Nat. Hist., ser. 2, vol. 8, 1851, p. 199.

— Hudson and Gosse, Eotifera, 1886, vol. 2, p. 42, pi. 20, fig. 4.

Furcularia ensifera Gosse, in Hudson and Gosse, Eotifera, 1886, vol. 2, p. 43,

pi. 20, fig. 3.

Fiaschiza paeta Gosse, in Hudson and Gosse, Eotifera, 1886, vol. 2, p. 79,

pi. 22, fig. 11. — Bilpinger, Jahresh. Ver. Naturk. Wiirttemberg, vol. 50,

1894, p. 53. — Levander, Acta Soc. Fauna et Flora Fennica, vol. 12, No. 3,

1895, p. 44. — Eunnstrom, Zool. Anz., vol. 34, 1909, p. 270. — Lie-

Pettersen, Bergens Mus. Aarbog (for 1909), 1910, j). 59.

Fiaschiza acronota Gosse, Journ. Eoyal Micr. Soe., 1887, p. 867, pi. 15,

fig. 15. — Hudson and Gosse, Eotifera, Suppl., 1889, p. 37, pi. 31, fig. 29.

Fiaschiza caeca Dixon-Nuttall and Freeman, Journ. Eoyal Mier. Soc.,

1903, p. 134, pi. 4, fig. 11. — Voigt, Forschungsber. Biol. Stat. Plon, vol. 11,

1904, p. 64. — Voronkov, Trudy Hidrobiol. Stants. Glubokom Oz., vol. 2,

1907, p. 105. — Von Hofsten, Arkiv Zool., Stockholm, vol. 6, No. 1, 1909,

p. 53. — Sachse, Siisswasserfauna Deutschlands, pt. 14, 1912, p. 121, figs.

235, 236. — Jakubskt, Eozpr. Wiad. Muz. Dzieduszyckich, vol. 1, No. 3-4,

1915, p. 20. — Weber and Montet, Cat. Invert. Suisse, pt. 11, 1918, p. 141.

FiascMza forficata Harring, Bull. 81 U. S. Nat. Mus., 1913, p. 34; Proc.

U. S. Nat. Mus., vol. 47, 1913, p. 528.

500 Wisconsin Academy of Sciences, Arts, and Letters.

The body is elongate, fairly slender, somewhat prismatic and

slightly compressed laterally. The head is large and obliquely

truncate anteriorly. The neck is well marked. The abdomen is

very nearly parallel-sided and slightly convex dorsally ; the lorica

is firm and the plates well marked ; the lateral clefts are narrow and

parallel-sided. The foot is conical and rather short; the tail is

small and rounded. The toes are fairly wide apart at the base,

moderately short, stout, very slightly recurved and taper gradu¬

ally to acute points ; their length is a little less than one fourth of

the total length. The foot glands are moderately large and pyri¬

form.

The corona is strongly oblique and convex without projecting

lips.

The mastax is large and of the normal type; the fulcrum is

slightly expanded at the posterior end and the manubria are

crutched. The gastric glands are large and pigmented red.

The ganglion is long and saccate ; at its posterior end is a small

retrocerebral sac with a distinct duct, bifurcate anteriorly, but

not reaching the surface of the corona. There is no eyespot.

Total length 175-184/x; toes 36-40jLi.

Cephalodella forficata is common everywhere in weedy ponds.

We have used Ehrenberg ’s name for this species, as it is undoubted¬

ly the same as Gosse’s Diaschiza caeca; the form of the toes, often

crossed, size 150/x, and the very large ' ‘ eye, ’ ’ that is : gastric glands,,

all agree with Gosse’s description.

(7. forficata, tenuior, collactea and intuta show considerable simi¬

larity; the form of the body is approximately the same, all have

pigmented gastric glands and crutched manubria. They are readily

distinguished by the form of the toes; C. tenuior, collactea and

intuta are also much smaller than normal specimens of C. forficata.

CEPHALODELLA INTUTA Myers, new species.

Plate XXXV, figures 2-5.

The body is moderately elongate and slender, laterally com¬

pressed and slightly gibbous posteriorly. The head is relatively

long on the dorsal side and short on the ventral side on account of

the strongly oblique corona. The neck is well marked. The abdo¬

men increases gradually in depth for about two thirds of its length

and from there tapers rapidly towards the short, conical foot ; the

Earring & Myers — Rotifer Fauna of Wisconsin — II. 501

tail is small and rounded. The lorica is fairly rigid and the plates

well marked. The lateral clefts are narrow anteriorly and increas

gradually in width towards the posterior end of the lorica. The

toes are faintly recurved, wide apart at the base, very long and

slender, tapering slightly to acute, clawlike points with transverse

basal septa ; their length is about one fourth of the total length.

The corona is convex and strongly oblique, without projecting

lips.

The mastax is large and of the normal type; the fulcrum is

broadly expanded at the posterior end and the manubria crutched.

The gastric glands are tinted red.

The ganglion is long and saccate ; at its posterior end is a small

sac with well marked duct, bifurcate at the anterior end, but not

reaching the surface of the corona. There is no eyespot.

Total length 115-125/x ; toes 30-35/x ; trophi 40/x.

Cephalodella intuta has been collected among submerged sphag¬

num and other aquatic vegetation in Loon Lake, about one mile

south of Eagle Eiver, Vilas County, Wisconsin, and in soft, acid

water ponds around Atlantic City, New Jersey. Its nearest rela¬

tive is C. forficata, from which it differs in the form of the body,

shorter foot and much longer and more slender toes, as well as in

its smaller size.

CEPHALODELLA COLLACTEA Myers, new species.

Plate XXXIV, figure 5.

The body is moderately elongate, nearly cylindrical and slightly

gibbous dorsally. The head is very short and strongly oblique

anteriorly. The neck is well marked. The abdomen increases

gradually in width for nearly three fourths of its length and from

this point tapers rapidly to the large, robust foot; the tail is a

small rounded boss. The lorica is fairly rigid and the plates well

marked ; the lateral clefts are narrow anteriorly and increase grad¬

ually in width towards the posterior end. The toes are long,

straight, slender and very slightly tapering; the dorsal edge is

straight throughout; the ventral edge is straight for about five

sixths of the length of the toe; at this point it bends abruptly at

an obtuse angle to meet the dorsal line and this tapering portion

of the toe ends in a bristle-like point. The length of the toes is

502 Wisconsin Academy of Sciences, Arts, and Letters.

one fourth of the total length. The foot glands are fairly large

and pyriform.

The corona is oblique and strongly convex without projecting

lips.

The mastax is small; the fulcrum is straight and slightly ex¬

panded posteriorly, the manubria slender and crutched. The

gastric glands are frequently tinted red.

The ganglion is moderately long and saccate ; eyespot and retro-

cerebral organ are absent.

Total length 130/x ; toes 32/x.

Cephalodella collactea has been found only in Loon Lake, Vilas

County, Wisconsin. Its nearest relative is C. intuta, from which it

is readily distinguished by the form of the toes.

CEPHALODEIjLA INQTTILINA Myers, new species.

Plate XXXVI, figure 1.

The body is moderately elongate, nearly parallel-sided and later¬

ally compressed. The head is short and broad, somewhat longer

dorsally than ventrally. The neck is well marked. The abdomen

is either parallel-sided or slightly wider posteriorly; the lorica is

very thin and flexible and the edges of the plates indistinct; the

lateral clefts are narrow anteriorly and increase gradually in width *

towards the posterior end. The foot is large and very broad at

the base ; the tail is very small. The toes are long, slender, nearly

straight, somewhat enlarged at the base and very slightly tapering,

suddenly reduced at the ends to minute, but very distinct claws;

their length is about one fourth of the total length. The foot glands

are moderately large and pyriform.

The corona is decidedly oblique and strongly convex without

projecting lips.

The mastax is relatively small ; the posterior end of the fulcrum

is broadly and abruptly expanded ; the manubria are crutched.

The ganglion is long and saccate; eyespot and retrocerebral

organ are absent.

Total length 250-270/x; toes 62-68/>i.

Cephalodella inquilina is common among sphagnum and sub¬

merged plants in soft-water ponds ; we have collected it in Vilas and

Oneida Counties, Wisconsin, and around Atlantic City, New Jer-

Barring (& Myers — Rotifer Fauna of Wisconsin — II. 503

sey. It is a very rapid swimmer, readily recognized by its large

size and the peculiar, clawed toes.

CEPHALODELLA LICINIA Myers, new species.

Plate XXXIV, figure 4.

The body is elongate, slender and somewhat prismatic. The

head is a little longer than wide and obliquely truncate anteriorly.

The neck is not strongly marked. The abdomen is very slightly

arched dorsally, almost parallel-sided; the lorica is fairly rigid

and the plates well marked ; the lateral clefts are rather narrow

anteriorly and increase in width towards the posterior end. The

foot is conical and moderately large ; the tail is small and rounded

and unusually far back on the foot. The toes are long and slender ;

they are nearly parallel-sided and straight for about three fourths

of their length; the terminal portion is strongly decurved and

tapers to very slender, bristle-like points ; their length is about one

fourth of the total length. The foot glands are small and pyri¬

form.

The corona is oblique and slightly convex without projecting

lips.

The mastax is relatively small and weak ; the fulcrum is slightly

expanded at the posterior end and the manubria are slender, rod¬

like and not crutched. The gastric glands are small.

The ganglion is very long and saccate. The eyespot and retro-

cerebral organ are absent.

Total length 105/>t; toes 28^.

C ephalodella licinia is found in small numbers among Fontinalis

novae-angliae in bog pools at Bargaintown, near Atlantic City,

New Jersey.

CEPHALODELLA VACUNA Myers, new species.

Plate XXXV, figure 6.

The body is elongate, fairly slender, nearly parallel-sided and

somewhat prismatic. The head is long and obliquely truncate

anteriorly. The neck is well marked. The abdomen increases

very slightly in size towards the posterior end and is faintly curved

dorsally ; the lorica is flexible and the plates well marked ; the

lateral clefts are fairly wide anteriorly and become more so towards

the posterior end. The body is obliquely truncate posteriorly ; the

504 Wisconsin Academy of Sciences, Arts, and Letters,

foot is large and conical, projecting slightly beyond the small,

rounded tail. The toes are very long, faintly recurved and slender,

tapering gradually to acute points; their length is more than one

fourth of the total length. The foot glands are small.

The corona is convex and strongly oblique without projecting

lips.

The mastax is large and of the normal type; the fulcrum is

slightly expanded posteriorly and the manubria strongly crutched.

The ganglion is large and elongate saccate. Neither eyespot

nor retrocerebral organ are present.

Total length 220/x ; toes 62/x.

Cephalodella vacuna has been collected in small numbers among

submerged sphagnum in a pond with soft, acid water at Gravelly

Run, near Atlantic City, New Jersey.

CEPHALODELLA SPECIOSA Myers, new species.

Plate XXXIV, figure 6.

The body is moderately elongate and tapers gradually to the

base of the toes. The head is very large; its dorso-ventral width

is greater than the width of the abdomen. The neck is well marked.

The abdomen tapers rapidly from the neck to the base of the foot ;

the plates of the lorica are moderately flexible and separated by

well marked lateral clefts. The foot is of normal length; the tail

is small and knoblike and does not quite reach the posterior end

of the foot. The toes are very long, slender, tapering and very

slightly decurved ; near the blunt tip is a transverse septum, giving

the toes an appearance of being clawed; their length is but little

less than half the length of the body.

The corona is oblique and without projecting lips.

The mastax is large and of normal type ; the fulcrum is crutched

and the manubria slender and rodlike.

The ganglion is large and saccate ; neither eyespot nor retrocere¬

bral organ are present.

Total length 145-155/a; toes 45-47/x.

Cephalodella speciosa is found in a large pond at Oceanville, New

Jersey, among Biccia and floating sphagnum in soft, acid water.

Earring & Myers — Rotifer Fauna of Wisconsin — II, 505

CEPHALODELLA CUNEATA Myers, new species.

Plate XXIX, figure 2.

The body is fairly short and conical, tapering gradually from

the corona to the base of the toes. The head is very large, nearly

half the length of the entire body and considerably wider than the

abdomen. The neck is indicated by a slight constriction. The

abdomen is very short and tapers evenly and rapidly to the base

of the foot; the lorica is very flexible, but the plates are well

marked; the lateral clefts are fairly wide and flaring at the pos¬

terior end. The foot is relatively long and conical and rather small

at the base ; the tail is rudimentary. The toes are very long, slender

and decurved, slightly expanded at the base and gradually taper¬

ing to acute, bristle-like points; their length is one third of the

total length. The foot glands are very small and pyriform.

The corona is strongly oblique and convex with prominent, beak¬

like lips.

The mastax is huge; the fulcrum is nearly half as long as the

entire body ; the manubria are very slender and not crutched.

The ganglion is elongate and saccate ; neither eyespot nor retro-

cerebral organ are present.

Total Ingth 105/x ; toes 32/x.

Cephalodella cuneata is fairly common among sphagnum in a

shallow ditch with soft, acid water, about five miles north of Egg

Harbor, New Jersey. It has considerable resemblance to 0. speciosa,

but is much smaller, the head is relatively larger and the toes

quite different.

CEPHALODELLA HYALINA Myers, new species.

Plate XXXII, figure 4.

The body is elongate, fairly slender and slightly compressed later¬

ally. The head is large and strongly oblique anteriorly. The neck

is well marked. The abdomen is very nearly parallel-sided, slightly

rounded dorsally at the extreme posterior end; the integument is

very flexible and the body virtually illoricate; the edges of the

plates are very obscure and difficult to trace; the lateral clefts

appear to be narrow and parallel-sided. The foot is fairly large

and robust; the tail is prominent and near mid-length. The toes

are long, slender, wide apart at the base, recurved and slightly

tapering, with conical points formed by an abrupt downward

506

Wisconsin Academy of Sciences, Arts, and Letters.

curvature of the dorsal edge; at the apex is a short, bristle-like

nib forming a continuation of the ventral edge; the length of

the toe is about one fourth of the total length. The foot glands

are large and pyriform.

The corona is strongly oblique and decidedly convex without

projecting lips.

The mastax is large and of the normal type; the fulcrum ends

in a triangular expansion and the manubria are strongly crutched.

The ganglion is very long and saccate ; eyespot and retrocerebral

organ are absent.

Total length 200-215/^ ; toes 45-50)a.

Cephalodella hyalina is not rare in ponds with soft, acid water;

we have collected it among submerged sphagnum in Lake Kathan

and Starvation Lake, Oneida County, Wisconsin, and in Fontinalis

novae-angliae at Bargaintown, near Atlantic City, New Jersey.

CEPHALODELLA PAPILLOSA Myers, new species.

Plate XXXIV, figure 7.

The body is relatively short, laterally compressed and strongly

gibbous dorsally. The head is fairly large and obliquely truncate

anteriorly. The neck is deeply constricted. The abdomen in¬

creases rapidly in depth for about two thirds of its length and is

gently rounded posteriorly; the transverse width is only one half

of the dorso-ventral depth of the body at its highest point. The

lorica is fairly rigid and the plates well marked; the lateral clefts

are fairly wide and nearly parallel-sided throughout their length.

The foot is moderately short and broad at the base; the tail is

very small. The toes are very long, slender and slightly decurved ;

from the base they taper evenly and gradually to acute points-

their length is nearly half the length of the body.

The lateral antennae are of unusual form ; small tufts of sensory

setae are seated on small, slender conical tubules.

The corona is oblique and strongly convex without projecting

lips.

The mastax is rather large and of the normal type ; the fulcrum

is slightly expanded posteriorly ; the manubria are rodlike and not

crutched.

Earring (& Myers — Rotifer Fauna of Wisconsin — 11. 507

The ganglion is elongate saccate ; retrocerebral organ and eyespot

are absent.

Total length 135/x ; toes 43ju,.

Cephalodella papillosa is rare ; we have collected it only in a large

pond at Oceanville, New Jersey, among Riccia and floating sphag¬

num.

CEPHALODELLA EVA (Gosse) .

Plate XXXV, figure 8.

Furcularia eva Gosse, Journ. Eoyal Micr. Soe., 1887, p. 864, pi. 14, fig. 9.

— Hudson and Gosse, Eotifera, SuppL, 1889, p. 26, pi. 31, fig. 17. — Wier-

ZEJSKI, Rozpr. Akad. Umiej., Wydz. Mat.-Przyr., Krakow, ser. 2, vol. 6,

1893, p. 230, pi. 5, fig. 41. — Stenroos, Acta Soc. Fauna et Flora Fennica,

vol. 17, No. 1, 1898, p. 134. — EunnstrOM, Zool. Anz., vol. 34, 1909, p. 269.

Furcularia semisetif era Glasscott, Proc. Eoyal Dublin Soe., new ser., vol. 8,

1893, p. 55, pi. 4, fig. 2.

Proales tigridia Weber, Eev. Suisse Zool., vol. 5, 1898, p. 468, pi. 18, figs.

18-20; not Proales tigridia Gosse.

Biaschiza eva Dixon-Nuttall and Freeman, Journ. Eoyal Micr. Soe., 1903,

p. 137, pi. 3, fig. 8. — Voigt, Forschungsber. Biol. Stat. Plon, vol. 11, 1904,

p. 64. — Von Hofsten, Arkiv Zool., Stockholm, vol. 6, No. 1, 1909, p. 54.

Lucks, Eotatorienfauna Westpreussens, 1912, p. 93, fig. 26. — Sachse,

Siisswasserfauna Deutschlands, pt. 14, 1912, p. 122, figs. 237, 238. —

Montet, Eev. Suisse Zool., vol. 23, 1915,' p. 331. — Jakubski, Eozpr. Wiad.

Muz. Dzieduszyckich, vol. 1, No. 3-4, 1915, p. 20. — ^Weber and Montet,

Cat. Invert. Suisse, pt. 11, 1918, p. 142.

The body is moderately slender, strongly compressed laterally

and gibbous dorsally. The head is short, broad and obliquely trun¬

cate anteriorly. The neck is well marked. The abdomen is arched

dorsally and widest at mid-length; the lorica is very flexible, but

the plates are well marked ; the lateral clefts are very wide posteri¬

orly. The foot is large and conical and the tail minute; the foot

glands are large and pyriform. The toes are very long, nearly one

third of the total length; the proximal third is tapering and the

remainder very slender, almost bristle-like, flexible and strongly

decurved. The foot glands are large and ovate.

The corona is oblique and without projecting lips.

The mastax is large and of the normal type; the manubria are

nearly as long as the fulcrum and strongly crutched.

The ganglion is large and elongate saccate; eyespot and retro-

cerebral organ are absent.

Total length 275-285ja; toes 80-85ja.

508 Wisconsin Academy of Sciences, Arts, and Letters.

Cephalodella eva is fairly common in weedy ponds with soft'

water. We have collected it in Vilas and Oneida Counties, Wis¬

consin, and around Atlantic City, New Jersey. According to

Dixon-Nuttall and Freeman this species is very variable, especially

in the form of the toes; we have not found this to be the case:

all the specimens that we have seen are very constant, both in form

and size. We are inclined to believe that the varieties described by

them may belong to one or more different species.

CEPHALODELLA TENUISETA (Burn).

Plate XXXV, figure 7.

Furcularia tenuiseta Burn, Science Gossip, vol. 26, 1890, p. 34, text fig.

DiaschiBa tenuiseta Dixon-Nuttall and Freeman, Journ. Eoyal Micr.

Soc., 1903, p. 138, pi. 1, fig. 2.

The body is elongate, fairly slender and nearly cylindric. The

head is large, but relatively short, and obliquely truncate anteriorly.

The neck is well marked. The abdomen is unusually long and

slightly gibbous posteriorly; the lorica is very flexible and the

plates faintly marked; the lateral clefts are narrow anteriorly

and slightly wider at the extreme posterior end of the lorica. The

foot is short and bluntly conical with a small, rounded tail at mid¬

length. The foot glands are very small and pyriform. The toes

are half as long as the body, very slender, slightly recurved, and

gradually tapering to acute, conical points.

The corona is convex and slightly oblique without projecting

lips.

The mastax is large and of the normal type; the fulcrum is

slightly expanded posteriorly and the manubria rodlike, not

crutched. The oesophagus is very long and slender.

The ganglion is very long and saccate. The retrocerebral organ

and eyespot are absent.

Total length SSO-SQOju; toes 120-125ju.

Cephalodella tenuiseta was originally described from freshwater;

we have found it only among algae and detritus in brackish and

saltwater ditches around Atlantic City, New Jersey. Specimens

collected by the late C. F. Eousselet at Stowmarket, Suffolk, Eng¬

land, agree with our material in every way except in having much

longer toes. We have not been able to find any other differences

and believe that they are simply local variations of the same species.

Earring & Myers — Rotifer Fauna of Wisconsin — 11. 509

CEPHALODELLA APOCOLEA Myers, new species.

Plate XXXIII, figures 1-2.

The body is moderately elongate, nearly cylindric and slightly

compressed laterally. The head is large and relatively short. The

neck is not strongly marked. The abdomen is nearly parallel¬

sided and slightly convex dorsally; the lorica is very thin and

flexible, but the plates are well marked; the lateral clefts are

moderately wide and parallel-sided. The foot is large and robust ;

the small tail is near the posterior end. The toes are extremely

far apart at the base, cylindric and straight for about two thirds

of their length, abruptly recurved and ending in slender, sickle¬

shaped, acute points ; their length is about one fourth of the total

length. The foot glands are very large and pyriform.

The corona is oblique, moderately convex and without projecting

lips.

The mast ax is large and of the normal type, but the trophi are

slender; the fulcrum is slightly incurved at the extreme end,

but not expanded ; the manubria are delicate, slightly curved rods,

and not crutched. The gastric glands are small.

The ganglion is moderately elongate and pyriform; eyespot and

retrocerebral organ are absent.

Total length 125-135/x ; toes 32-35/x.

Cephalodella apocolea is common in weedy ponds and bogs;

we have collected it in Vilas and Oneida Counties, Wisconsin;

around Washington, District of Columbia; Atlantic and Ocean

counties. New Jersey, and Polk County, Florida; also in material

collected by Dr. H. S. Jennings around Ann Arbor, Michigan.

CEPHALODELLA STEEPTA Myers, new species.

Plate XXXV, figure 1.

? Furcularia macrodactyla Stenroos, Acta Soc. Fauna et Flora Fennica,

vol. 17, No. 1, 1898, p. 134, pl‘. 2, fig. 15.

The body is elongate, fairly slender, slightly gibbous dorsally

and faintly curved. The head is relatively long and tapers very

slightly from the neck towards the corona ; it is protected by a dis¬

tinct lorica, which falls into definite folds when the head is re¬

tracted, as in the genus Diurella and others. The neck is well

marked. The abdomen is somewhat prismatic and slightly arched

dorsally; its greatest depth is near mid-length. The plates of the

;,510 Wisconsin Academy of Sciences, Arts, and Letters.

lorica are very firm and the lateral cleft wide. The foot is large

and conical ; the tail is very small and near the middle of the foot.

The toes are very long, recurved, stout at the base and taper

gradually to acute points; their length is about two fifths of the

total length. The foot glands are large and pyriform.

The corona is slightly oblique and without projecting lips.

The mastax is moderately large and of the normal type, the ful¬

crum and manubria rodshaped and slightly curved, but not

crutched. The gastric glands are large and usually of a deep pink

color.

The ganglion is fairly large and saccate. Neither eyespot nor

retrocerebral organ are present.

Total length 100-110/x, ; toes 40-45ja.

Cephalodella strepta is not rare in weedy ponds with soft, acid

water. We have collected it at Eagle River, Vilas County, Wiscon¬

sin, and around Atlantic City, New Jersey. It swims very slowly

and deliberately, nearly always in a straight course, so that, when

once seen, it is easily recognized in spite of its minute size; its

behavior suggests the movements of a mechanical toy rather than

the erratic course of other rotifers. It is possible that Stenroos’s

Furcularia macrodactyla may have been described from speci¬

mens of this species ; according to his description and figure his ani¬

mal differs in the form of the body and foot, as well as in its

greater size and relatively longer toes.

CEPHALODELLA MUCEONATA Myers, new species.

Plate XXXVI, figures 2-4.

The body is elongate, slender and slightly gibbous dorsally. The

head is moderately long and obliquely truncate anteriorly. The

neck is well marked. The abdomen is protected by a very rigid

lorica, extending over the foot and projecting beyond it ; the longi¬

tudinal clefts are deep and terminate at the foot sheath, which has

a triangular ventral point and a stout dorsal spine, separated by

a deep, rounded sinus. The foot does not project beyond the ven¬

tral point of the foot sheath. The toes are nearly as long as the

body, unusually slender, slightly recurved and taper very grad¬

ually to acute points.

The corona is slightly oblique without projecting lips.

Harring & Myers — Rotifer Fauna of Wisconsin — II. 511

The mastax is of the normal type of the genus ; the inner edge^

of the rami are finely denticulate ; the fulcrum is slightly expanded

posteriorly and the rami are crutched.

The ganglion is large and saccate ; there is a small retrocerebrai

sac, curving over the posterior end of the ganglion; the duct ap¬

proaches the corona, but does not quite reach it. There is no

eyespot.

Total length 265-275/x; toes 120-125ja; trophi 36/x.

Cephalodella mucronata is not rare in weedy, soft-water ponds;

we have collected it in Vilas and Oneida Counties, Wisconsin,

around Atlantic City, New Jersey, and in Polk County, Florida.

It was first found by the late James Murray in preserved material

brought home from New Zealand; the contracted specimens, one

of which he sent to us, were believed by him to be Monommata

appendiculata Stenroos.

CEPHALODELLA PARASITICA (Jennings).

Plate XXXVI, figure 6.

Pleurotrocha constricta Jennings, Bull. Michigan Fish Comm., No. 3, 1894,

p. 14; not Pleurotrocha constricta Ehrenberg.

Pleurotrocha parasitica Jennings, Bull. U. S. Pish Comm., vol. 19 (for

1899), 1900, p. 84, pi. 16, figs. 13, 14. — Be Beauchamp, Bull. Soc. Zool.

France, vol. 30, 1905, p. 117, figs. 1, 2; Arch. Zool. Exper., ser. 4, vol. 10,

1909, p. 202.

Piaschiza parasitica Harking, Bull. 81 U. S. Nat. Mus., 1913, p. 34.

The body is fairly slender, distinctly curved and gibbous dor-

sally. The head is unusually long and tapers from the neck towards

the corona. The neck is well marked. The abdomen increases grad¬

ually in width for about two thirds of its length and from this

point tapers rapidly to the base of the foot. The integument is

very flexible and there is no trace of the usual fissured lorica. The

foot is short and conical, its base somewhat smaller than the pos¬

terior end of the abdomen; the tail is rudimentary. The toes are

very slightly decurved; the anterior half is nearly cylindric and

the posterior tapers gradually to acute points ; their length is about

one sixth of the total length. The foot glands are minute and

virtually atrophied.

The corona is strongly oblique and without projecting lips.

The mastax is large and the trophi of normal type; two large

salivary glands are attached to its posterior end.

512 Wisconsin Academy of Sciences, Arts, and Letters,

The ganglion is relatively small and saccate; neither eyespot

nor retrocerebral organ are present.

Total length 200/x; toes 35/^.

Cephalodella parasitica is usually found attached to the cuticle

of various oligochaetes (Stylaria, Chaetogaster). There is no rea¬

son for considering it a true parasite; it leaves the host readily

and swims as well as other members of the genus; in all prob¬

ability the temporary attachment is simply a method of trans¬

portation. We have collected this species in shallow bays of Mamie

Lake, Vilas County, and Lake Kathan, Oneida County, Wisconsin.

CEPHALODELLA EUPODA Myers, new species.

Plate XXXVI, figure 7.

The body is moderately slender and gibbous dorsally. The

head is large, somewhat longer than wide. The neck is well

marked. The abdomen increases gradually in width for about half

its length, and from this point tapers towards the base of the foot,

which is marked by a slight constriction. The integument is very

flexible and there is no trace of the lorica. The foot is large and

conical; the minute tail is somewhat beyond mid-length. The toes

are blade-shaped, short and slightly decurved ; they increase

slowly in width for nearly half their length, then decrease more

rapidly to conical, acute points about one third the length of the

toes ; their entire length is one seventh of the total length.

The corona is very slightly oblique and without projecting lips.

The mastax is fairly large and of normal type; the manubria

are somewhat stouter than usual and bent at an obtuse angle

near mid-length. The oesophagus is very long and convoluted.

The ganglion is large and saccate; there is no eyespot and no

trace of the retrocerebral organ.

Total length ISO^u,; toes 22/x.

A few specimens of Cephalodella eupoda were collected some

years ago at Four Mile Kun, near Washington, District of Co¬

lumbia; it has not been found elsewhere.

CEPHALODELLA LIPAEA Myers, new species.

Plate XXXVI, figure 5.

The body is extremely short, stout and nearly cylindric. The

head is enormous ; its length is nearly haK the length of the body.

Harring & Myers — Rotifer Fauna of Wisconsin — 11, 513

The neck is marked by a slight constriction. The abdomen is very

short and abruptly truncate posteriorly; the lorica is very flexible,

and the dorsal and ventral plates separated by a very wide cleft.

The foot is indistinct and the tail very small. The toes are short,

stout and blade-shaped; the dorsal edge is decurved and the

ventral nearly straight, with a slightly decurved, clawlike tip ;

their length is about one sixth of the total length of the body.

The corona is slightly oblique without projecting lips.

The mastax is huge, but of the normal type of the genus; the

fulcrum is slightly expanded and decurved at the posterior end;

the manubria are slender and rodlike. The total length of the

mastax is fully half the length of the body.

The ganglion is very large and saccate; there is no eyespot and

no trace of the retrocerebral organ.

Total length 130-140/x; toes 22-24/x.

Cephalodella lipara was collected among floating sphagnum in

a ditch with soft, acid water, about five miles north of Egg Harbor,

New Jersey. It has a superficial resemblance to C. physalis, but is

readily distinguished by the short toes, very stout body and the

absence of the eyespot.

Genus DORYSTOMA Harring and Myers.

Notommatid rotifers with short, stout, gibbous, illoricate body,

with a distinct constriction between head and abdomen; the foot

is very short and apparently two-jointed; the toes are short and

decurved ; at the base of the foot there is a short spine.

The corona is oblique and consists of a marginal wreath of cilia

with lateral, auricle-like tufts of cilia adapted to propulsion.

The mastax is a specialized form of the virgate type ; the trophi

are modified as supports for the walls and a highly specialized

epipharynx is present and serves to pierce the body wall of the

prey by projection through the mouth.

The eyespot is single and at the posterior end of the ganglion

Type of the genus. — Dory stoma caudata (Bilfinger) = Proales

caudata Bilfinger.

This species was listed in volume twenty as a member of the

Wisconsin rotifer fauna on the strength of a preliminary identi¬

fication, which later proved to be incorrect. However, enough is

514 Wisconsin Academy of Sciences, Arts, and Letters.

known of this species to warrant the provision of a separate genus

for it. The external form has been well described by Bilfinger,

Voigt and Lie-Pettersen, and the mastax by De Beauchamp.

Genus ROUSSELETIA Harring.

Notommatid rotifers with short, gibbous, illoricate body, with

a slight constriction separating head and abdomen; the tail is

large and collar-like, surrounding the base of the long, un jointed

foot; the two toes are short and conical.

The corona is slightly oblique; the marginal cilia are short,

except on two lateral, auricle-like areas; on the unciliated apical

plate are two slightly decurved papillae; the mouth is near the

ventral edge.

The mastax is virgate and very large ; the fulcrum expands

fan-wise towards the posterior end and is in the transverse plane

of the body; the rami are large and dome-shaped, without mar¬

ginal denticulations ; the mallei are simple, curved rods with a

ventral spur; unci are absent; a rodshaped epipharynx, decurved

at the ends, is imbedded in the anterior wall of the mastax.

The retrocerebral sac is large and pyriform ; no subcerebral

glands are present. The eyespot is cervical.

Type of the genus. — Bousseletia corniculata Harring.

ROUSSELETIA CORNICULATA Harring.

Plate XXXVII, figures 1-4.

'Bousseletia corniculata Harring, Proc. U. S. Nat. Mus., vol. 46, 1913, p. 393,

pi. 37, figs. 1-3.

The body is short, moderately stout and gibbous dorsally; its

greatest width is nearly one third of the total length. The in¬

tegument is very flexible, but the outline is quite constant. The

entire body is colored green by symbiotic zoochlorellae.

The head and abdomen are separated by a well marked con¬

striction. The width of the head segment is nearly equal to its

length, about two thirds of the greatest width of the body. The

abdomen increases gradually in width for about two thirds of its

length and is rounded posteriorly; a short, sleeve-like tail sur¬

rounds the base of the foot. The dorsal surface is marked with

faint longitudinal striations, continued about half way down the

sides. The foot is relatively long, slightly tapering and without

Harring & Myers — Rotifer Fauna of Wisconsin — II, 515

joints. The toes are short, conical and acutely pointed; their

length is about one twelfth of the total length. Only the ventral

half of the truncate posterior end of the foot is occupied by the

toes ; on the dorsal margin is a minute papilla with a few long setae.

The dorsal antenna is a small, setigerous pit in the normal

position; the lateral antennae are on minute tubules, only a short

distance above and in front of the tail.

The corona is very slightly oblique ; the circumapical band of

cilia is without any dorsal gap and has laterally two strongly

ciliated, auricle-like areas. The buccal field is evenly covered

with very short cilia. On the unciliated apical plate are two

conspicuous, slightly decurved papillae, resembling those of the

genus Ploesoma. The mouth is near the ventral margin of the

corona.

The mastax is virgate and almost half as long as the body

proper. The fulcrum is narrow at the base and its sides gradually

diverge towards the posterior end, enclosing a thin, ovate lamella

with fanshaped striations; it is unique on account of being in the

transverse plane of the body. The rami are large and roughly

triangular in ventral view; the edges are curved towards the

dorsal side, so that a roughly dome-shaped cavity is formed. The

mallei are simple, strongly curved rods, with a short spur near

mid-length; there is no trace of the unci. The epipharynx is rod¬

shaped, with decurved ends. The piston is very large and fills

the entire cavity of the mastax.

The oesophagus is slender and moderately long. There is no

constriction between the stomach and intestine. The gastric

glands, ovary and bladder are normal. The foot glands are long

and clubshaped.

The retrocerebral organ consists of a large, pyriform sac, which

never contains bacteroids; there are no subcerebral glands. The

ganglion is large and saccate, and the small eyespot is at the pos¬

terior end.

Total length 130/x; toes 10/x; trophi 40/x.

Rousseletia corniculata occurs in weedy ponds, but never in large

numbers. It seems to be widely distributed; we have collected it

at Washington, District of Columbia, at Atlantic City, New Jersey,

in Oneida and Vilas Counties, Wisconsin, in Fairmount Park,

Philadelphia, Pennsylvania, and Polk County, Florida.

516 Wisconsin Academy of Sciences, Arts, and Letters,

Genus TYLOTROCHA Harring and Myers.

Notommatid rotifers with spindle-shaped, illoricate body, with

a distinct constriction between the head and abdomen ; two lateral,

knoblike, retractile processes near mid-length; the tail is rudi¬

mentary, the foot fairly long and unjointed; the toe is single,

formed by the fusion of two originally separate toes. The dorsal

antenna is double.

The corona is slightly oblique and has a marginal row of mod¬

erately long cilia and two lateral, auricle-like tufts of long cilia;

the buccal field is sparsely ciliated; on the apical plate are two

unciliated, retractile elevations without definite form.

The mastax is a specialized form of the virgate type; all the

normal elements are fused into a dome-shaped structure, serving

for the support of the walls of the mastax during the pumping

action.

At the posterior end of the ganglion is a lenticular pigment

body, which is probably a rudimentary retrocerebral sac enclosing

the eyespot.

Type of the genus. — Tylotrocha monopus (Jennings) = Notom-

mat a monopus Jennings.

TYLOTROCHA MONOPUS (Jennings).

Plate XXXVII, figures 5-8.

Notommata monopus Jennings, Bull. Michigan Fish Comm., No. 3, 1894,

p. 14, figs. 5, 6; No. 6, 1896, p. 87.

Tylotrocha monopus Harking and Myers, Trans. Wisconsin Acad. Sci., vol.

20, 1922, p. 555.

The body of this peculiar species is spindle-shaped and slender;

its greatest width is about one fourth of the total length. The

integument is very flexible and the outline varies constantly in re¬

sponse to the contractions of the animal. The entire body is of a

brilliant, translucent reddish-purple or crimson color.

The head and abdomen are separated by a shallow constriction;

there is no distinct transverse fold. The head segment decreases

slightly in width towards the neck; its length is nearly equal to

the width. The abdomen is fusiform, largest in the middle and

tapering to the rudimentary tail. Near the middle are two knob¬

like lateral elevations, retractile at the will of the animal and con¬

stantly changing in form. The foot is relatively long and conical,

continuing the outline of the body ; it is without joints, but deeply

Earring & Myers — Rotifer Fauna of Wisconsin — II. 517

wrinkled. The toe is single^ slender and conical, slightly reduced

at the base and apparently formed by the fusion of the two normal

toes, as there are two well developed mucus glands; its length is

one twentieth of the total length.

There are two dorsal antennae, slightly elevated papillae with

a minute tuft of sensory setae ; they are on the posterior part of

the head and about 15/i, apart. The lateral antennae are in the

normal position and very small.

The corona consists of a circumapical band of moderately long

cilia ; laterally there are two tufts of long cilia, resembling auricles.

The buccal field is evenly and somewhat sparsely ciliated; on its

upper edge there is a median tuft of long cilia or setae. The mouth

is near the ventral edge. Near the lateral margins of the apical

plate are two unciliated retractile elevations without any very

definite form ; they are constantly being thrust out and withdrawn,

thus resembling the enigmatic lateral humps ; when fully extended,

they appear to be bluntly conical.

The mastax is a highly specialized form of the virgate type; all

the normal elements are firmly fused into a roughly dome-shaped

structure. The outlines of the incus are still recognizable : a very

slender, rodshaped fulcrum and two elongate triangular rami, sep¬

arated by an elongate oval space. The mallei are no longer di¬

visible in unci and manubria ; a single, roughly semicircular lamella

with a posterior, rodlike extension representing the posterior part

of the median cell, is all that is actually present, and the ventral

margin is firmly united to the rami. The animal has not been

observed while feeding, and the operation of the mastax is un¬

known ; the sclerified framework may act simply as a support to

the walls of a pump, the piston moving in the cavity. The possi¬

bility is not excluded that it may serve as the real piston, as in the

genus Lindia, and produce the vacuum necessary for suction by

a rocking motion; this would, however, appear to require some

form of an epipharynx, and no trace of this has been found.

The oesophagus is long and slender. The stomach and intestine

are separated by a deep constriction. The gastric glands are re¬

placed by a cluster of small rounded bodies of high refractive in¬

dex, apparently enclosed in a membranous investment ; these highly

refractive globules are also found floating freely in the body cavity,

and there is always a cluster in the lateral humps; their nature

and function are unknown. The bladder is a simple expansion of

518 Wisconsin Academy of Sciences, Arts, and Letters.

the cloaca. Two well developed foot glands are present, indicating

that the single toe is of recent origin. The ovary is normal.

The ganglion is large and nearly circular; at its posterior end

is a lenticular, pigmented body which seems to be partly eyespot

and partly a rudimentary retrocerebral sac.

Total length 175-250/x; toes 18-24/x; trophi 18/x.

Tylotrocha monopus is fairly common in certain localities. We

have collected it in shallow ponds in Vilas and Oneida Counties,

and limnetic in lakes in Washburn and Sawyer Counties, Wiscon¬

sin, and Polk County, Florida; it is common in ponds around At¬

lantic City, New Jersey. Jennings found it in Lake Michigan, as

well as in inland lakes in Michigan.

Genus E.ESTICULA Harring and Myers, new genus.

Notommatid rotifers with very slender, spindle-shaped, illoricate

body, nearly cylindric anteriorly and tapering gradually from

mid-length to the base of the toes, without reduction in diameter

of the foot ; the two toes are short and usually have a bulbous en¬

largement at the base, containing a small mucus reservoir.

The corona is frontal or slightly oblique, with a marginal wreath

of short cilia and two lateral, auricle-like tufts of long cilia for

propulsion ; the buccal field is covered with short, close-set cilia.

The mastax is virgate and adapted to prehension, but still re¬

tains the pumping action unimpaired; the fulcrum is long and

slender; the rami are triangular and nearly symmetrical, with a

right-angled bend near mid-length; the unci have a single well

developed tooth; the epipharynx is rudimentary or absent. Two

large salivary glands are present.

The retrocerebral organ consists of a small, rounded, ductless

sac. The eyespot, when present, is a loosely aggregated mass of

red pigment granules, diffusing among the vacuoles of the sac.

Type of the genus. — Besticula melandocus (Gosse) = Furcularia

melandocus Gosse.

RESTICUIiA MELANDOCUS (Gosse).

Furcularia melandocus Gosse, Journ. Royal Micr. Soc., 1887, p. 2, pi. 1,.

fig. 4. — Hudson and Gosse, Rotifera, Suppl., 1889, p. 27, pi. 31, fig. 18.

— Bilfinger, Jahresh. Naturk. Wiirttemberg, vol. 50, 1894, p. 47. — VoiGT^

‘Siisswasserfauna Deutschlands, pt. 14, 1912, p. 102, fig. 189.

Notommata melandocus Harking, Bull. 81 U. S. Nat. Mus., 1913, p. 79.

Harring (& Myers — Rotifer Fauna of Wisconsin — II. 519

Eosphora melandocus Harking and Myers, Trans. Wisconsin Acad. Sci.,

vol. 20, 1922, p. 644, pi. 59, figs. 6-10.

For description of this species see volume twenty, cited above;

we have finally come to the conclusion that a new genus is neces¬

sary for this and related species.

SESTICTJIjA GELIDA (Harring and Myers).

Eosphora gelida Harring and Myers, Trans. Wisconsin Acad. Sci., vol. 20,

1922, p. 642, pi. 60, figs. 1-6.

For description of this species see volume twenty, cited above.

It is closely related to R. melandocus, especially in the structure of

the mastax.

EESTICITEA ANCEPS Harring and Myers, new species.

Plate XXXVIII, figures 5-8.

The body is elongated, spindle-shaped and very slender; its

greatest width is about one fifth of the total length. The integu¬

ment is very flexible and the outline is constantly changing in re¬

sponse to the contractions of the animal. The entire body is very

transparent.

The head and neck segments are of about equal length and width,

approximately three fourths of the greatest width of the body.

The transverse folds limiting the two anterior segments are well

marked. The abdomen is very nearly parallel-sided for half its

length ; from there it tapers very gradually to the base of the foot,

ending in an indistinct, broadly rounded tail. The foot continues

the general outline of the body without any marked reduction; it

has two joints, the proximal very large and tapering to the short

terminal joint. The toes are short and conical; their external

margins are slightly curved and the inner straight; their length

is about one twentieth of the total length.

The dorsal and lateral antennae are small setigerous papillae in

the normal positions.

The corona is slightly oblique and consists of a marginal wreath

of cilia, closed on the ventral side immediately below the mouth;

laterally there are two strong, auricle-like tufts especially adapted

to swimming. The buccal field is covered with short, close-set cilia ;

the small apical plate is unciliated.

The mastax is virgate, but of a modified type; although it still

retains the pumping action seemingly unimpaired, the unci may

520 Wisconsin Academy of Sciences, Arts, and Letters.

be used for prehension. The fulcrum is very long and slender,

slightly incurved and expanded at the posterior end for the at¬

tachment of the muscles of the piston. The rami are broadly tri¬

angular in ventral view and have large, somewhat asymmetric

alulae with a complicated system of reenforcing ribs. The basal

apophysis is prominent and very broad; behind this there is a

large oval ventral opening between the rami, continuing to the

point where the teeth of the unci normally rest. The dorsal, curved

portion of the rami is deeply striate and obtusely dentate on its

inner margin. The unci have a very small, subsquare basal plate,

traversed diagonally by a rudimentary tooth, finely striate at the

tip ; only one tooth in each uncus is functional. Five or six very

short accessory teeth are attached to the tip of the left uncus, but

none to the right. The central section of the manubrium tapers

gradually to the posterior end; the basal plate is somewhat pent¬

agonal in outline. A slender, curved rod is imbedded in the walls

of the mastax on each side just below, and parallel to, the posterior

edge of the ramus; these rods assist in the support of the mastax

during the pumping action. The salivary glands are very large

and nearly of the same size.

The oesophagus is slender and moderately long. The gastric

glands are very small. A slight constriction separates the stomach

from the intestine. The ovary and bladder are normal. The foot

glands are small and pyriform; they discharge into a very small,

spherical mucus reservoir at the base of the toes.

The ganglion is large and saccate. The retrocerebral sac is fairly

large, pyriform and vacuolate; it does not appear to contain bac-

teroids. No subcerebral glands are present. The eyespot is a

huge, lens-shaped disc attached to the posterior end of the gang¬

lion ; its diameter is two thirds of the width of the ganglion itself.

Total length 250-300/1,; toes 12-15/i,; trophi 45/^.

Besticula anceps is rare; we have only found a few specimens

in a boggy area along Helen Creek, near Mamie Lake, Vilas County,

Wisconsin. Its nearest relative is B. nyssa, but it is readily rec¬

ognized by its smaller size, the form of the corona and the eyespot,

as well as the simpler toes.

Earring (& Myers — Rotifer Fauna of Wisconsin — II. 521

HESTICULA NYSSA Harring and Myers, new species.

Plate XXXVIII, figures 1-4.

The body is elongate, spindle-shaped or subcylindric and very

slender; its greatest width is only one sixth of the total length.

The integument is very flexible and the outline changes with the

contortions of the animal. The body is transparent.

The head segment is rounded anteriorly and this portion is sep¬

arated from the head proper by a slight constriction; it corre¬

sponds to the rostrum of other Notommatids. The length of the

neck segment is a little greater than its width ; the transverse folds

limiting it anteriorly and posteriorly are well marked. The ab¬

domen is very nearly cylindric ; it is longitudinally striate and has

three indistinct, transverse folds. The tail is quite prominent,

short and very broad. The foot has two very short, broad joints

of about equal length. The toes have a large basal, bulbous en¬

largement, similar to, but smaller than R. melandocus; the poste¬

rior half is conical, acute and slightly decurved. A very faint

transverse line separates the basal bulb and the clawlike tip of

the toes ; their length is about one twentieth of the total length.

The dorsal antenna is a small setigerous papilla in the normal

position; the lateral antennae have not been observed.

The corona is virtually ventral and consists of a simple, evenly

ciliated oval area without any auricles or any cilia specialized for

swimming. The mouth is near the posterior edge.

The mastax is virgate with modiflcations adapting it for pre¬

hension. The fulcrum is long and slender, slightly incurved and

expanded at the posterior end for the attachment of the muscles

of the piston. The rami are roughly triangular in ventral view

and have large asymmetric alulae. The basal apophysis is very

prominent; above this there is a large oval ventral opening be¬

tween the rami. At the apex of the rami there are three or four

very small teeth, traceable for some distance from the inner edges

of the rami as faint striae. The dorsal portion of the rami is

marked with a marginal band of fine, closely spaced, convergent

striae; near the dorsal tips there appears to be complete fusion

of the rami. The unci have two teeth each; the ventral tooth is

well developed, but not clubbed at the tip. The second tooth is

strongly curved and only half the length of the main tooth; it

rests in a well marked depression or socket in the ramus and evi¬

dently serves only as a hinge for the entire malleus. Five acces-

522 Wisconsin Academy of Sciences, Arts, and Letters.

sory teeth are attached to the tip of the left uncus ; there are none

on the right side. The basal plate of the manubria is rather small

and rounded; the central portion is long and slender, somewhat

irregularly curved and tapers gradually towards the posterior

end. The slender reenforcing rods imbedded in the walls of the

mastax of so many Notommatids are in this species fibrillate at

the ends and fused to the rami. The piston is very large and pow¬

erful and, as far as can be judged from the structure of the

mastax, the pumping function is of primary importance and pre¬

hension secondary. The salivary glands are very large and of

the same size.

The oesophagus is very long and slender. The gastric glands,

ovary and bladder are normal. There is no distinct separation be¬

tween stomach and intestine. The foot glands are small and pyri¬

form ; a minute mucus reservoir may be concealed in the basal en¬

largement of the toes.

The ganglion is large and saccate ; the spherical, ductless, vacuo¬

late retrocerebral sac is attached to its posterior end. No sub¬

cerebral glands are present. There is no true eyespot ; the anterior

portion of the sac is filled with a mass of red pigment granules,

diffusing among its vacuoles, but not enclosed in any capsule.

Total length 300-350/x; toes 14-18/x; trophi

Besticula nyssa has been found only at Bargaintown, near At¬

lantic City, New Jersey. Its nearest relatives are B. melandocus

and B. anceps; it is readily recognized by its large size, slender

body and the very simple, ventral corona, as well as the structure

of the eyespot.

Genus EOSPHORA Ehrenberg.

Notommatid rotifers with fusiform, illoricate body, nearly cyl-

indric anteriorly and tapering to a distinct tail ; the foot is usually

tubular and moderately long; the two toes are short.

The corona is frontal, with a marginal wreath and a lateral, in¬

ner arc of cilia joined to it and simulating auricles; the mouth is

at the ventral margin of the buccal field and the intra-marginal

ciliation has largely disappeared.

The mastax is virgate, but the pumping action has been partly

lost; the fulcrum is short and very broad; the rami are triangular

and bent at a right angle near mid-length, where one or two large^

blunt teeth are usually present ; the unci have a single, very large

Harring (fr Myers — Rotifer Fauna of Wisconsin — II. 523

tooth, adapted to prehension ; a rudimentary epipharynx is usually

present. There are two salivary glands, the right much larger than

the left and frequently curving under the fulcrum.

The retrocerebral organ is small, but both sac and glands are

present. The eyespot is cervical, at the posterior end of the gang¬

lion; two accessory frontal eyespots are found in some species.

Type of the genus. — Eosphora najas Ehrenberg.

The genera Eothinia and Besticula are very closely related to

Eosphora, but seem to form natural groups with the included

species agreeing in so many important characters (corona, mastax,

retrocerebral organ, form of body, etc.), that generic separation

appears advisable.

EOSPHORA THOA Harring and Myers, new species.

Plate XXXIX, figures 1-5.

The body is broad and very robust; its greatest width is one

third of the total length. The integument is very flexible and the

animal highly contractile. The entire body is hyaline.

The head and neck segments are fused and separated from the

abdomen by a well marked constriction; the corona is as wide as

the body at its widest point. The abdomen tapers gradually from

a point near mid-length to the base of the foot, and ends in a broad,

but not very prominent tail. The foot is conical and very stout;

it is without joints, but irregularly wrinkled. Its length is about

one fourth of the entire length. The two toes are heart-shaped in

dorsal view, ending in very fine points ; the ventral edge is straight,

while the dorsal is strongly curved; their length is about one fif¬

teenth of the total length. In young animals the abdomen is

faintly striate or plicate dorsally.

The corona is frontal and consists of a marginal wreath of cilia,

interrupted dorsally and passing in a curve to the lateral angles

where it is joined by an inner arc, starting also from the dorsal

gap ; from the angles the corona continues as a single band, closed

ventrally and passing immediately below the mouth. The buccal

field is evenly ciliated. On the apical plate are two large papillae,

each with a tuft of sensory setae. There is no indication of the

accessory eyespots usually present on these papillae in other species

of this genus.

524 Wisconsin Academy of Sciences, Arts, and Letters,

The dorsal antenna is on the posterior portion of the head seg¬

ment, just in front of the transverse neck fold; the distance from

the neck to the lateral antennae is about two thirds of the length of

the abdomen.

The mastax is of a modified virgate type, in which the pumping

action has become subordinate and the trophi adapted to the

seizure of prey by prehension. The fulcrum is very short and

broad. The rami are elongate, roughly triangular and strongly

curved longitudinally ; near the base there is a very large apophysis,

projecting as a double spur towards the ventral side. The inner

edges of the rami have near mid-length a blunt tooth, interlocking

with its mate on the opposite side; immediately in front of this

tooth there are some faint denticulations, and nearer the base a

strong transverse rib forms a slightly projecting knob. The poste¬

rior half of the rami are armed with about twenty small, conical

teeth, separated by relatively large interspaces. The unci have a

single, very robust, clubshaped ventral tooth, which is connected

to the upper end of the manubrium by an excessively thin lamella

of irregular outline. The central section of the manubrium is

nearly parallel-sided and slightly curved; the basal plate is tri¬

angular. The epipharynx consists of two symmetrical, roughly

li-shaped pieces, imbedded in the walls of the mastax; the shorter

branch is directed diagonally outwards and towards the ventral

side; the longer, broadly expanded posteriorly, is nearly parallel

to the longitudinal axis of the mastax.

The epipharynx has been turned through an angle of approxi¬

mately 90 degrees in figure 5, to avoid obscuring the form of the

rami; it is consequently somewhat foreshortened, and its true

length is shown in figure 4. The right lobe of the mastax is con¬

siderably larger than the left; this condition usually indicates the

presence of a salivary gland, but there are no structural re¬

mains of it.

The oesophagus is relatively short. The gastric glands are

very large and rounded. There is no constriction between the

stomach and intestine. The nuclei of the ovary are unusually

large and irregular in outline ; they appear to have separate yolk-

masses. The bladder is normal. The foot glands are nearly cylin-

dric and as long as the foot itself; they discharge into a minute,

spherical mucus reservoir at the base of the toes.

The ganglion is large and saccate. The retrocerebral organ con¬

sists of a small, vacuolate sac and two very small subcerebral

Earring c§ Myers — Rotifer Fauna of Wisconsin — 11. 525

glands. The eyespot is large and at the posterior end of the

ganglion.

Total length 300-500/x ; toes 20-35^ ; trophi 50/x long, 60/x wide.

Eosphora thoa is rare; we have found only a few specimens in

‘‘Cemetery pond,” near Eagle River, Vilas County, Wisconsin,

and at Bargaintown, near Atlantic City, New Jersey. In external

appearance it resembles E. anthadis so closely that it is virtually

impossible to separate them. However, their behavior is very differ¬

ent; E. anthadis is a very slow swimmer and generally sluggish

in its movements, while E. thoa swims very fast and is constantly

changing its direction. The eyespot is sufficient to distinguish the

species, and the trophi show but slight resemblances. It should be

noted that E. anthadis is found only in moderately hard waters,

while E. thoa occurs only in very soft water, so that there is really

no great danger of confusion.

Genus ENTEROPLEA Ehrenberg.

Notommatid rotifers with short, very stout, saccate, illoricate

body; the head is short and broad, the neck well marked; the ab¬

domen is slightly pyriform and ends in a short, sleevelike tail ; two

deep dorsal grooves at the points of attachment of the retractor

muscles; the foot is short and indistinctly two-jointed; the toes

are short and bladeshaped.

The corona is frontal and consists of a marginal wreath of strong

cilia and a transverse arc of moderately long cilia separating the

apical plate and the unciliated buccal field.

The mastax is a modified form of the virgate type, adapted to

prehension ; the fulcrum is short and broad, the rami lyrate and

very powerful, denticulate on the inner margin; the unci have a

single, very large tooth, clubbed at the point ; the manubria are

short and broad; two rudimentary salivary glands are present.

The gastric glands are very long and ribbonlike ; on the dorsal

side of the stomach are four long, tubular, glandular appendages.

The retrocerebral organ is rudimentary; both sac and glands

are present. Two eyespots are seated on small knobs on the apical

plate.

Type of the genus. — Enteroplea lacustris Ehrenberg.

526 Wisconsin Academy of Sciences, Arts, and Letters.

ENTEROPLEA LACUSTEIS Ehrenberg.

Plate XL, figures 1-5.

Enteroplea lacustris Ehrenberg, Abh. Akad. Wiss. Berlin, 1830, p. 46; in

Hemprich and Ehrenberg, Symb. Phys. Anim. Evert., Phytozoa, 1832 (?),

pi. 3, fig. IV, 11. — Barring, Bull. 81 U. S. Nat. Mus., 1913, p. 44.

Diglena lacustris Ehrenberg, in Hemprich and Ehrenberg, Symb. Phys.

Anim. Evert., Phytozoa Polypi, 1832 (?), fob h (second page); Abh.

Akad. Wiss. Berlin (for 1831), 1832, p. 136, pi. 3, fig. 10, pi. 4, fig. 14;

(for 1833), 1834, p. 335, pi. 10, fig. 2; Infusionsthierchen, 1838, p. 442,

pi. 54, fig. 4.— Dujardin, Hist. Nat. Zooph., 1841, p. 652.— Toth, Math.

Termesz. Kozl., vol. 1, 1861, p. 187, (pi. 4) fig. 32.

Triphylus lacustris Hudson, in Hudson and Gosse, Eotifera, Suppl., 1889,

p. 19, pi. 32, fig. 16. — ^Western, Journ. Quekett Micr. Club, ser. 2, vol.' 4,

1890, p. 107, pi. 10, fig. 1; 1892, p. 374, pi. 25, fig. 5. — Bilfinger, Jahresh.

Ver. Naturk. Wiirttemberg, vol. 48, 1892, p. 113. — Wierzejski, Eozpr.

Akad. Umiej., Wydz. Mat.-Przyr., Krakow, ser. 2, vol. 6, 1893, p. 224.

— Kertesz, Budapest Eotat. Faun.. 1894, p. 28. — Jennings, Bull. U. S.

Fish Comm., vol. 19 (for 1899), 1900, p. 84 — Voigt, Forschungsber. Biol.

Stat. Plon, vol. 11, 1904, p. 40; Siisswasserfauna Deutschlands, pt. 14,

1912, p. 110, figs. 207, 208. — Voronkov, Trudy Hidrobiol. Stants. Glu-

bokom Oz., vol. 2, 1907, p. 95. — De Beauchamp, Arch. Zool. Exper., ser.

4, vol. 10, 1909, pp. 24, 228, 341, figs. II A, XXXVI, LVII, pi. 9, figs.

124—131. — Lucks, Eotatorienfauna Westpreussens, 1912, p. 47. fig. 6.

—Weber and Montet, Cat. Invert. Suisse, pt. 11, 1918, p. 124, fig. 35.

The body is short, saccate and very stout; its greatest width is

nearly two thirds of the length of the body proper. The integu¬

ment is very soft and flexible, but the outline remains quite con¬

stant. The entire body is very hyaline.

The head is short and very broad; it is separated from the ab¬

domen by a well defined constriction or neck immediately behind

the mastax. The abdomen is slightly pyriform, ending posteriorly

in a short, sleeve-like tail surrounding the base of the foot ; postero-

dorsally it has two parallel, deep, V-shaped grooves, caused by the

pull of the dorsal retractor muscles, the broadened ends of which

are attached to the internal apex of the V. The foot is indistinctly

two- jointed, tapering and rather short, about one fifth of the length

of the body ; the terminal joint is nearly twice as long as the

basal joint. The toes are nearly cylindric for one half of their

length, with conical, acute points; their length is about one six¬

teenth of the total length.

The dorsal antenna is a small, setigerous papilla in the normal

position ; the lateral antennae are quite close together, immediately

in front of and above the tail.

Harring & Myers — Rotifer Fauna of Wisconsin — II. 527

The corona is frontal and consists of a marginal wreath of

strong cilia, interrupted dorsally and closed behind the mouth,

which is near the ventral edge. The buccal field is in this species

unciliated and somewhat depressed; its margin is indicated by a

horseshoe-shaped band of cilia, separating it dorsally from the

apical plate.

The mastax belongs to the virgate type, but is somewhat modi¬

fied, as the animal is carnivorous and the mallei have become

adapted to the seizure of prey by prehension through the mouth

opening. The rami are lyrate in ventral view and have prominent

basal apophyses for the attachment of the abductor muscles; lat¬

erally they are expanded into broad, thin, somewhat asymmetric

lamellae. Near mid-length they are bent at a right angle; this

median section is lamellar and denticulate on its inner edge, with

fine striae continuing for some distance from the base of the den¬

ticles. At the apex of the angle there is on each ramus a strongly

developed tooth, fitting into a groove on the opposite side. The

dorsal ends of the rami form two powerful, curved, opposing teeth,

separated from the central, denticulate section by an elongate oval

opening. The fulcrum is a moderately long, broad lamella, ex¬

panded at the base of the rami and nearly parallel-sided poste¬

riorly. The unci have a single, powerful tooth, somewhat curved

and clubbed at the point, and a small accessory tooth, only half the

length of the main tooth; the basal plate is roughly triangular.

The manubria have a subsquare basal plate ; the central section is

continued as a nearly straight, slightly compressed rod, curved to¬

wards the ventral side at the extreme end. The epi pharynx con¬

sists of two triangular plates, bent at a somewhat obtuse angle

along the median line, and projecting forward at a right angle

to the rami at the anterior apex. The piston is very large and

almost completely fills the cavity of the mastax. There are two

rudimentary salivary glands in the ventral angles of the mastax.

The oesophagus begins high up on the dorsal side of the mastax ;

its walls are muscular and may be greatly distended to allow the

passage of entire animals into the stomach. The gastric glands

are long and ribbon-like; their ends are bifurcate. In addition

to the glands there are four very long, slender gastric appendages,

no doubt glandular in nature ; two of these are a short distance

below the gastric glands, the others near the posterior end of the

stomach. There is no constriction between the stomach and in-

528 Wisconsin Academy of Sciences, Arts, and Letters,

testine. The bladder is large. The foot glands are very long and

slender, extending almost the entire length of the foot.

The ganglion is relatively small and saccate. The retrocerebral

sac is rudimentary and fused to the lower surface of the ganglion.

Two small subcerebral glands are present; in the young animals

they contain bacteroids and are consequently opaque, but v/ith

age they gradually become transparent. The two eyespots are on

small knobs on the apical plate, a short distance below and outside

the arc of cilia limiting the buccal field dorsally.

Total length 500-600/x; toes 30-35/^; trophi 70/x long, 60/x wide.

Enteroplea lacustris is widely, but somewhat sporadically dis¬

tributed; where it does occur, it is usually present in enormous

numbers. It is very often found in company with Epiphanes

{= Notops) clavnlata, although they are not to be considered in¬

separable.

Genus EOTHINIA Harring and Myers.

Notommatid rotifers with moderately elongate, fusiform, illori-

cate body, a distinct neck segment separating head and abdomen;

the tail is moderately large or rudimentary, the foot two- jointed

and short ; the toes are rather short.

The corona is obliquely frontal and may have a ventral chin;

the marginal cilia are fairly short except on two lateral, auricle¬

like areas.

The mastax is virgate, but somewhat specialized ; the inner*' edges

of the rami are provided with numerous short, closely spaced,

needlelike teeth ; the piston is well developed.

The retrocerebral organ is well developed; both sac and glands

are present; there is a cervical eyespot at the posterior end of the

ganglion and two frontal eyespots on the apical plate.

Type of the genus. — Eothinia elongata (Ehrenberg) ^ Eosphora

elongata Ehrenberg.

EOTHINIA TEIPHAEA Harring and Myers, new species.

Plate XLI, figures 1-5.

The body of this species is fusiform and moderately elongate;

its greatest width is about one fourth of the total length. The in-

Harring & Myers- — Rotifer Fauna of Wisconsin — II. 529

tegument is very flexible and the outline changes according to the

contractions of the animaL The body is very transparent.

The transverse folds limiting the head and neck segments are

unusually deep. The head segment is broadest anteriorly and

tapers towards the neck; its width is but little less than the width

of the abdomen. The neck segment is a little narrower than the

head, but approximately the same length, slightly more than half

the width. The abdomen increases very slightly in width for about

half its length and then tapers gradually to the base of the foot;

the tail is represented by a very slight dorsal elevation. The two

short foot joints continue the outline of the abdomen without any

sudden reduction in width. The two toes are very long and slender,

slightly decurved and incurved ; they have a nearly hemispherical,

slightly compressed, bulbous enlargement at the base. The length

of the toes is about one seventh of the total length.

The corona extends down on the ventral side about one fourth

of the length of the body; the post-oral portion projects as a

minute chin. The unciliated apical plate is strongly convex; the

buccal field has a well marked, troughlike median depression in

which the mouth is located. The marginal cilia are short except

on two lateral, auricle-like areas.

The dorsal antenna is a short, stubby boss with a central depres¬

sion, at the center of which is a tuft of sensory setae. The lateral

antennae have not been observed.

The mastax is virgate and resembles closely the type of the

genus, but in the development of the unci it is a little nearer the

normal Notommata-type, The fulcrum is extremely broad at the

base and tapers somewhat abruptly to a slender, rodlike, slightly

incurved posterior section. The rami are broadly triangular and

nearly symmetrical; their inner edges are provided with short,

close-set, needle-like teeth, beginning near the base and continuing

to the apex. The unci have a subsquare basal plate with a well

developed ventral tooth, clubbed at the tip, and a rudimentary

second tooth, beginning near the base of the ventral tooth and

crossing the basal plate diagonally to its dorsal edge. Two short

and very slender accessory teeth are attached to the ventral edge

of the principal tooth in each uncus. The basal plate of the manu¬

brium is large and subsquare ; the posterior portion is fairly stout

and decurved at the tip. Two slender, slightly curved rods are

imbedded in the walls of the mastax below the posterior edges of

530 Wisconsin Academy of Sciences, Arts, and Letters.

the rami ; they serve as supports during the pumping action. The

piston is well developed.

The oesophagus is slender and moderately long. The stomach

and intestine are not separated by any constriction. The gastric

glands and ovary are normal. The cloaca appears to function as a

bladder. The foot glands are large, nearly circular and slightly

compressed; they discharge into a small, spherical mucus reser¬

voir, which is partly in the bulbous, basal enlargement of the toe.

The ganglion is moderately large and nearly spherical. The

retrocerebral sac is pyriform and very small ; the subcerebral glands

are short and always contain a rounded mass of bacteroids at the

level of the cervical eyespot, thus producing the appearance of

three eyespots in a transverse row. There are two accessory fron¬

tal eyespots on the apical plate in addition to the cervical eyespot

at the posterior end of the ganglion.

Total length 175-250^; toes 25-35/x; trophi 28/x long, 35 wide.

Eothinia triphaea occurs in small numbers among sphagnum

growing on the margins of shallow ponds. We have found it

widely distributed in Vilas and Oneida Counties, Wisconsin, and

also in ponds around Atlantic City, New Jersey, and at Hyatts-

ville, near Washington, District of Columbia.

EOTHINIA ARGUS Harring and Myers, new species.

Plate XLI, figures 6-11.

The body of this species is fusiform and moderately elongate;

its greatest width is about one fourth of the total length. The in¬

tegument is moderately flexible and the outline fairly constant.

It is a very transparent animal.

The transverse folds limiting the head and neck segments are

well marked. The head segment is very broad anteriorly and

tapers towards the neck; its width is equal to the width of the

abdomen. The neck segment is considerably narrower than the

head. The abdomen is nearly cylindric for three fourths of its

length and rounded posteriorly. The tail is small and three-lobed ;

the median lobe is subsquare and the lateral lobes rounded. The

foot has two short joints of nearly equal length. The toes are

slender, conical, acutely pointed and slightly decurved ; their length

is about one fourteenth of the total length.

The corona is slightly oblique and terminates a short distance

behind the mouth. The apical plate is strongly convex and un-

Harring & Myers — Rotifer Fauna of Wisconsin — II. 531

ciliated; the buccal field is covered with very short, close-set cilia

and has a median depression around the mouth. The marginal

cilia are short except on two laterally projecting, auricle-like areas.

The dorsal and lateral antennae are small setigerous papillae in

the normal positions.

The mastax is virgate and slightly asymmetric. The fulcrum is

long and broad at the base, tapering gradually to the slightly in¬

curved posterior end. The rami are roughly triangular and have

moderately large alulae. There is a heartshaped opening just

above the fulcrum; this is followed in the right ramus by four

small teeth, two strong, apical teeth and two small teeth on the

dorsal portion; the left ramus has five small teeth above the ven¬

tral opening and one large, apical tooth, followed by two small

dorsal teeth. The rami are prolonged dorsally by two thin, blade¬

like curved pieces, not observed in the mastax of any other Notom-

matid. The unci have a single, large tooth with two small acces¬

sory teeth attached to the clubbed tip on the ventral side; the

basal plate is irregularly oval and has a narrow, denticulate por¬

tion adjoining the tooth on the dorsal side. The basal plate of the

manubria is large ; the posterior branch is nearly straight and has

a slight terminal expansion. Two slender, double-curved rods are

imbedded in the walls of the mastax just below the posterior edges

of the rami; they serve as supports during the pumping action.

The piston is large and muscular. On the left side is a granular

area which appears to represent an atrophied salivary gland.

The oesophagus is very long and slender. There is no constric¬

tion between the stomach and intestine. The gastric glands and

ovary are normal. The cloaca functions as a bladder. The foot

glands are pyriform and very small.

The ganglion is large and saccate. The retrocerebral organ con¬

sists of a very small, pyriform sac and two short subserebral glands,

which contain a rounded mass of bacteroids at the level of the eye-

spot. There are two accessory frontal eyespots on the apical plate

in addition to the cervical eyespot at the posterior end of the

ganglion.

Total length 250-300ju,; toes 18-22jtA; trophi 38/x.

Eothinia argus was collected in small numbers in Lenape Lake,

at Mays Landing, New Jersey. It is very closely related to

E. triphaea, but is readily distinguished by the much shorter toes

and the long retrocerebral sac.

532 Wisconsin Academy of Sciences, Arts, and Letters.

Genus SPHYRIAS Harring.

Notommatid rotifers with short, broad, saccate, illoricate body;

the head is very broad and truncate anteriorly and has two short,

stumplike lateral projections; it is separated from the abdomen by

a well marked constriction ; the abdomen is subsquare and ends in

a collarlike tail; the foot is long and wrinkled; the toes are short,

conical and obtusely pointed.

The corona consists of two roughly circular wreaths of long cilia

at the edges of the short, lateral projections of the head; the

anterior surface of the head is unciliated; the mouth is near the

ventral margin and strongly protrusile.

The mastax is virgate; the rami are roughly triangular and

strongly curved, the inner margins armed with numerous short,

closely spaced, needlelike teeth; the fulcrum and manubria are

long, straight rods; the unci have only a single functional tooth;

the piston is very large.

The retrocerebral organ is rudimentary; the two eyespots are

on the lateral projections of the head.

Type of the genus. — Sphyrias lofuana CRousselet) = Notops

lofuana Rousselet.

SPHYRIAS LOFUANA (Rousselet).

Plate XLII, figures 1-5.

Notops lofuana Rousselet, Proe. Zool. Soe. London, 1910, p. 795, pi. 75,

figs. 1-3.

Sphyrias lofuana Harking, Bull. 81 U. S. Nat. Mus., 1913, p. 96; Proe.

U. S. Nat. Mus., vol. 46, 1913, p. 400, pi. 37, figs. 4-8.

The body of this species is short, stout and truncate anteriorly;

its greatest width is about one half of the total length. The in¬

tegument is moderately flexible and the outline is constantly

changing in response to the violent contractions of the animal. It

is a moderately transparent species.

The head and abdomen are separated by a deep transverse fold.

The outline of the head is variable; when the animal is at rest (as

represented in the figures), it is broadly triangular and wider than

the abdomen; when it is swimming, the mouth region is retracted

and the head becomes squarely truncate. The abdomen is sub¬

square in dorsal view, truncate posteriorly and ends in a short,

very wide, collar-like tail, surrounding the base of the foot; it is

Earring (& Myers — Rotifer Fauna of Wisconsin — II. 533

marked dorsally with strong, longitudinal folds, gradually dis¬

appearing on the sides. The foot is fairly long and wrinkled, but

not jointed. The two toes are somewhat obtusely pointed, slightly

decurved and laterally compressed; their length is about one tenth

of the total length.

The tubular dorsal antenna is on the abdomen, immediately be¬

hind the transverse fold, and joined to the integument in its en¬

tire length. The lateral antennae are small, rounded, knoblike,

setigerous papillae just above and in front of the tail.

The corona is perhaps easier to understand if it is considered as

a simplified or specialized E osphora-QomnsL. As explained under

this genus, its corona consists of a circumapical band of cilia, in¬

terrupted dorsally, with a lateral arc of very long cilia for pro¬

pulsion, and an inner arc of fairly long cilia joined to the circum¬

apical band at both ends. In Sphyrias the dorsal gap in the corona

is almost equal to the width of the head, and all that remains of

the circumapical band is the lateral, auricle-like arc of long cilia ;

in front of this is a short arc of fairly long cilia, representing the

inner arc of Eosphora. Nothing remains of the latero-ventral arc

of the circumapical band normally joining the auricles and passing

below the mouth except a few short cilia at the sides of the ver¬

tical, slitlike mouth. The mouth regio^ is retracted when the ani¬

mal swims, and the inner arc of cilia thus forced out even with

the external or posterior arc; this produces an appearance recall¬

ing the form of the hammerhead shark {Sphyrna, in Kafinesque’s

Greek!). At the point of attachment of the dorsal longitudinal

muscles are four small tufts of sensory setae.

The mastax is virgate, but adapted also to prehension. The ful¬

crum is very long and straight; it is formed of two plates joined

together at the edges and forming a Y-shaped trough with the

apex dorsal. The rami are symmetrical and roughly triangular;

their inner edges are armed with about twenty long, needle-like

teeth. The unci have only a single, long tooth, clubbed at the tip

and with a minute basal plate. The manubria are long, rodlike

and nearly straight; they diverge from the incus at an angle of

nearly 45 degrees. Two straight rods, expanded at their dorsal

ends into triangular plates, are imbedded in the walls of the

mastax below the dorsal branch of the rami and aid in the support

of the mastax during the pumping action. The piston is large

and powerful ; its striate longitudinal muscles are very conspicuous.

534 Wisconsin Academy of Sciences, Arts, and Letters.

In the ventral angles of the mastax are two well developed salivary

glands.

The oesophagus is moderately long and slender. There is no

constriction between the stomach and intestine. The ovary is very

long and ribbon-shaped and passes dorsally over the stomach into

the head segment. The gastric glands and bladder are normal.

The foot glands are slender, slightly clubshaped and nearly as

long as the foot.

The ganglion is large and saccate. At its posterior end is a

small, spherical, ductless retrocerebral sac and two granular areas

at the external angles of the ganglion appear to represent the re¬

mains of the subcerebral glands. The two eyespots are on rounded,,

knoblike projections between the lateral arcs of cilia.

Total length 275-300/x; toes 35-30/^; trophi 62/^.

Sphyrias lofuana is not rare in weedy ponds around Washing¬

ton, District of Columbia; we have found it also near Atlantic

City, New Jersey, and at Dock Lake, near Spooner, Wisconsin.

In spite of the somewhat aberrant external form this species

is closely related to Eosphora and Eotkinia, as shown by a com¬

parison of the mastax and corona. Its food seems to consist prin¬

cipally of the smaller Bdelloids and ConocJiilus, whose trophi are

often found in the stomach.

Grenus MONOMMATA Bartsch.

Notommatid rotifers with slender, elongate ovate, spindle-shaped,

illoricate body, with a slight constriction behind the mastax, sep¬

arating the head and abdomen; the foot is very short and ob¬

scurely two-jointed; the toes are extremely long, nearly twice the

length of the body, and unequal, the right toe longer than the left.

The corona is slightly oblique and consists of a marginal wreath

of cilia with lateral, auricle-like tufts of longer cilia adapted to

propulsion; the apical plate is unciliated and the buccal field

evenly covered with short, close-set cilia; the mouth is somewhat

below the center of the corona.

The mastax is intermediate between the virgate and forcipate

type; the rami are lyrate and the inner margins armed with one

or more strong teeth immediately below the mouth opening; the

unci have three unequally developed, long, slender, clubbed teeth j

the manubria are broad and lamellar at the base; the piston is

Harring & Myers — Rotifer Fauna of Wisconsin — 11. 535

weak and plays only a subordinate part in the operation of the

mastax.

The retroeerebral organ is imperfectly developed; the sac is

ductless and the glands rudimentary or absent; the eyespot is at

the posterior end of the ganglion.

Type of the genus. — Monommata longiseta (Muller) = Yorticella

longiseta Muller.

One of the most interesting results of this survey of the notom-

matid rotifers is the discovery of the remarkable type of mastax

of this genus; a more perfect intermediate or ^‘missing link” be¬

tween the forcipate and virgate mastax than that of Monommata

grandis could hardly be imagined. If only the ventral view were

available, it would unhesitatingly be called forcipate; its real

affinities become evident at once in the lateral view, showing the

curvature of the rami found only in the virgate mastax and also

the large basal plate of the manubria. Both are developed as

supports for the walls of the pumping type of mastax. However,

the changes necessary to transform it into a perfectly good forci¬

pate mastax are small indeed.

MONOMMATA LONGISETA (Muller).

Plate XLIII, figures 1-5.

? Cercaria ortis Muller, Zool. Dan. Prodr., 1776, p. 280; Anim. Infus.,

1786, p. 138, pi. 20, fig. 7.

Yorticella longiseta Muller, Anim. Infus., 1786, p. 295, pi. 42, figs. 9, 10.

? Trichoda Mcaudata Schrank, Fauna Boica, vol. 3, pt. 2, 1803, p. 87.

? Vaginaria hrachiura Schrank, Fauna Boica, vol. 3, pt. 2, 1803, p. 144.

^ Furcocerca orMs Lamarck, Hist. Nat. Anim. sans Vert., vol. 1, 1815,

p. 448.

Furcularia longiseta Lamarck, Hist. Nat. Anim. sans Vert., vol. 2, 1816,

p. 39. — Hudson and Gosse, Eotifera, 1886, vol. 2, p. 46, pi. 18, fig. 16.

— Anderson, Journ. Asiatic Soc. Bengal, vol. 58, pt. 2, 1889, p. 53.

— Wierzejski, Eozpr. Akad. Umiej., Wydz. Mat.-Przyr., Krakow, ser. 2,

vol. 6, 1893, p. 230. — Skorikov, Trav. Soc. Nat. Kharkow, vol. 30, 1896,

p. 295. — Weber, Eev. Suisse Zool., vol. 5, 1898, p. 476, pi. 19, fig. 3.

— Hempel, Bull. Illinois State Lab. Nat. Hist., vol. 5, 1898, p. 370.

— Lie-Pettersen, Bergens Mus. Aarbog (for 1909), 1910, No. 15, p. 46.

— Mola, Ann. Biol. Lac., vol. 6, 1913, p. 245.

? Trichocerca orhis Bory de St. Vincent, Class. Anim. Mier., 1826, p. 42.

F Lecane orhis Nitzsch, Enc. Wiss. u. Kiinste, sect. 1, vol. 16, 1827, p. 68.

F BracMonus orhis Blainville, Diet. Sci, Nat., vol. 60, 1830, p. 149.

Trichocerca longiseta Blainville, Diet. Sci. Nat., vol. 60, 1830, p. 150.

536 Wisconsin Academy of Sciences, Arts, and Letters.

Notommata longiseta Ehrenberg, Abh. Akad. Wiss. Berlin, 1830, p. 46; In-

fusionsthierchen, 1838, p. 432, pi. 53, fig. 2.

Notommata longiseta aequalis Ehrenberg, Abh. Akad. Wiss. Berlin (for

1831), 1832, p. 134.

Notommata longiseta inaequalis Ehrenberg, Abh. Akad. Wiss. Berlin (for

1831) 1832, p. 134.

Notommata aequalis Ehrenberg, Abh. Akad. Wiss. Berlin (for 1831) 1832,

p. 134; Infusionsthierchen, 1838, p. 432, pi. 53, fig. 3.

Scaridium longisetum Schoch, Mier. Thiere Sussw.-Aquar., 1868, p. 30.

Monommata longiseta Bartsch, Jahresh. Naturk. Wiirttemberg, vol. 26, 1870,

p. 344. — Eyferth, Einf. Lebensf., 1878, p. 84, pi. 5, fig. 12; 1885, p. 109,

pi. 7, fig. l2. — Bergendal, Acta Univ. Lundensis, vol. 28, 1892, sect. 2,

No. 4, p. 75, pi. 1, fig. 14. — Stenroos, Acta Soc. Fauna et Flora Fennica,

vol. 17, No. 1, 1898, p. 134. — Voigt, Forschungsber. Biol. Stat. Plon,

vol. 11, 1904, p. 45; Siisswasserfauna Deutschlands, pt. 14, 1912, p. 104,

fig. 193. — Lucks, Eotarienfauna Westpreussens, 1912, p. 54. — Montet,

Rev. Suisse Zool., vol. 23, 1915, p. 324. — Jakubski, Rozpr. Wiad. Muz.

Dzieduszyckich, vol. 1, No. 3-4, 1915, p. 18. — ^Weber and Montet, Cat.

Invert. Suisse, pt. 11, 1918, p. 118.

Monommata aequalis Eyferth, Einf. Lebensf., 1878, p. 84; 1885, p. 109.

— Voigt, Siisswasserfauna Deutschlands, pt. 14, 1912, p. 104, fig. 194.

Furcularia aequalis Hudson and Gosse, Eotifera, 1886, vol. 2, p. 46, pi. 18,

fig. 5.

Monommata orlis Harring, Bull. 81 U. S. Nat. Mus., 1913, p. 72.

The body of this species is slender, elongate ovate and spindle-

shaped; its greatest width is less than one fourth of the length.

The integument is very flexible, but the outline is fairly constant.

The entire body is very transparent.

The head segment is relatively short and broad; the length is

slightly greater than the width. It is separated from the abdomen

by a well marked constriction. The abdomen is elongate and ovate,

slightly gibbous dorsally and rounded posteriorly; the integument

is covered with minute, interrupted, very closely spaced longi¬

tudinal striae. The foot is short, stout and obscurely two-jointed.

The toes are extremely long and unequal; the left toe is always

shorter than the right, but the relative length is variable, the usual

proportion being four to five, but may be as much as one to two

and individuals are occasionally found with nearly equal toes.

The basal portion of the toes is fairly stout and very nearly cylin-

dric, tapering gradually to the cylindrical, extremely slender

posterior portion, which is fully half the length of the entire toe.

The right toe is virtually straight and in the axis of the body;

the left toe curves slightly upwards and to the left.

Earring & Myers — Rotifer Fauna of Wisconsin — II. 537

The dorsal antenna is a small setigerons papilla in the normal

position; the lateral antennae are on the posterior fourth of the

body.

The corona is very slightly oblique and consists of a marginal

wreath of cilia with lateral, auricle-like tufts of longer cilia for

propulsion; the apical plate is unciliated and the buccal field

evenly covered with short, close-set cilia.

The mastax is of a modified virgate type. The fulcrum is long

and slender and tapers gradually towards the posterior end. The

rami are triangular at the base and bent near mid-length at an

approximately right angle; the dorsal portion is long, slender,

tapering and slightly incurved. Each ramus has at the angle a

long, slender tooth, interlocking with its mate on the opposite side.

The right uncus has three very long, slender teeth, the posterior

resting on the tip of the ramus, the two anterior on the ramus just

below the inner tooth; the left uncus has two teeth, one resting on

the posterior tip of the ramus and the other just below the inner

tooth. The manubria are broad and lamellar, ending in a rod¬

shaped posterior portion, slightly recurved at the end. The piston

is large, but apparently not very powerful.

The oesophagus is rather short. The gastric glands are small

and rounded. There is no constriction between stomach and in¬

testine. The ovary and bladder are normal. The foot glands are

very small and pyriform.

The ganglion is moderately large and saccate. The retrocerebral

sac is small and apparently ductless; it contains a small number

of transparent, globular bodies and can not be stained. Near mid¬

length of the ganglion are two small, rounded masses representing

the subcerebral glands, indicated in the figures, but invisible with¬

out intra-vitam staining. The eyespot is at the posterior end of

the ganglion.

Total length 200-250/a; length of body 75-95/a; length of right

toe 125-155/a; left toe two thirds to four fifths of right toe;

trophi 18/a.

Monommata longiseta is abundant everywhere in weedy ponds

all over the world.

Muller ^s Cercaria orMs was undoubtedly a contracted Monom¬

mata. Accepting as fact the frequently asserted identity of the

two ^ Varieties ' in every respect except size, the species was listed

as Monommata orhis by Harring in the Synopsis of the Rotatoria

538 Wisconsin Academy of Sciences, Arts, and Letters.

in accordance with accepted rules of nomenclature. However, it is

very evident that the genus includes two perfectly distinct species

and it is not now possible to refer C. orhis definitely to one or the

other. For this reason we have used the later name longiseta, as it

unquestionably belongs to the smaller species.

MONOMMATA GRANDIS Tessin.

Plate XLIII, figures 6-10.

Monommata grandis Tessin, Arch. Naturg. Mecklenburg, vol. 43, 1890,.

p. 151, pi. 1, figs. 11, 12. — Levander, Acta See. Pauna et Flora Fennica,.

vol. 12, No. 3, 1895, p. 35.

Furcularia longiseta grandis Rousselet, Journ. Quekett Micr. Club, ser. 2,

vol. 6, 1895, p. 124, pi. 7, fig. 3.

Monommata longiseta grandis Stenroos, Acta. Soc. Fauna et Flora Fennica,

vol. 17, No. 1, 1898, p. 135. — Voigt, Forschungsber. Biol. Stat. Plon,.

vol. 11, 1904, p. 56; Siisswasserfauna Deutschlands, pt. 14, 1912, p. 104.

— Weber and Montet, Cat. Invert. Suisse, pt. 11, 1918, p. 119.

Monommata appendiculata Stenroos, Acta. Soc. Fauna et Flora Fennica,

vol. 17, No. 1, 1898, p. 135, pi. 1, figs. 33, 34.

Monommata orhis grandis Harring, Bull. 81 U. S. Nat. Mus., 1913, p. 72.

The body of this species is moderately slender, elongate ovate

and spindle-shaped; its greatest width is less than one third of

the length. The integument is rather flexible and the outline

is quite constant. The body is moderately transparent.

The head segment is moderately large and the width nearly

equal to the length. It is separated from the abdomen by a well

marked constriction. The abdomen is elongate and ovate, distinctly

gibbous dorsally and rounded posteriorly ; the integument is

marked with very closely spaced, unbroken longitudinal striae,

acute-angled at the bottom of the grooves and the top of the ridges.

The foot is short, stout and obscurely two- jointed. The toes are

extremely long and unequal; their length, both absolute and rela¬

tive, is highly variable ; the left toe is always shorter than the

right, the difference varying betw^een one third and one fifth of

the length of the right toe. The basal portion of the toes, from

one third to one half of its length, is very nearly straight for the

greater part of its length; this is followed by a tapering section,

which passes into the very nearly cylindrical, extremely slender

posterior portion, which is from one half to two thirds of the entire

length. The right toe is straight and in the axis of the body; the

left toe curves slightly upwards and to the left.

Harring (& Myers — Rotifer Fauna of Wisconsin — II, 539

The dorsal antenna is a small setigerous papilla in the normal

position; the lateral antennae are on the posterior fourth of the

body.

The corona is very slightly oblique and consists of a marginal

wreath of cilia with lateral, auricle-like tufts of longer cilia for

propulsion; the apical plate is unciliated and the buccal field

evenly covered with short, close-set cilia.

The mastax belongs to a type intermediate between the virgate

and the forcipate, the piston playing but a very subordinate role

in its function. The fulcrum is rather short and very broad at

the base, tapering rapidly to the blunt posterior end. The rami

are broadly triangular at the base and the posterior portion lyrate ;

the basal apophysis is abnormally large, somewhat curved and

separated from the rami proper by a deep, rounded sinus. The

inner margins of the rami are equipped with a unique and com¬

plicated dental armature. The teeth are naturally divisible into

three groups: ventral, oral and posterior. The ventral group

consists of 12-14 comblike teeth; their length increases gradually

to a point somewhat beyond mid-length, decreasing from there

towards the oral group. This consists of four extremely large,

stout, slightly curved and acutely pointed, interlocking teeth; the

relative length and development of the individual teeth is some¬

what variable, especially in the case of the second and third

tooth on each side. The posterior group consists of three long,

slender, acutely pointed, interlocking needle-like teeth in each

ramus. Each uncus has three long, slender, unequal teeth, clubbed

at the tips; two rest on the rami at the base of the comb-like teeth

and one on the posterior end, beyond the oral group of teeth.

In the right uncus the ventral tooth is largest and the second

somewhat smaller; in the left uncus the ventral tooth is relatively

small and the second as large as the right ventral. The basal

portion of the manubria is broad and plate-like, the posterior

rod-like and decurved at the end. The piston is relatively small

and weak.

The oesophagus is short. The gastric glands are rather small

and rounded. There is no constriction between the stomach and

instine. The ovary and bladder are normal. The foot glands are

small and pyriform.

The ganglion is large and saccate. The retrocerebral sac is

fairly large and pyriform; it is usually crowded with bacteroids

and opaque to transmitted light. The duct can not be traced be-

540 Wisconsin Academy of Sciences, Arts, and Letters,

yond the dorsal antenna ; snbcerebral glands are not present. The

eyespot is at the posterior end of the ganglion.

Greatest length 680ja; body 210ft; right toe 470ft trophi

Monommata grandis is abundant everywhere in weedy ponds in

the United States; judging from published records it is not com¬

mon in Europe, but it is uncertain whether this is to be attributed

to actual rarity or to reiterated assertions of its specific identity

with M. longiseta.

Subfamily TETRASIPHONINAE.

Genus TETRASIPHON Ehrenberg.

Notommatid rotifers with fusiform, illoricate body, without con¬

striction between head and abdomen; the tail is rudimentary; the

foot is short and two- jointed, with two long, slender toes.

The corona is an oblique, weakly ciliated area without auricles

and used for carrying food to the mouth only, not for propulsion ;

the mouth is a little below the center of the corona.

The antennae are long and tubular, the dorsal antenna double.

The mastax is an aberrant form of the virgate type ; the fulcrum

is short and the rami very large and dome-shaped; the mallei are

imperfectly developed. The epipharynx consists of four pieces

of complicated form, which apparently serve to expand the mouth

opening. The piston is very bulky, but weak; it is attached to

the ventral floor of the mastax.

The retrocerebral organ is well developed ; the subcerebral glands

are longer than the sac. The eyespot is at the posterior end of

the ganglion.

Type of the genus. — Tetrasiphon hydrocora Ehrenberg.

Subfamily LINDIINAE.

Genus LINDIA Dujardin.

Notommatid rotifers with spindle-shaped, elongate, illoricate

body, usually with several indistinct annulations, and without con¬

striction at the base of the foot, which has two very small toes;

the cloaca opens under a small tail, at the base of the foot.

The corona is an elongate, oval area covering the oblique anterior

surface of the head and continuing beyond the mouth on the ven-

Earring & Myers — Rotifer Fauna of Wisconsin — II. 541

tral surface as a slightly projecting chin; the marginal cilia are

relatively short, except on two latero-frontal areas provided with

long and powerful cilia adapted to swimming, in the majority of

species seated on evertile auricles; the apical plate is enclosed by

the marginal ciliation and has occasionally a projecting rostrum;

the mouth is near the center of the corona.

The mastax is adapted to suction or ‘"pumping” by oscillating

as a complete unit on a transverse axis near the posterior end

of the fulcrum; the name “cardate” is proposed for this peculiar

type. The fulcrum is short and narrow, usually tapering towards

the posterior end; the rami are lyrate, resembling the forcipate

type; the ventral branch or cell of the manubria is very large,

equaling or exceeding in length the median or principal cell, with

which it forms a rather acute angle ; the unci are feebly developed,

the teeth being rudimentary and usually united into a thin plate.

Nearly all the species included have an epipharynx of very com¬

plicated form; it may consist of one, two or four separate pieces,

which act as supports for the edges of the mouth, while the oscil¬

lation of the entire mastax produces behind it the vacuum neces¬

sary for the pumping action. Two salivary glands occupy the

posterior portion of the mastax in some species.

The retrocerebral organ consists of a rather small, hemispherical,

ductless sac at the posterior end of the ganglion; it is filled with

red pigment granules and encloses the large, disc-shaped eyespot,

seated on the ganglion.

Type of the genus. — Lindia torulosa Dujardin.

Subfamily BIRGEINAE.

Genus BIRGEA Harring and Myers.

Notommatid rotifers with short, broad, illoricate, anteriorly

truncate body; the head is short and very broad, separated from

the abdomen by a slight constriction; the abdomen is ovate and

ends posteriorly in a short, broad tail ; the foot is long, very slender

and three- jointed; the toes are fairly long and lanceolate.

The corona is frontal with a circumapical band of short cilia,

interrupted dorsally, and two lateral, auricle-like areas with long

cilia adapted to propulsion; the buccal field is evenly ciliated and

the mouth is near the ventral edge.

542 Wisconsin Academy of Sciences y ArtSy and Letters.

The mastax is very aberrant; the trophi, which normally seize

and subdivide the food, are virtually atrophied and their func¬

tions transferred to a pair of hooked ‘ ‘ pseudunci ’ probably a

highly specialized form of the epipharynx; no gastric glands are

present, but the walls of the stomach are produced into a number

of gastric caeca and contain symbiotic zoochlorellae.

No retrocerebral organ is present; the ganglion is small and

the eyespot at its posterior margin.

Type of the genus. — Birgea enantia Harring and Myers.

Harring & Myers — Rotifer Fauna of Wisconsin — II, 543

Explanation of Plates.

All the figures are highly magnified. For actual measurements see text.

The epipharynx is stippled in order to distinguish it more readily from the

trophi proper.

PLATE XVI.

Fig. 1. Proales similis, dorsal view; page 434.

Fig. 2. Proales similis, lateral view.

Fig. 3. Proales similis, trophi, ventral view.

Fig. 4. Proales similis, trophi, lateral view.

Fig. 5. Proales similis, trophi, frontal view.

Fig. 6. Proales reinhardti, dorsal view; page 431.

Fig. 7. Proales reinhardti, lateral view.

Fig. 8. Proales reinhardti, trophi, ventral view.

Fig. 9. Proales reinhardti, trophi, lateral view.

Fig. 10. Proales reinhardti, unci, frontal view.

PLATE XVII.

Fig. 1. Proales wernecTcii, dorsal view; page 426.

Fig. 2. Proales wernecTcii, lateral view.

Fig. 3. Proales wernecTcii, trophi, ventral view.

Fig. 4. Proales wernecTcii, trophi, lateral view.

Fig. 5. Proales wernecTcii, trophi, frontal view.

Fig. 6. Proales gigantea, dorsal view; page 424.

Fig. 7. Proales gigantea, lateral view.

Fig. 8. Proales gigantea, trophi, ventral view.

Fig. 9. Proales gigantea, trophi, lateral view.

Fig. 10. Proales gigantea, trophi, frontal view.

PLATE XVIII.

Fig. 1. Proales daphnicola, dorsal view; page 430.

Fig. 2. Proales daphnicola, lateral view.

Fig. 3. Proales daphnicola, trophi, ventral view.

Fig. 4. Proales daphnicola, trophi, lateral view.

Fig. 5. Proales daphnicola, trophi, frontal view.

PLATE XIX.

Fig. 1. Proales brevipes, dorsal view; page 428.

Fig. 2. Proales brevipes, lateral view.

Fig. 3. Proales doliaris, dorsal view; page 437.

Fig. 4. Proales doliaris, lateral view.

Fig. 5. Proales doliaris, trophi, ventral view.

544

Wisconsin Academy of Sciences, Arts, and Letters.

EXPLANATION OF PLATES-— Continued

Fig. 6. Proales doliaris, trophi, lateral view.

Fig. 7. Proales doliaris, rami, frontal view.

PLATE XX.

Fig. 1. Proales minima, dorsal view; page 435.

Fig. 2. Proales minima, lateral view.

Fig. 3. Proales minima, tropM, ventral view.

Fig. 4. Proales minima, trophi, lateral view.

Fig. 5. Proalinopsis staurus, dorsal view; page 439.

Fig. 6. Proalinopsis staurus, lateral view.

Fig. 7. Proalinopsis staurus, trophi, ventral view.

Fig. 8. Proalinopsis staurus, trophi, lateral view.

Fig. 9. Proalinopsis staurus, trophi, frontal view.

PLATE XXI.

Fig. 1. Notommata epaxia, dorsal view; page 443.

Fig. 2. Notommate epaxia, lateral view.

Fig. 3. Notommata epaxia, trophi, ventral view.

Fig. 4. Notommata epaxia, trophi, lateral view.

Fig. 5. Notommata epaxia, unei, frontal view.

Fig. 6. Notommata codonella, dorsal view; page 444.

Fig. 7. Notommata codonella, lateral view.

Fig. 8. Notommata codonella, trophi, ventral view.

Fig. 9. Notommata codonella, trophi, lateral view.

Fig. 10. Notommata codonella, unci, frontal view.

PLATE XXII.

Fig. 1. Notommata doneta, dorsal view; page 448.

Fig. 2. Notommata doneta, lateral view.

Fig. 3. Notommata doneta, trophi, ventral view.

Fig. 4. Notommata doneta, trophi, lateral view.

Fig. 5. Notommata thopica, dorsal view; page 446.

Fig. 6. Notommata thopica, lateral view.

Fig. 7. Notommata thopica, trophi, ventral view.

Fig. 8. Notommata thopica, trophi, lateral view.

Fig. 9. Notommata thopica, unei, frontal view.

PLATE XXIII.

Earring Myers — Botifer Fauna of Wisconsin — II.

Fig. 7.

Fig. 8.

Fig. 9.

Fig. 10.

Fig. 1.

Fig. 2.

Fig. 3.

Fig. 4.

Fig. 5.

Fig. 6.

Fig. 7.

Fig. 8.

Fig. 9.

Fig. 1.

Fig. 2.

Fig. 3.

Fig. 4.

Fig. 5.

Fig. 6.

Fig. 7.

Fig. 8.

Fig. 1.

Fig. 2.

Fig. 3.

Fig. 4.

Fig. 5.

Fig. 6.

Fig. 7.

Fig. 1.

Fig. 2.

Fig. 3.

Fig. 4.

Fig. 5.

Fig. 6.

Fig. 7.

EXPLANATION OF PLATES— Continued

Taphrocampa annulosa, lateral view.

Taphrocampa annulosa, trophi, ventral view.

Taphrocampa annulosa, trophi, lateral view.

Taphrocampa annulosa, unci, frontal view.

PLATE XXIV.

Taphrocampa clavigera, dorsal view; page 455.

Taphrocampa clavigera, lateral view.

Taphrocampa clavigera, trophi, ventral view.

Taphrocampa clavigera, trophi, lateral view.

Taphrocampa selenura, dorsal view; page 454.

Taphrocampa selenura, lateral view.

Taphrocampa selenura, trophi, ventral view.

Taphrocampa selenura, trophi, lateral view.

Taphrocampa selenura, unci, frontal view.

PLATE XXV.

Fleurotrocha petromyzon, dorsal view; page 459.

Pleurotrocha petromyzon, lateral view.

Fleurotrocha petromyzon, trophi, ventral view.

Fleurotrocha petromyzon, trophi, lateral view.

Fleurotrocha rohusta, dorsal view; page 461.

Fleurotrocha rohusta, lateral view.

Fleurotrocha rohusta, trophi, ventral view.

Fleurotrocha rohusta, trophi, lateral view.

PLATE XXVI.

Cephalodella mineri, lateral view; page 471.

Cephalodella elongata, lateral view; page 471.

Cephalodella innesi, lateral view; page 470.

Cephalodella innesi, trophi, ventral view.

Cephalodella innesi, trophi, lateral view. ,

Cephalodella paxilla, lateral view; page 468.

Cephalodella marina, lateral view; page 469.

PLATE XXVII.

Cephalodella gracilis, lateral view; page 473.

Cephalodella angusta, lateral view; page 467.

Cephalodella catellina, lateral view; page 465.

Cephalodella catellina, trophi, ventral view.

Cephalodella catellina, trophi, lateral view.

Cephalodella sterea, lateral view; page 474.

Cephalodella epitedia, lateral view; page 468.

545

546

Wisconsin Academy of Sciences^ ArtSy and Letters,

Fig. 1.

Fig. 2.

Fig. 3.

Fig. 4.

Fig. 5.

Fig. 6.

Fig. 1.

Fig. 2.

Fig. 3.

Fig. 4.

Fig. 5.

Fig. 6.

Fig. 7.

Fig. 1.

Fig. 2.

Fig. 3.

Fig. 4.

Fig. 5.

Fig. 6.

Fig. 7.

Fig. 1.

Fig. 2.

Fig. 3.

Fig. 4.

Fig. 5.

Fig. 6.

Fig. 7.

Fig. 8.

Fig. 1.

Fig. 2.

Fig. 3.

Fig. 4.

Fig. 5.

Fig. 6.

Fig. 7.

EXPLANATION OP PLATES — Continued

PLATE XXVIII.

CepJialodella hoodii, lateral view; page 482.

Cephalodella exigua, lateral view; page 481.

Cephalodella plicata, lateral view; page 483.

CepJialodella plicata, cross section.

CepJialodella ventripes, lateral view; page 484.

CepJialodella auriculata, lateral view; page 479.

PLATE XXIX.

CepJialodella nana, lateral view; page 491.

CepJialodella cuneata, lateral view; page 505.

CepJialodella pJiysalis, lateral view; page 484.

Cephalodella physalis, trophi, ventral view.

Cephalodella physalis, trophi, lateral view.

Cephalodella xenica, lateral view; page 492.

Cephalodella strigosa, lateral view; page 485.

PLATE XXX.

Cephalodella compressa, lateral view; page 487.

Cephalodella tantilla, lateral view; page 486.

Cephalodella piulca, lateral view; page 488.

CepJialodella gihha, lateral view; page 472.

Cephalodella gibha, trophi, ventral view.

Cephalodella gibba, trophi, lateral view.

Cephalodella dorseyi, lateral view; page 487.

PLATE XXXI.

Cephalodella galbina, lateral view; page 490.

Cephalodella belone, lateral view; page 490.

Cephalodella globata, lateral view; page 475.

CepJialodella panarista, lateral view; page 478.

CepJialodella panarista, trophi, ventral view.

Cephalodella panarista, trophi, lateral view.

Cephalodella panarista, toe with spine.

Cephalodella elegans, lateral view; page 489.

PLATE XXXII.

Cephalodella nelitis, lateral view; page 493.

Cephalodella melia, lateral view; page 493.

Cephalodella pheloma, lateral view; page 496.

Cephalodella hyalina, lateral view; page 505.

Cephalodella megalocephala, lateral view; page 494.

Cephalodella megalocephala, trophi, ventral view.

Cephalodella megalocephala, trophi, lateral view.

Earring (& Myers — Rotifer Fauna of Wisconsin — II,

Pig.

Fig.

EXPLANATION OF PLATES— Continued

PLATE XXXIII.

1. CepJialodella apocolea, lateral view; page 509.

2. Cephalodella apocolea, toes, dorsal view.

3. Cephalodella tenuior, lateral view; page 497.

4. Cephalodella dixon-nuttalli, lateral view; page 498.

5. Cephalodella dixon-nuttalli, toes, dorsal view.

6. Cephalodella retusa, lateral view; page 498.

7. Cephalodella forficata, lateral view; page 499.

PLATE XXXIV.

1. Cephalodella forficula, lateral view; page 476.

2. Cephalodella forficula, tropM, ventral view.

3. Cephalodella forficula, trophi, lateral view.

4. Cephalodella licinia, lateral view; page 503.

5. Cephalodella collactea^ lateral view; page 501.

6. Cephalodella speciosa, lateral view; page 504.

7. Cephalodella papillosa, lateral view; page 506.

PLATE XXXV.

1. Cephalodella strepta, lateral view; page 509.

2. Cephalodella intuta, lateral view; page 500.

3. Cephalodella intuta, trophi, ventral view.

4. Cephalodella intuta, trophi, lateral view.

5. Cephalodella intuta, toes, dorsal view.

6. Cephalodella vacuna, lateral view; page 503.

7. Cephalodella tenuiseta, lateral view; page 508.

8. Cephalodella eva, lateral view; page 507.

PLATE XXXVI.

1. Cephalodella inquilina, lateral view; page 502.

2. Cephalodella mucronata, lateral view; page 510.

3. Cephalodella mucronata, trophi, ventral view.

4. Cephalodella mucronata, trophi, lateral view.

5. Cephalodella lipara, lateral view; page 512.

6. Cephalodella parasitica, lateral view; page 511.

7. Cephalodella eupoda, lateral view; page 512.

PLATE XXXVII.

1. Rousseletia corniculata, dorsal view; page 514.

2. Rousseletia corniculata, lateral view.

3. Rousseletia corniculata, trophi, ventral view.

4. Rousseletia corniculata, trophi, lateral view.

5. Tylotrocha monopus, dorsal view; page 516.

548 Wisconsin Academy of Sciences, Arts, and Letters,

EXPLANATION OP PLATES— Continued

Pig. 6. Tylotrocha monopus, lateral view.

Fig. 7. Tylotrocha monopus, trophi, ventral view.

Fig. 8. Tylotrocha monopus, trophi, lateral view.

PLATE XXXVIII.

Fig. 1. Besticula nyssa, dorsal view; page 521.

Fig. 2. Eesticula nyssa, lateral view.

Fig. 3. Eesticula nyssa, trophi, ventral view.

Fig. 4. Eesticula nyssa, trophi, lateral view.

Fig. 5. Eesticula anceps, dorsal view; page 519.

Fig. 6. Eesticula anceps, lateral view.

Fig. 7. Eesticula anceps, trophi, ventral view.

Fig. 8. Eesticula anceps, trophi, lateral view.

PLATE XXXIX.

Pig. 1. Eosphora thoa, dorsal view; page 523.

Fig. 2. Eosphora thoa, lateral view.

Fig. 3. Eosphora thoa, trophi, ventral view.

Fig. 4. Eosphora thoa, trophi, lateral view.

Fig. 5. Eosphora thoa, trophi, frontal view.

PLATE XL.

Fig. 1. Enteroplea lacustris, dorsal view; page 526.

Fig. 2. Enteroplea lacustris, lateral view.

Fig. 3. Enteroplea lacustris, trophi, ventral view.

Fig. 4. Enteroplea lacustris, trophi, lateral view.

Pig. 5. Enteroplea lacustris, incus, oblique frontal view.

PLATE XLI.

Earring & Myers — Rotifer Fauna of Wisconsin — II, 549

EXPLANATION OP PLATES— Continued

Sphyrias lofuana, trophi, ventral view.

Sphyrias lofuana, trophi, lateral view.

Sphyrias lofuana, trophi, oblique frontal view.

PLATE XLIII.

Monommata longiseta, dorsal view; page 535.

Monommata longiseta, lateral view.

Monommata longiseta, trophi, ventral view.

Monommata longiseta, trophi, lateral view.

Monommata longiseta, trophi, oblique frontal view. '

Monommata grandis, dorsal view; page 538.

Monommata grandis, lateral view.

Monommata grandis, trophi, ventral view.

Monommata grandis, trophi, lateral view.

Monommata grandis, trophi, oblique frontal view.

I'

TRANS. WIS. ACAD., VOL. XXI.

PLATE XVI

HARRING AND MYERS. -NOTOMMATID ROTIFERS

TRANS. WIS. ACAD., VOL. XXL

PLATE XVIl

HARPING AND MYERS.— NOTOMMATID ROTIFERS

Op

TRANS. WIS. ACAD., VOL. XXI.

PLATE XVIII

MARRING AND MYERS.— NOTOMMATID ROTIFERS

■mAMS. wrs. ACAD., VOL. XX!.

PLATE XIX

HARKING AND MYERS.— NOTOMMATID ROTIFERS

TWANS. WI8. ACAD., VOL. XXI.

PLATE XX

■1

TRANS. WIS. ACAD.. VOL. XXI.

PLATE XXI

HARKING AND MYERS— NOTOM M ATI D ROTIFERS

TRANS. WIS. ACAD., VOL, XXi.

HARRING AND MYERS.— NOTOMMATIO ISOTIFERS

TRANS. W!S. ACAD.. VOL. XXI.

PLATE XXm

HARRSNG and MYERS.— NOTOMM ATI O ROTIFERS

PLATE XXIV

MARRING AND MYERS —NOTOMMATID ROTIFERS

TRANS. WIS. ACAD.. VOL. XXI.

HARKING AND hdVERS— NOTOWIMATlD ROTIPBRS

TRANS. WIS. ACAD., VOL. XXI.

PLATE XXVI

MARRING AND MYERS.— NOTOMMATID ROTIFERS

WARRING AND MYERS.— NOTOMMATiD ROTIFERS

'■'r'

TRANS. WiS. ACAD., VOL. XXI.

PLATE XXVIII

HARKING AND MYERS.— NOTOMMATID ROTIFERS

r

MARRING AND MYERS- NOTOMMATID ROTIFERS

TRANS. WIS. ACAD., VOL. XXI.

PLATE XXX

MARRING AND MYERS.- NOTOMMATID ROTIFERS

TRANS. WIS. ACAD., VOL. XXI.

PLAtE XXXI

HARRIN6 AND MYERS.— NOTOMMATID ROTlFKfl®

TRANS. WIS. ACAD.. VOL. XXI.

PLATE XXXII

MARRING AND MYERS.— NOTOMMATID ROTIFEfJS

TRANS. WIS. ACAD., VOL. XXI

PLATE XXXIII

HARPING AND MYERS.- NOTOMMATID ROTIFERS

TRANS. WIS. ACAD., VOL. XXI.

PLATE XXXIV

HARRINO AND MYKRS.-NOTOMMATID ROTIFERS

TRANS. WIS. ACAD., VOL. XXI.

PLATE XXXV

MARRING AND MYERS.— NOTOMMATID ROTIFERS

TRANS. WIS. ACAD.. VOL. XX!.

PLATE XXXVI

HARRING AND MYERS.— NOTOMMATJD ROTIFERS

HARRING AND MYERS.— NOTOMMATID ROTIFERS

■TOANS. WFS. ACAD.. VOL. XX!

PLATE XXXVIII

HARRSNG AND MYERS.— NOTOMM ATI D ROTIFERS

TRANS. WIS. ACAD., VOL. KXl

PLATE XXXIX

HARRtNS AND MYERS.- NOTOMMATID ROTIFERS

TRANS. WiS. ACAD.. VOL. XXI.

PLATE XL

HARKINS AND MYER«.--NOTOIMMATiD WOTI^RS

TRANS. WIS. ACAD., VOL. XXI.

TRANS. WIS. ACAD.. VOL. XXI.

PLATE XLII

HARRiNG AND MYERS.— NOTOMMATID ROTIFERS

TRANS. WIS. ACAD., VOL. XX!.

HARRiMO AND MYERS.— NOTOMMATID ROTIFERS

PROCEEDINGS OF THE ACADEMY

1921 TO 1923

FIFTY-FIRST ANNUAL MEETING, 1921

The fifty -first meeting of the Wisconsin Academy of Sciences,

Arts, and Letters, in joint session with the Wisconsin Archeological

Society, was held at Madison, in Room 301, Biology Building,

University of Wisconsin, on Friday and Saturday, April 15 and

16, 1921.

Under the direction of the president the following program was

presented :

First Session, Friday, April 15, 3 :00 P. M.

General Business.

Presentation of Papers.

1. Pood Value of some Aquatic Organisms found in Lake Mendota.

Chancey Juday.

2. Notes on the Herpetology of Wisconsin. George Wagner.

3. The Distribution of the Pishes in Six Wisconsin Lakes. A. S. Pearse,

4. The Eadiation Hypothesis of Chemical Action. Parrington Daniels.

Illustrated.

5. The Polarization Method of Measuring the Gloss of Paper. L. E. Inger-

SOLL. Illustrated.

6. Notes on the Chemical Composition of some of the Larger Aquatic Vege¬

tation of Lake Mendota. Henry A. Schuette. By title.

7. A Preliminary List of the Water Mites of Wisconsin. Euth Marshall.

By title.

Second Session, Saturday, April 16, 9 :30 A. M.

8. Bisexual Organs in Bryum medium. George S. Bryan. Illustrated.

9. Some Cytological Studies on Beloniella BehniL E. M. Gilbert.

10. Aquatic Pungi of the Madison Lakes. Progress Eeport. E. M. Gilbert.

11. The Anther ozoid of Biccardia pinguis. W. N. Steil.

12. Notes on Parasitic Pungi in Wisconsin, IX. J. J. Davis.

13. Stone Celts from Southern Colorado. C. E. Brown.

Effigy Mound Photographs. George E. Pox.

14. Jacob Wimpfelings Germania, Strassburg 1501 and Thomas Murners

Germania Nova, 1502. A Controversy about the Nationality of Alsace

(at the beginning of the Sixteenth Century). Ernst Voss.

552 Wisconsin Academy of Sciences, Arts, and Letters.

Third Session, Saturday, April 16, 2 :30 P. M.

11. The American Housing Situation. Leonard S. Smith.

12. The Declining Use of Lake Michigan as a Waterway. E. H. Whitbeck.

13. Some Observations on the Volcano of Kilauea. G. B. Culver. Illus¬

trated.

14. Eoad Oil and Its Use. A. F. Galman. By title.

15. Carl Winkler and his Meteorological Observations. Edward Kremers.

By title.

The Secretary presented the following applications for mem¬

bership. On motion he was instructed to cast the ballot in their

favor :

Lloyd Slote Dancey, Waukesha

Howard Greene, Milwaukee

E. E. Hubert, Madison

F. F. Lewis, Janesville

Stanley Walker Eockwood, Waukesha

James Elcana Eogers, Waukesha

Harry Linn Starr, Waukesha

Clarence L. Turner, Beloit

He then presented the secretary’s report for the year, which was

approved :

Report of the Secretary for the Year 1920

Honorary Members . 7

Life Members . 17

Corresponding Members . 34

Active Members . 247

Total . 305

Changes since last report:

Active members reported for 1919 . 244

Members reinstated . 3

New Memberships enrolled in 1920 . 16

263

Deaths of Active Members . 5

Eesignations . 1

Dropped for non-payment of dues . 7

Active Members made Life Members . 3 16

Present Active Membership . 247

Mr. George Merrick was enrolled as an Honorary Member in

1920. Seven new Life Members were enrolled in 1920; Thomas

E. Brittingham, Madison; Prank P. Hixon, La Crosse; A. J. Hor-

Proceedings of the Academy.

553

lick, Racine; C. K. Leith, Madison; Frank A. Logan, Chicago;

Mrs. Charles M, Norris, Milwaukee ; and M. S. Slaughter, Madison.

During 1920 arrangements were completed whereby the Academy

and the American Association for the Advancement of Science

were affiliated.

There are 66 new members who have paid their dues to be en¬

rolled after their applications have been acted upon by this meet¬

ing. Of this number, 58 will become members of both societies.

I regret to report the loss of five active members by death: E. T. Harper,

Geneseo, Ill.; Herman W. Kunz, Milwaukee; Publius V. Lawson, Menasha;

W. A. P. Morris, Madison; and Dr. C. H. Vilas, Madison.

Arthur Beatty,

Secretary.

The treasurer’s report for the year was next presented, adopted,

and audited, as follows:

Report of the Treasurer for the Year 1920

RECEIPTS

Received from Dues and Initiations . $1,034.50

Received from sale of Transactions . 14.58

Received from Interest on Certificates of Deposit . 16.00

Received from Interest on Liberty Bonds . 31.25

Received from Interest on City of Madison Bonds . 118.00

Received from 5 City of Madison Bonds matured April 1, 1921 . 500.00

Total . $1,714.33

Balance on hand April 23, 1920 . 152.95

Total . $1,867.28

DISBURSEMENTS

Secretary- Treasurer ’s Allowance . $200.00

6 City of Madison Bonds purchased April 1, 1921 . 600.00

Warrant sent A. A. A. S. in adjustment of dues . 540.00

1,340.00

. $ 527.28

Arthur Beatty,

Treasurer.

Balance on hand April 16, 1921

554 Wisconsin Academy of Sciences^ Arts, and Letters,

We have examined the securities owned by the Academy and find them in

accord with the Treasurer's record.

We approve the statement of Eeceipts and Disbursements of the Treasurer

as shown by his record.

E. H. Whitbeck,

E. G. Smith,

Auditors.

The next order of business was the report of the nominating

committee :

Your committee on nominations reports the following nomina¬

tions for officers and committees for the ensuing term:

President, Melvin A. Brannon, Beloit.

Vice President, Sciences, S. A. Barrett, Milwaukee.

Vice President, Arts, Grant Showerman, Madison.

Vice President, Letters, Karl Young, Madison.

Secretary, C. Juday, Madison.

Treasurer, C. Juday, Madison.

Curator, C. E. Brown, Madison.

Librarian, W. M. Smith, Madison.

COMMITTEE ON PUBLICATIONS

The Secretary, ex-officio.

The President, ex-officio.

W. E. Tottingham, Madison.

COMMITTEE ON LIBRAEY

The Librarian, ex-officio.

George Wagner, Madison.

W. Harley Barber, Eipon.

A. E. Whitford, Milton.

A. A. Trever, Appleton.

COMMITTEE ON MEMBERSHIP

The Secretary, ex-officio.

P. W. Boutwell, Beloit.

V. E. McCaskill, Superior.

H. H. Smith, Milwaukee.

G. S. Bryan, Madison.

We also recommend that the present Secretary and C. E. Allen of the Com¬

mittee on Publication be continued for a period of six months.

Eespectfully submitted,

Charles E. Brown,

E. M. Gilbert,

Wm. S. Marshall.

Committee on Nominations.

Proceedings of the Academy.

555

The report of the Nominating Committee was adopted and the

Secretary was instructed to cast the ballot for these officers.

It was moved by W. S. Marshall that the Nominating Committee

be continued and be directed to draft an amendment to Article 6

of the constitution of the Academy, relating to the appointment

of committees by the President. Carried.

It was moved by Edward A. Birge that those who contributed

$100 to the Fiftieth Anniversary Medal be placed on the list of

the Life Members of the Academy, in accordance with the earlier

practice of the Academy. Carried.

This motion applies to the following persons, who were duly

elected Life Members:

Hon. Thomas E. Brittingham, Madison

Hon. F. P. Hixon, La Crosse

Hon. A. J. Horlick, Eacine

Dr. Charles K. Leith, Madison

Hon. Prank A. Logan, Chicago

Mrs. Charles M. Norris, Milwaukee

Dr. M. S. Slaughter, Madison

Dr. E. A. Birge, Madison, and Dr. C. S. Slichter, Madison, who

donated $100 each to the Medal Fund, were already Life Members

of the Academy.

Fourth Session, Saturday, 6 :30 P. M.

The annual dinner was held at the University Club, at which

54 persons were present. The retiring President, Edward A.

Birge, delivered an address on ‘‘A Lake as a Going Concern.”

The President had specimens of the Fiftieth Anniversary Medal,

which were inspected and highly approved by the guests.

The 1921 meeting was then adjourned.

556 Wisconsin Academy of Sciences , Arts, and Letters.

FIFTY-SECOND ANNUAL MEETING, 1922

The fifty-second annual meeting of the Wisconsin Academy was

held in the Trustee’s Room of the Public Museum, Milwaukee,

Wisconsin, on Thursday and Friday, April 13 and 14, 1922.

First Session, Thursday, April 13, 2 :00 P. M.

Presentation of Papers.

1. Some Aspects of British Colonial Policy during the Nineteenth Century.

Paul Knaplund.

2. An Aspect of Samuel Johnson’s Criticism of Shakespear. Karl Young.

3. An Experimental Study of the Relativity of Spacial Perception. Mar¬

garet Wooster.

4. Home Economics and the College of Classical Traditions. Florence P.

Robinson.

5. A Blackfoot Sweat Lodge. S. A. Barrett. Illustrated.

6. A New Method of Museum Technique. R. E. Tyrrell. Illustrated with

specimens.

7. Notes on the Distribution and Occurrences of Insects in Wisconsin. T.

E. B. Pope. Illustrated with specimens.

8. Development of the Eye of the Confused Flour Beetle, Trilobium con-

fusum. W. S. Marshall.

9. On the Anatomy of Atropos divinatoria Mull. Ruth W. Chase.

10. Typhoid Agglutinins in Rabbits. Elizabeth A. Smith.

11. Some Phenomena in a Dying Lake. M. A. Brannon.

12. The Sangamon River — A Study in Stream Pollution. Minna E. Jewell.

13. Crayfishes in the Eastern United States, with Special Reference to Ohio.

C. L. Turner.

Second Session, Friday, April 14, 9:00 A. M.

Presentation of Papers.

14. The Piskum or Plains Indian Buffalo Drive. S. A. Barrett. Illustrated.

15. Menominee Customs Concerning Children. Alanson Skinner.

16. A Preliminary Report on the Ethno-botany of the Menominee Indians.

Huron H. Smith. Illustrated.

17. Notes on Parasitic Fungi in Wisconsin, X. J. J. Davis.

18. Cytological Studies on the Lower Basidiomycetes, II. Aurieularia. E.

M. Gilbert.

19. A Census of the Water Molds of a Portion of Lake Monona. E. M. Gil¬

bert.

20. Inheritance in a Simple Plant. C. E. Allen. Illustrated.

21. Progress Report on the Flora of Wisconsin. Huron H. Smith.

22. Notes on the Occurrences of Certain Wisconsin Fishes. T. E. B. Pope.

23. The Parasites of Lake Pishes. A. S. Pearse.

Proceedings of the Academy.

557

24. Observations Concerning the Eespiration of Turtles. Frances E. Die-

bold.

25. A Quantitative Study of the Bacteria of Lake Mendota. E. B. Fred and

Frank C. Wilson.

26. The Hydrogen Ion Concentration in Wisconsin Lake Waters. Frank C.

Wilson, E. B. Fred, and C. Juday.

Third Session, Friday, April 14, 2:00 P. M.

Presentation of Papers.

27. The Significance of an Economic Fisheries Exhibit and Its Eelation to

Conservation. T. E. B. Pope.

28. The Analytical Chemistry of Selenium and Tellurium. V. Lenher.

29. The Summer Temperature of the Bottom Water in Some Deep Lakes of

the Western United States. Geo. I. Kemmerer.

30. Unusual Veinings in the Eichardton Meteorite. E. N. Buckstapp.

Illustrated with specimens.

31. The Progress of Topographic Mappings in Wisconsin. W. O. Hotchkiss.

32. Eoad Material Surveys in Wisconsin. E. F. Bean.

33. The Lime Industry in Wisconsin. E. Steidtmann.

34. The Mining of Sulphur at Freeport, Texas. Ira Edwards. Illustrated.

35. The Upper Mississippi River as a Commercial Waterway. E. H. Whit-

beck.

36. The Cytology of Venturia. C. N. Frey.

The annual dinner was held on Thursday evening at the Hotel

Wisconsin, with 86 persons in attendance.

Following the dinner brief addresses were made in honor of

three of the early members of the Academy who are commemorated

on the Semi-centennial Medallion. President E. A. Birge spoke

on Dr. George W. Peckham, Mr. William W. Wight on Dr. I. A.

Lapham, and Dr. John J. Davis on Dr. P. K. Hoy.

The Secretary presented the following applications for member¬

ship. On motion he was instructed to cast the ballot in their favor :

Irving M. Addleman, Wausau

B. P. Churchill, Milwaukee

H. H. Conwell, Beloit

Percy M. Dawson, Madison

H. D. Densmore, Beloit

Edward Evans, La Crosse

Geo. J. Fiebiger, Waterloo

Arthur C. Foster, Ealeigh, N. C.

Edwin B. Fred, Madison

Chas. N. Frey, New York, N. Y.

W. A. Hamilton, Beloit

E. G. Hastings, Madison

558

Wisconsin Academy of Sciences, Arts, and Letters,

Thomas M. Jasper, Urbana, Ill.

W. A. Kenyon, Madison

Clifford S. Leonard, Madison

Fred. L. Musbach, Marshfield

Lowell E. Noland, Madison

Henry V. Ogden, Milwaukee

C. E. Patzer, Milwaukee

Geo. E. Potter, Durham, N. H.

John Walker Powell, Milwaukee

Jessie P. Eose, Madison

Arthur Simon, Milwaukee

Kurt Stock, Pish Creek

Noel P. Thompson, Madison

Pred. T. Ullrich, Platteville

David E. W. Wenstrand, Milwaukee

J. W. White, Platteville

Clyde M. Woodworth, Urbana, Ill.

John A. Jeske, Milwaukee

Chas. E. Porteus, Milwaukee

^ Stephen J. Majerowski, Milwaukee

Edward E. Tyrrell, Milwaukee

Tenus Tuttrup, Milwaukee

S. Cheifetz, Milwaukee

George Peter, Milwaukee

E. S. Haynes, Beloit

G. D. Shallenberger, Beloit

Clarence L. Clark, Beloit

Kenneth B. Barnes, Beloit

Florence P. Eobinson, Beloit

Wm. J. Trautman, Beloit

E. N. Buckstaff, Oshkosh

E. A. Goessl, Milwaukee

The following individuals were elected to Life Membership:

Arthur Beatty, Madison

Mrs. Elizabeth G. Peckham, Milwaukee

Report of Secretary, November 1, 1921 to April 10, 1922

Honorary Members . 6

Life Members . 17

Corresponding Members . 33

Active Members — Old . 275

Active Members — New . 44

Members reinstated . 3

Total . 378

Eesigned . 4

Proceedings of the Academy.

559

The Illinois State Academy of Science extended a cordial in¬

vitation to the members of the Wisconsin Academy to attend its

meeting at Eockford, Illinois, April 27 to 29, 1922. It was voted

that greetings and good wishes for a successful meeting be ex¬

tended to the Illinois State Academy of Science.

Being an affiliated society, the Wisconsin Academy is entitled

to a representative on the Council of the American Association for

the Advancement of Science. It was voted that the Secretary

represent the Wisconsin Academy on the Council of the American

Association and that he be authorized to appoint some member to

represent the Academy when he does not attend the meetings of

the American Association.

It was voted that the Academy express its thanks to the Mil¬

waukee Public Museum for its interest in the meeting and for

the use of a room for the sessions.

Chancey Juday,

Secretary.

Report of Treasurer for 1922

RECEIPTS

Eeceived from previous Treasurer, November 15, 1921 . ...$ 367.58

Eeceived from Dues and Initiations . . . 944.10

Eeceived from sale of Transactions. . . . . 96.00

Eeceived from Interest on Bonds . . * . . . 50.62

$1,458.30

DISBURSEMENTS

Patterns for Medallion . . . $ 12.10

Postage . . . . . . . . . 58.00

Printing . . . . . . . . 13.36

Safety Deposit Box Rent. . . . . 4.00

Warrant sent A. A. A. S. in Adjustment of Dues . 671.10

$ 758.56

Balance April 10, 1922 . . . . 699.74

$1,458.30

Audited and found correct.

C. E. Allen,

Ira Edwards,

Auditors.

560 Wisconsin Academy of Sciences, Arts, and Letters.

Securities owned by the Academy April 10, 1922:

Government Bonds . . . $ 700.00

City of Madison Bonds . 2,600.00

Certificate of Deposit . 76.38

$3,376.38

Chancey Juday,

Treasurer.

FIFTY-THIRD ANNUAL MEETING, 1923

The fifty-third annual meeting of the Wisconsin Academy was

held at Beloit College, Beloit, Wisconsin, on April 6 and 7, 1923,

in joint session with the Wisconsin Archeological Society.

The following program was presented:

First Session, Friday, April 6, 10 :00 A. M.

Presentation of Papers.

1. Augustine of Hippo qua Patriot. E. K. Eichardson.

2. Chartiludium Institute Summarrie Doctore Thoma Murner Memorante et

Ludente. Strassburg, 1518. Ernst Voss.

3. The Sentimental Eeturn to Nature in the Eighteenth Century. William

E. Alderman.

4. Botanical Collecting in Southwestern Wisconsin in 1922. Huron H.

Smith. Illustrated.

5 Eacial Characters in Sphaerocarpus and their Inheritance. C. E. Allen.

Illustrated.

6. Physiological Stability in Maize. W. E. Tottingham.

7. Notes on Parasitic Fungi in Wisconsin. J. J. Davis.

8. On the Nature of Disease Eesistance in Plants. J. C. Walker.

9. The Nuclear Phenomena in Some of the Short-cycled Eusts. E. M. Gil¬

bert.

10. The Corrected Names of Certain Milk Bacteria. Euth W. Chase and

W. D. Frost.

11. The Characteristics of Certain Fecal Bacteria as shown by the Little

Plate Method. Ola E. Johnston and W. D. Frost.

Second Session, Friday, April 6, 2 :00 P. M.

Presentation of Papers.

12. Beloit Mound Groups. George L. Collie.

13. Winnebago Chieftains and Villages of the Lower Eock Eiver Eegion.

N. W. JiPSON.

14. Eemoval of the Eock River Winnebago in 1833. Louise P. Kellogg.

15. The Applications of Amerind Decorative Art. Mrs. W. E. Tylor.

16. Prehistoric Archeology in Prance. A. W. Pond. Illustrated.

Proceedings of the Academy.

561

17. Stage Coaches and Taverns of the Baraboo Eegion. H. E. Cole.

18. Wood County Potawatomi. A. Gerend.

19. The Manufacture of Stone Axes and Celts. H. L. Skavlem.

20. Indian Cave, Eichland County. C. E. Brown.

21. Wisconsin Caves. W. C. English.

22. The Glaciation of Northern Illinois. M. M. Leighton, University of

Illinois.

23. Some Eecent Discoveries of Wisconsin Graptolites. Eufus M. Bagg.

24. The Diplomatic Crisis of our Civil War. E. B. Way.

Third Session, Saturday, April 7, 9 :00 A. M.

Presentation of Papers.

25. Floundering in Modernity. George C. Clancy.

26. Scientific Pupil-classification. C. G. P. Franzen.

27. Milton as a Writer on Education. Oliver M. Ainsworth.

28. Symposium on Eugenics.

a. General Statement of Problems. M. F. Guyer.

b. Pauperism and Crime in Wisconsin. J. L. Gillin.

c. The Inheritance of Mental Traits. V. A. C. Henmon.

Fourth Session, Saturday, April 7, 1 :30 P. M.

29. Further Notes on Wisconsin Eep tiles. George Wagner.

30. A Case of Arrested Development in a Frog’s Heart. George Wagner.

31. N^otes on the Biology of the Book Louse. Euth W. Chase.

32. The Amount of Food Eaten by Four Wisconsin Fishes. A. S. Pearse,

Illustrated.

33. The Chemical Composition of Trout and Perch. A. S. Pearse.

34. A Mutation in the Moth Fly (Psychoda) and its Method of Inheritance.

C. L. Turner. Illustrated with specimens.

35. A new Arrhenurus from Washington State. Euth Marshall. By title.

36. Water Mites from Alaska and The Canadian Northwest, with an Account

of the Collecting Grounds. Euth Marshall. Illustrated.

3^7. Landmarks Work in Winnebago County. A. P. Kannenberg.

The annual dinner was held on Friday evening, April 6, at

Beloit College. Following the dinner Professor W. S. Bayley,

President of the Illinois State Academy of Science, gave an ad¬

dress on The Function of a State Academy of Science.

The Secretary presented the following applications for member¬

ship. On motion he was instructed to cast the ballot in their favor :

R. A. Brink, Madison

Wm. A. Clark, Stevens Point

Thos. A. Rogers, Stevens Point

G. Steiner, Washington, D. C.

562 Wisconsin Academy of Sciences, Arts, and Letters.

Report of Secretary, April 10, 1922 to April 2, 1923

Honorary Members . 7

Life Members . 17

Corresponding Members . 32

Active Members . 328

384

Eesigned . 3

Deaths . 4

The following deaths have been reported to the Secretary: Professor Henry

Prentiss Armsby, Corresponding Member, State College, Pa.; Professor A. S.

Flint, Life Member, Madison; Dr. Paul H. Dernehl, Active Member, Mil¬

waukee; Professor Fritz Wilhelm Well, Active Member, Berkeley, California.

The Winnebago County Archeological and Historical Society

extended a cordial invitation to the members of the Wisconsin

Academy and the Wisconsin Archeological Society to attend its

field meeting at Oshkosh on June 8 and 9, 1923.

The following resolution was presented:

Whereas there are in the State of Wisconsin a considerable number of in¬

teresting caves, and

Whereas some of these, of notable size and beauty, containing stalactites

and other interesting geological formations as well as Indian pictographs and

petrographs of scenic and historical interest to the general public and to tour¬

ists, are being mutilated and otherwise defaced by thoughtless persons,

Be it resolved that the Wisconsin Academy of Sciences, Arts and Letters

and the Wisconsin Archeological Society, in joint meeting assembled at

Beloit College, Beloit, Wisconsin, on this sixth day of April, 1923, greatly

deplore the neglect into which these scenic monuments have fallen and the

continuance of such destructive practices and depredations and urge upon the

owners of these caves that they in every way protect them against further

mutilations and misuse, and upon the State of Wisconsin itself to take such

measures as shall make such vandalism an offense against its laws.

Charles E. Brown,

W. O. Hotchkiss,

A. S. Pearse,

Committee on Besolutions.

The resolution was unanimously adopted.

It was moved that the thanks of the Wisconsin Academy of

Sciences, Arts, and Letters and of the Wisconsin Archeological

Society be extended to Beloit College for its great interest in the

meetings and for the hospitality shown to the members of the two

societies. The motion was unanimously adopted.

Chancey Juday,

Secretary.

Proceedings of the Academy,

563

Report of Treasurer, April 10, 1922 to April 2, 1923

The publication of Volume XX of the Transactions left a deficit

of nearly $700 so that it was necessary to cash the certificate of

deposit amounting to $76.38 and to sell $600 in government bonds

in order to pay the printing deficit. This action was approved

by the Council of the Academy.

RECEIPTS

Balance in State Treasury, July 1, 1922 . $ 28.48

Annual appropriation for fiscal year 1922-1923 . 1,500.00

Sale of Transactions . 29.95

Annual dues of Active Members . 912.00

$2,470.43

DISBURSEMENTS

Dues of members of A. A. A. S . $ 668.00

Freight on Transactions to Washington, D. C . . 13.09

Labor in packing and mailing Transactions . 13.60

Postage on letters and Transactions . 29.48

Allowance of Secretary for 1922 . 200.00

$ 924.17

Balance in State Treasury, April 2, 1923 . 1,546.26

$2,470.43

Owing to the high cost of printing and to the limited funds, it

has been thought best to omit the membership list in this volume

of the Transactions. It will be printed in the next volume.

Chancey Juday,

Treasurer,

Audited and found correct.

C. E. Allen,

Ira Edwards,

Auditors,

8