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TRANSACTIONS

OF THE

WISCONSIN ACADEMY

OF

SCIENCES, ARTS AND LETTERS

VOL. XXVII

TRANSACTIONS

OF THE

WISCONSIN ACADEMY

OF

SCIENCES, ARTS AND LETTERS

VOL. XXVII

NATURAE SPECIES RATIOOUE

MADISON, WISCONSIN

OFFICERS OF THE WISCONSIN ACADEMY OF

SCIENCES, ARTS AND LETTERS

President

Charles E. Allen, University of Wisconsin

Vice-Presidents

In Science : Rufus M. Bagg, Lawrence College

In the Arts: Otto L. Kowalke, University of Wisconsin

In Letters: William E. Alderman, Beloit College

Secretary-Treasurer

Lowell E. Noland, University of Wisconsin

Librarian

Walter M. Smith, University of Wisconsin

Curator

Charles E. Brown, State Historical Museum

Council

The president, ex officio

The vice-presidents, ex officio

The secretary-treasurer, ex officio

The librarian, ex officio

E. A. Birge, past president

Charles S. Slichter, past president

John J. Davis, past president

Louis Kahlenberg, past president

L. J. Cole, past president

S. A. Barrett, past president

Committee on Publication

The president, ex officio

The secretary, ex officio

Arthur Beatty, University of Wisconsin

Committee on Library

The librarian, ex officio

Howard Greene, Milwaukee

Mrs. Angie K. Main, Fort Atkinson

Rollin C. Mullenix, Lawrence College

George Van Biesbroeck, Yerkes Observatory

Committee on Membership

The secretary, ex officio

E. F. Bean, State Geological Survey

Ralph N. Buckstaff, Oshkosh

Anselm M. Keefe, St. Norbert College

William N. Steil, Marquette University

Correspondence relating to publication in the Transactions or to other Academy-

business should be directed to the secretary, Lowell E. Noland, Biology Building,

University of Wisconsin, Madison, Wis. Publications intended for the Library

of the Academy should be sent directly to the Librarian, Walter M. Smith, Uni¬

versity of Wisconsin Library, Madison, Wis.

3> o , 7 3

. iyu^ Uj L3

CONTENTS

Page

The Geography of the Upper Rock River Valley; a Study in Cho-

rography. (With 14 text figures). H. O. Lathrop __ _ _ _ 1

The Two Creeks Forest Bed, Manitowoc County, Wisconsin. (With 4

text figures). L. R. Wilson _ _ _ _ _ _ _ 31

Varved Clays of Wisconsin. (With 2 text figures). Elmer W. Ell¬

sworth _______ _ _ _ . _ _ _ 47

The Distribution of Cloudiness in Wisconsin. (With 6 text figures).

Eric R. Miller _ _ _ _ _ _ _ _ _ 59

The Development of the Ice Cream Freezer. (With 6 text figures).

H. A. Schuette and Francis J. Robinson _ _ _ _ _ _ 71

Shakespeare’s Use of English and Foreign Elements in the Setting of

The Two Gentlemen of Verona. Julia Grace Wales _ , _ 85

Cowley as a Man of Letters. Ruth Waller stein _ 127

Wisconsin Myxomycetes. (With Plates I to VI). H. C. Greene _ 141

Notes on Parasitic Fungi in Wisconsin. XIX (With an index to Notes

XVIII and XIX.) J. J. Davis _ _ _ _ 183

Impermeability in Mature and Immature Sweet Clover Seeds as Af¬

fected by Conditions of Storage. Earl A. Helgeson - 193

Preliminary Reports on the Flora of Wisconsin. XV. Polygonaceae.

(With 42 text figures). Kenneth L. Mahony _ 207

Preliminary Reports on the Flora of Wisconsin. XVI. Xyridales.

(With 6 text figures). Norman C. Fas sett - - - - 227

Preliminary Reports on the Flora of Wisconsin. XVII. Myriacaceae,

Juglandaceae. (With 6 text figures). Norman C. Fas sett - 231

Preliminary Reports on the Flora of Wisconsin. XVIII. Sarraceniales.

(With 6 text figures). Florence B. Livergood „ _ _ _ 235

Preliminary Reports on the Flora of Wisconsin. XIX. Saxifragaceae.

(With 18 text figures). Norman C. Fassett - - ,_ 237

Preliminary Reports on the Flora of Wisconsin. XX. Malvales. (With

6 text figures). Alice M. Hagen _ _ _ 247

Standards for Predicting Basal Metabolism: I. Standards for Girls

from 17 to 21. (With 4 text figures). Marian E. Stark - ,_ 251

The Decapod Crustaceans of Wisconsin. (With 13 text figures).

Edwin P. Greaser - - - - - 321

Preliminary List of the Hydracarina of Wisconsin. Part II. (With

Plates VII to X) . Ruth Marshall _ _ _ _ _ 339

A Report on the Mollusca of the Northeastern Wisconsin Lake Dis¬

trict. (With 127 text figures). J. P. E. Morrison - - 359

iv

Contents .

Studies on the Life History of A cello, haldemani (“Desh.” Binney.)

(With 2 text figures, and Plates XI and XII). J. P. E. Morrison 397

Dissolved Oxygen and Oxygen Consumed in the Lake Waters of

Northeastern Wisconsin. (With 85 text figures). C. Juday and

E. A. Birge _ _______ _ _ _____ - - - — _ 415

Races, Associations and Stratification of Pelagic Daphnids in some

Lakes of Wisconsin and other Regions of the United States. (With

Plates XIII to XVIII). Richard Woltereck _ _ _______ _ 487

Solar Radiation and Inland Lakes. Fourth Report. Observations of

1981. (With 7 text figures). E. A. Birge and C. Juday - — 528

Proceedings of the Academy — _________ — — - 563

Constitution and By-Laws of the Academy - .____ - - — — - 569

Subject and Author Index to the Papers published by the Academy,

1870-1932. Lowell E. Noland _ _ — - - ._ 573

THE GEOGRAPHY OF THE UPPER ROCK RIVER

VALLEY: A STUDY IN CHOROGRAPHY

H. 0. Lathrop

Introduction

The Upper Rock River Valley is a region of middle-latitude

mixed farming, diversified dairy production, and a multiform

small-scale industrial life imposed upon a variety of glacial

topographic features left by the late Wisconsin ice sheet. The

landscape varies considerably in detail, but there are broad,

unifying elements which give it homogeneity and differentiate

it from the adjacent regions. The Region under discussion

includes part of the area drained by the Rock River and its

tributaries, extending south almost to Janesville, and small

marginal areas on the west and north lying outside the Rock

River drainage system (Fig. 1). The greatest north-south

extent of the Region from Janesville to Berlin is approximately

85 miles, and the maximum east-west extent along a line from

Middleton to Delafield is about 60 miles, giving an area of about

3500 square miles. The western limits are marked by a transi¬

tion to rougher land, while to the northwest the Region merges

into a sandy country having a less uniform surface with large

areas of marshland. To the east and southeast the Niagara

Escarpment and the interlobate Kettle Moraine, with its north¬

ward extension, mark the beginning of a region of decidedly

stronger relief. The southern extent of the Region is delimited

by the outer margin of the terminal moraine of the Green Bay

Glacier, beyond which the flattish-to-gently-rolling country is in

marked contrast to the region to the north, and partakes strik¬

ingly of corn belt characteristics.

In this Region the unifying elements of the natural and cul¬

tural landscapes serve to set it apart from the adjacent regions.

Marginal areas are generally sufficiently different in character

so that boundaries can be drawn with considerable accuracy.

However, geographic unconformities are the exception rather

than the rule and, for the most part, narrow zones transitional

in character, mark the limits of the Region.

OCT 1 1332

2 Wisconsin Academy of Sciences , Arts, and Letters .

The broad unifying characteristics of the Region vary suffi¬

ciently in detail to make it desirable to divide it into several

sub-regions. It is areally differentiated chiefly by man’s cultural

imprint in the various parts. In Dodge County and the mar¬

gins of adjacent counties, the cheese industry is a conspicuous

development which distinguishes this section from the sur-

Lathrop — Geography of Rock River Valley .

3

rounding areas (Fig. 11). A narrow peninsula of the cheese

district extends in a westerly direction from the northwest cor¬

ner of Dodge County to the margin of the Region. In the south¬

western part the tobacco industry is areally concentrated in

such a fashion as to give a small area individual characteristics

and to set it apart (Fig. 5). Lying between these two and

extending around the cheese section on two sides is a general

dairy district where the milk is marketed at condenseries,

creameries* cheese factories* and city markets. Lying to the

north of the cheese area and extending to the limits of the

Region is a second area where general dairying prevails. Hence,

the following agricultural sub-regions may be designated.

I. The Cheese District

II. The Tobacco District

III. The General Dairy District

IV. The Northern General Dairy District

The Cheese District

The cheese industry of the Region is centered in a striking

way in Dodge County (Figs. 1 and 11) . The northern boundary

of the county is almost coextensive with the limits of the Cheese

District in that direction. To the west the shading off of the

cheese industry occupies approximately one third of Columbia

County, and the southwestern corner of Green Lake County,

where the narrow belt referred to above reaches entirely across

to the western boundary of the Region. The thinning margins

of the industry also extend into the northern border of Dane

County, the northeast corner of Jefferson, and the northwest

corner of Waukesha counties. In nearly all of these marginal

areas cheese making occupies a distinctly secondary position as

compared with other dairy interests, and indicates the decreas¬

ing importance of the industry outward from Dodge County

(Fig. 11). On the south these border areas also represent the

retreating margin of cheese production. To the east and north¬

east thru Washington County the cheese industry joins with

the lake-shore cheese area, but the connecting zone shows a

relatively lighter cheese production than the centers to the east

and west.1

1 Bulletin 90, Wisconsin Department of Agriculture, Madison, Wisconsin, 1928,

p. 75.

4 Wisconsin Academy of Sciences , Arts , amZ Letters .

Types of Relief and their Utilization

Drumlins

Most of the Cheese District has the strongest relief of any

part of the Region. The drumlins cover a greater percentage

of the surface and are higher than elsewhere, and the indivi¬

dual drumlins have a larger areal extent. They also exert a

Lathrop— Geography of Rock River Valley .

5

stronger influence upon man’s utilization of the Region than in

other parts. The long north-south or northeast-southwest axes

are roughly parallel, and the drumlins are located en echelon

so that transportation across or around them is difficult. In

some cases the bases of the adjacent drumlins are almost con¬

tiguous, but the higher ones are usually from one to four miles

apart with lower drumlin-like ridges, rolling lands of gentle

relief, or marshlands lying between. The profound influence of

the drumlins upon man’s utilization of the Region is evident

everywhere. Most of the higher ones have slopes running up

to 15 degrees; 5.8 per cent of the area surveyed falls into the

classification of “steep”, slopes which, by the technique of the

survey, runs from 10 to 15 degrees inclusive.2 The lower gentle

slopes are used for all types of agriculture, but the intermediate

slopes and summits are generally wooded. The timber is com¬

posed of mixed hardwoods of the oak-hickory association. In

many cases the smaller timber has been taken out and the under¬

brush cropped and killed by pasturing so that the woodlands

present an open appearance which permits the growth of blue-

grass or a mixture of similar grasses, thus providing good per¬

manent pastures. However, in some cases rather heavy stands

of timber prevail and the land is little used for pasture because

the density of tree growth prevents the development of grasses

upon the forest floor.

The production of good agricultural crops upon the steeper

slopes and summits of some of the drumlins indicates that the

present utilization of such lands is temporary. It is not clear

that steepness of slope or sterility or rockiness of soil prohibits

agriculture in most cases. The drumlins are composed of a mix¬

ture of unconsolidated glacial clays, sands, and gravels, with

some boulders intermixed. Such soil texture permits the absorp¬

tion of large quantities of water so that erosion and soil wash

are not serious problems, even upon slopes of 10 to 15 degrees.

Due, however, to soil texture and excellent drainage many of

such lands are inclined to be droughty in dry spells in summer.

2 Detailed field surveys were made of three typical areas shown on Figure 1.

These surveys covered a total of 13 square miles. Details such as crops, soil,

topography, slope, and drainage were observed and mapped. The data thus col¬

lected were assembled and totaled and reduced to percentages. These percentages

are considered as typical of the areas in which the survey was made and are

referred to frequently thruout the paper.

6 Wisconsin Academy of Sciences , Arts , and Letters .

Inter-Drumlin Areas

The land between the higher drumlins consists chiefly of two

topographical types, gently-rolling, ridge-like ground moraine

and low-lying, flattish lands, some of which are marshy. The

former are the best agricultural lands in the Cheese District

and comprise about half of the total area. They are well drained

and their gently-rolling configuration is almost ideal for plow

agriculture. The light-brown loamy soils are fertile and pro¬

duce abundantly. Originally they were all covered with timber,

but there is practically none remaining on such lands because

of their high value for agriculture. The glacial influence is

again in evidence in the general north-south trend of the low

inter-drumlin ridges. Many of them are too low and their con¬

figuration is not sufficiently distinct to class them with the other

higher and well-formed drumlins, altho they undoubtedly belong

to the same class of glacial deposits and have a similar origin.3

They furnish good locations for many of the roads which in

some instances follow the crest of such elevations for miles. As

a consequence the highway pattern has a distinctly northeast-

southwest to north-south orientation. Because of their good

drainage, gentle gradients, and accessibility to roads, these

ridges furnish ideal sites for farmsteads and many of the farm

homes are located upon them.

The low-lying, flattish, inter-drumlin tracts range all the way

from well drained to poorly drained farm lands, thru marsh

hay, permanent wet pasture and peat bog, to marshlands cov¬

ered with cat-tails, coarse grasses, and tamarack and willow

thickets. Many of these areas are the beds of former glacial

lakes, the well known Horicon Marsh, containing about 60

square miles, being the largest. Considerable areas have been

drained by open ditches or tile. The water table is so high that

deep ditches of large capacity are necessary to be effective.

Where these lands are well drained they make splendid farm

lands, the soil survey classifying most of them as various mem¬

bers of the Clyde series. These are black soils, high in humus

content, and contain sufficient quantities of silt so that they are

fairly easy to work. Their high fertility makes them especially

desirable for corn, but their low-lying situation makes them

susceptible to unseasonable frosts. It is impossible to drain

3 W. C. Alden and his co-workers in the Quarternary Geology of Southeastern

Wisconsin classify the low ridges as drumlins.

Lathrop— Geography of Rock River Valley .

7

some areas because the intervening lands separating them from

streams is so high as to prohibit the construction of proper

drainage ditches.

Large areas of the wet lands are cut for wild or marsh hay.

In Dodge County, in 1926 and 1927, the 16,000 acres cut for

marsh hay was approximately one and a half times that of

alfalfa, and the yield per acre was more than two thirds as

great.4 The acreage of wild hay is about one fourth as great as

that of all tame hays with one sixth as large a total yield. Such

hay, however, lacks the high nutritious quality of tame hay.

The marsh hay lands occupy a little less than three per cent of

the total area of the county. After the hay is cut these lands

furnish excellent late-summer and fall pastures. They, together

with the adjacent permanent wet pastures many of which are

too wet even for the cutting of hay in midsummer, occupy an

important position in the economy of farm practice. As the

late summer rains decrease in frequency and intensity the

upland pastures often become short, and in times of extreme

drought are entirely inadequate to furnish pasture for the large

numbers of livestock. At such times the wet pastures are at

their best because the ground water is sufficiently high, even

during the most severe droughts, to furnish the grasses with

an adequate supply of water.

The Non-Drumlm Area

The part of the Cheese District lying southwest from Wau-

pun about Beaver Dam and to the west and north is outside the

drumlin country. In this section the relief is gently rolling ;

marshlands are smaller in area; there is less timber ; and the

land, much of which was formerly prairie, is almost ideally sit¬

uated for agriculture. These differences in the natural environ¬

ment have a number of consequent responses in man's utiliza¬

tion of the land. A higher percentage of the area is cropped ;

marsh hay and permanent woodland and wet pasture are almost

absent ; the roads and fields assume a truly rectangular pattern ;

and the linear northeast-southwest arrangement of farmsteads

gives way to a miscellaneous pattern, influenced by promiscuous

elevations in the ground moraine and the road layout.

4 Bulletin No. 90, Wisconsin Department of Agriculture, 1928, pp. 37-42.

8 Wisconsin Academy of Sciences, Arts, and Letters .

Crop Distribution in Relation to the Natural Environment

The crops produced in the Cheese District are similar to those

grown in the remainder of the Region and are typical of dairy

Fig. 3. Corn acreage. One dot equals 50 acres.

Fig. 4. Oat acreage. One dot equals 50 acres.

Fig. 5. Tobacco acreage. One dot equals 10 acres.

Fig. 6. Canning peas acreage. One dot equals 20 acres.

Lathrop— Geography of Rock River Valley.

9

countries in middle latitudes. The grains claim first place and

occupy 71 per cent of the area of cropped lands in Dodge

County5 (Figs. 3 and 4). Hay and rotation pasture cover 24

per cent of the cropped area, thus attesting the importance of

the dairy industry. The remaining five per cent of cropped lands

is given over largely to cash crops, peas for canning being the

chief one (Fig. 6) . Cash crops are not highly significant because

the farmer depends upon the milk check for his cash income.

Crops furnish but 13 per cent of the farmer’s income compared

with 87 per cent from livestock and animal products.6 Very

little hay or grain is sold from the farms. Practically all of it

is converted into the various kinds of animal feeds, and in poor

crop years the farmer is compelled to buy from outside sources

to supplement his own production. However, large purchases

over long periods are rare because the dairymen tends to adjust

the size of his dairy herd to the food producing capacity of his

farm.

Animal Products

The animal population in the Cheese District is large, averag¬

ing more than two animal units per acre for the total area of

Dodge County, which may be taken as typical of the entire

District. The relative importance of the leading animals in

1929 is shown in the following table:7

Table I

From the foregoing table the predominance of cattle and

cattle products is apparent at once. Milk furnishes more than

half of the gross income, to which may be added the returns

6 Bulletin 90, Wisconsin Department of Agriculture, 1928, p. 15.

6 Supplement No. 1 to Bulletin 90, Wisconsin Department of Agriculture, 1929,

p. 21.

1 Ibid , pp. 18-18. (An animal unit equals one cow, one horse, seven sheep, or

five swine.)

10 Wisconsin Academy of Sciences , Arts , and Letters .

from cattle and calves sold off the farm. This gives a total of

64 per cent or almost two thirds of the gross farm income that

is obtained from cattle and cattle products. Practically all the

cattle in the Cheese District are Holstein. This breed is well

Fig. 9. Swine. One dot equals 20 animals.

Fig. 10. Sheep. One dot equals 20 animals.

Lathrop — Geography of Rock River Valley .

11

adapted to the conditions of this District. They give large

quantities of milk which is an important consideration in a

country where most of the milk is marketed at cheese factories.

Holsteins are large animals and heavy feeders which enables

them to handle efficiently the bulky and watery grasses from

the wet pastures which figure so prominently in the farm econ¬

omy of the District.

Horses are used in large numbers thruout the District for

farm work, altho tractors are also utilized for this purpose.

Swine are found on practically every farm and are produced

extensively as a by-product of the dairy industry, being fed

largely upon skim milk from the creameries or whey from the

cheese factories. Their number diminishes markedly in the sec¬

tion where milk is sold to condenseries (Figs. 9 and 13).

The Tobacco-Dairy District

Delimitation of the District

The Tobacco-Dairy District is located in the southwestern

part of the Rock River Valley Region and occupies portions of

Dane and Rock counties. The most intensive cultivation of

tobacco centers in the townships of Dunkirk, Pleasant Springs,

Albion, and Christiana in Dane County, with a long narrow

tongue of heavy production extending northwestward thru the

townships of Cottage Grove, Burke, and Windsor (Fig. 5).

From this center of heavy production the industry shades off

in all directions but with unusual abruptness to the east and

vrest.

In the District thus outlined, tobacco production and dairy¬

ing are the first interests of the farming population and rural

life centers largely about these two activities. In some areas,

and with individual farmers over the entire District, tobacco

is of prime consideration with dairying holding a secondary

place, while in others the reverse is true. In general, the two

interests appear to work well together as the number of dairy

cows does not diminish appreciably in the Tobacco District. At

certain times of the year tobacco makes a heavy demand for la¬

bor and attention on the part of the farmer. However, during

the cultivating season this is partially met by sharemen and

temporary hired help, often women and children. At the time of

harvest most of the work in the tobacco fields must be done in

12 Wisconsin Academy of Sciences , Arts, and Letters

the middle of the day when the dairy herd is out on pasture.

Thus, it is seen that in many ways tobacco and dairying are

supplementary and not competitors for the farmer’s time and

attention.

General Landscape Characteristics

The topography of the Tobacco District varies considerably

in detail. The northeast half lies on the southwest margin of

the short-drumlin area. Scattered drumlins of ovoid configura¬

tion with patches of timber covering the crests of some of the

higher ones characterize the landscape. The long axes of the

drumlins extend northeast-southwest, having a maximum length

of about a mile, although most of them are less than a quarter

of a mile long. The short axes are only a few hundred feet in

length in most cases and the maximum elevation is about 100

feet above the surrounding country. However, the slopes break

abruptly from the contiguous lands and present small areas of

rougher topography. Steep slopes include but 1.4 per cent of

the area. Most of the drumlins are farmed and the patches of

timber on the summits are small. The soils of the drumlins are

coarser than those of lands lying in proximity to them and are

classified by the Soil Survey as fine sandy loams or gravelly

sandy loams. Such soils are distinctly less productive than

adjacent soil types and are especially droughty during dry

spells in summer.

The widely separated drumlins occupy only a small percent¬

age of the area and exert a minor influence upon the utilization

of the land. The prevailing rolling type of topography is broken

by poorly adjusted stream channels, along which are extensive

marshlands and peat bogs which serve as permanent pastures.

Of the area surveyed 9.2 per cent is used for this purpose. The

marshlands gradually become smaller in extent toward the

northwest and are almost absent at the northern limit of the

Tobacco District. Much of the uplands was originally prairie,

which probably occupied half of the total area of the Tobacco

District. These are deep, fertile, black soils of high productiv¬

ity and stand out among the lighter colored Miami soils, which

cover most of the original timbered areas. The prairie soils are

classified as Carrington loams, and their otherwise nearly level

surface is broken by occasional drumlins which become lower

Lathrop — Geography of Rock River Valley .

13

and more scattered to the northwest, and are entirely absent

in the northern extent of the District.

To the south and southeast the Tobacco District extends

across the terminal moraine of the Green Bay Glacier. This

area presents a rougher topographic aspect than other parts of

the District. Small areas of timber remain on the steeper slopes

of the ridges and hills which comprise the moraine. Most of

the land is cleared, however, and some of the finer tobaccos are

grown upon the well-selected slopes and soils of this type of

topography. Edgerton, the chief tobacco market for the Dis¬

trict, lies in the morainal lands. The northwest-southeast trend¬

ing belt of low hills composing the moraine are from one to two

miles in width, and their outer margin marks the southwestern

limit of the Region. South of them the dominantly prairie

lands of the Illinoian Drift present a contrasting type of land¬

scape and mark the beginning of the corn belt type of topog¬

raphy. Tobacco is grown in decreasing quantities south of the

moraine which indicates that the newer soils and varied topog¬

raphy to the north are probably factors favoring tobacco pro¬

duction.

Farm Crops in Relation to the Natural Environment

The usual crop association of middle-latitude grains and hays

is found in the Tobacco District. In the area surveyed 84 per

cent of the land was in crops. The percentage devoted to per¬

manent pasture and timber is correspondingly low, as previ¬

ously noted. Thus the Tobacco District has a high percentage

of cultivated land and a relatively intensive cultivation. Prac¬

tically all the area except the marshlands is in agricultural

crops or rotation pasture.

Tobacco is the only special crop and is the one that gives

distinguishing characteristics to the cultural landscape. Seed

beds covered with cheese cloth in early spring, heavy manur¬

ing of the land in winter, thorough working and preparation

of the soil, the unique planting implement with its tank of

water for watering the transplanted seedlings, intensive cul¬

tivation thruout the summer, the heavy hand labor of suckering,

topping, and cutting, and the final stringing of the heavy wilted

plants on laths to be transported to the tobacco barn, all give

distinction to the cultural landscape of this District. The rough

14 Wisconsin Academy of Sciences , Arts , and Letters.

tobacco barn planked up and down with unplaned unpainted

boards adds a further distinctive feature. Tobacco requires

the attention and interest of the farmer throughout the year

and his activities and equipment distinguish the Tobacco Dis¬

trict from the adjacent areas.

Most of the tobacco is of special quality and is produced for

cigar binders. Thin tough leaves of uniform quality and color

are demanded for this purpose, so that special attention is given

to the choice of soil and topography, to the curing process, and

to the variety of tobacco, in order that the product may enter

the market at a high price for this special use. Slight differ¬

ences in soil change the thickness, color, and flavor of the leaf.

Ignorance or carelessness in curing and processing the crop

after it is in the shed may mean a difference of ten cents a

pound in final receipts. Thus, the tobacco industry requires

much attention and forethought on the part of the grower and

careful selection of soil and topography. “It can truly be said

that tobacco culture is one of the fine arts of agriculture, and

patience, perseverance and care are the three graces which

lead to success.”8

The long practice of tobacco culture in this District gives the

necessary skill and the newly arrived Norwegian immigrant

has supplied much of the unskilled labor. Tobacco culture was

important before the Norwegian immigrants entered. As these

people with large families and but little money came into the

country there was a heavy demand for opportunity to work to

secure a livelihood and to buy a farm. Tobacco offered the

opportunity. They could become sharemen with the landowner

who furnished the experience, land, and capital. As rapidly as

possible they bought their own farms but continued to cultivate

tobacco. They have been willing to do the heavy exacting labor

required by tobacco culture for the large returns to be secured.

The Norwegians have thus become the chief tobacco growers

and have tended to perpetuate the industry wherever they have

gone.9

Tobacco from this belt is shipped all over the United States

for cigar binders. In order to facilitate marketing and to sta-

8 Report, Commissioner of Labor Statistics of Wisconsin, 1883-1884, p. 222.

9 Hibbard, B. H., History of Agriculture in Dane County. Bulletin No. 101,

University of Wisconsin, Economics and Political Science Series, Vol. 1, No. 2,

p. 172.

Lathr op— Geography of Rock River Valley. 15

bilize prices, a pool of tobacco growers has been formed which

claims to control about two-thirds of the acreage. Many large

growers do not belong because they prefer to do their own deal¬

ing and sell when they desire, while others find it difficult to

get sharemen who are willing to wait for the pool to sell the

tobacco.

Edgerton is the chief marketing center for tobacco. The out¬

standing feature in the city is the group of tobacco warehouses.

The tobacco is purchased by buyers and stored there awaiting

shipment elsewhere. This little city is the Mecca for the buyers

in this District. Tobacco and the tobacco market are the topics

of conversation everywhere and the city's chief importance is

due to this product. During the winter hundreds of laborers,

chiefly women, are employed in the warehouses. Edgerton's

leadership in the tobacco trade appears to be largely historical,

as it is not so near the geographical center of the tobacco belt

of today as Stoughton. In the early history of the industry it

was near the center of the tobacco growing district and its

importance has continued because of the momentum of its early

start. Good hotel accomodations for the buyers existed, the

warehouses were built there, and the marketing phase of the

industry appears to be permanently established.

The grains are the largest group of crops in acreage in the

Tobacco District. Corn ranks first as a crop and occupies almost

20 per cent of the cropped area, while oats is a close second

with almost 17 per cent. These two crops are grown largely in

rotation, and occupy the same lands over a period of years.

They are grown almost wholly on the flat and rolling lands.

Corn, in common with other cultivated crops, suffers at times

because so much care and attention are required by the tobacco

crop at the season when corn needs cultivation ; if tobacco needs

attention the other crops are often neglected. Neither corn nor

oats is sold off the farms in any considerable quantity, but

both are fed in the original form or ground with other grains

into mixed dairy feeds.

Wheat and barley are grains of secondary importance in this

belt, as elsewhere in the Rock River Valley Region. Together

they occupy but 6 per cent of the cropped land, of which 5.4

per cent is devoted to barley, leaving to wheat a negligible per¬

centage. This is an interesting transformation from 60 years

ago when wheat occupied the rich prairie lands of this section

16 Wisconsin Academy of Sciences , Arts , and Letters.

almost to the exclusion of all other crops.10 Both crops are fed

upon the farms. Wheat, barley, and oats are sometimes inter¬

mixed in the same field, and given the rather colorful name of

“succotash”.

Hay and rotation pasture together occupy 44 per cent of the

cropped land. If the area in permanent marsh and woodland

pastures be added, a total of 47 per cent of the land is used

for hay and the various types of pasture. This utilization of

land reflects in a striking way the importance of the dairy

industry in the District.

The Dairy Industry

The animal population in the Tobacco District is dense com¬

pared to most sections of Wisconsin or of the United States. It

is not so high, however, as in the Cheese District, the attention

to tobacco reducing the number of animals. A general indiffer¬

ence prevails toward animal products except those related to

dairying. Other animals are produced temporarily or for con¬

venience or work on the farm.

The three chief markets for milk in the Tobacco District are

creamery, city market, and condensery, their relative impor¬

tance being in the order named. Figures 12, 13, and 14 show

where each use is dominant. Competition for milk is keen

among the several markets. In the area surveyed, milk is sold

to all three markets although a slight preference is shown the

creamery.

The General Dairy District

Lying between the Cheese District and the Tobacco District,

extending far around to the east and west of the former, and

completely encompassing the latter, is a District devoted to

general dairying without any specialty such as characterizes

each of the two sub-regions already discussed. The area is lack¬

ing in any large urban influence, except for the section lying

near Madison, altho numerous villages and towns are scattered

over it.

The Short-Drumlin Area

Southeast of a line drawn from Columbus thru Sun Prairie

to Madison, short drumlins are the dominant characteristic of

10 Hibbard, B. H., History of Dane County, op cit, p. 121.

Lathrop — Geography of Rock River Valley ,

17

Fig. 11. Cheese. One dot equals 2 farms selling milk to cheese factories.

Fig. 12. Creameries. One dot equals 2 farms selling milk to creameries.

Fig. 13. Condenseries. One dot equals 2 farms selling milk to con-

denseries.

Fig. 14. City Market. One dot equals 2 farms selling milk to city markets.

the topography, while to the northwest of this line they are

almost absent. Since uses of the land are influenced in many

18 Wisconsin Academy of Sciences , Arts , and Letters

ways by this topographic difference, it seems wise to divide the

General Dairy District into two divisions for general descrip¬

tive purposes, and it is convenient to refer to a similar division

in the discussion of certain phases of human occupance. Here¬

after, these two divisions will be referred to as the drumlin and

the non-dr umlin areas. The drumlins are short and are similar

to those of the Tobacco District.11

Marshes

Drainage is even more poorly developed than in the long-

drumlin country and large areas of marshlands and some poorly

drained farm lands lie between the drumlins. Many of the

lower areas merge into tamarack swamps surrounded by sedges,

cat-tails, and other coarse marsh growth that have little value

even for pasture.12 Hence, there are considerable tracts of idle

or waste lands in some parts of the District. Large areas are

in permanent pasture and others are cut for marsh hay which

plays a similar role in the system of farm economy as in the

Cheese District.

The soil in most of the marshes is peaty. It has a depth of

two to five feet in the smaller, and five to fifteen feet in the

larger marshes.13 The peat soils are widely distributed over the

drumlin section and practically every township has some areas

of such land. Small tracts have been drained and are now used

for agricultural purposes. In general, such soils require con¬

siderable cultivation and attention after reclamation before

they are highly productive for general agriculture. When prop¬

erly conditioned, they produce good crops of corn, hay, and sim¬

ilar crops. Only a small percentage of such soils has been

reclaimed and many of the others are so located as to make it

physically impossible to drain them, while in others drainage

is so expensive as to be prohibitive.

The Rolling Lands

Lying in intermediate locations between the flat lands and

the scattered drumlins are large areas of rolling ground

moraine, similar to such lands already described in the Cheese

“Geib, W. F. and Others, Soil Survey of Jefferson County, U. S. Department

of Agriculture, 1914, p. 56.

12 Geib, W. F. and Others, Soil Survey of Jefferson County, U. S. Department

of Agriculture, 1914, p. 56.

13 Ibid, p. 56.

Lathrop- — Geography of Rock River Valley .

19

District. In the parts of the terminal moraine timber covers

a large per cent of the area. The moraines are much rougher

than the adjacent lands and consist of a miscellaneous collection

of hills and valleys having a variety of soils and numerous

small lakes scattered thru the depressions. The Lake Mills

recessional moraine extending thru Lake Mills, Jefferson, and

Hebron is a belt of hills roughly concentric with the terminal

moraine and marks a tract of rougher land, extending trans¬

verse to the general trend of the drumlins. In this area the

characteristic morainal features predominate.

The Non-Drumlin Area

The northwestern non-drumlin section consists of long

stretches of upland having a gently undulating to rolling topog¬

raphy and black prairie soils. The prairie soils are interspersed

with tracts of Miami soils which range all the way from silt

loams thru the sandy loams and, near Wyocena, merge into the

dominantly sandy soils, which mark the northwestern extent of

the Rock River Valley Region.14 The drumlins are absent from

the surface profile in this section and much of the topography

has an almost monotonous uniformity. The soil maps show

the various phases of soil types trending northeast-southwest,

corresponding to the movement of the ice.15 The chief varia¬

tions in the topography are the rounded rolling hills of the

ground moraine and small valleys where shallow streams have

cut their channels. Drainage is better developed than in the

drumlin section but small tracts of marsh and peat are found

along streams, in depressions which have not had time since

the glacial period to become drained by natural processes, and

in old glacial stream channels like the one extending from

Cambria to Pardeeville.16 However, the areas of marsh and

wet lands are small compared to those of the drumlin area.

In some places near the western margin of the Region, ledges

of sandstone protruding thru the mantle of glacial till show

the character of the bed rock and indicate that the glacial debris

is thinning as proximity to the western limit of glaciation is

reached. The lowest land values in the Region are found along

“Whitson, A. R., General Soil Map of Wisconsin ,

15 Gieb, W. J. and Others, Soil Surveys of Dane and Columbia Counties, U. S.

Department of Agriculture, 1915 and 1913, Soil Maps.

16 Gieb, W. J. and Others, op cit. Soil Map.

20 Wisconsin Academy of Sciences , Arts , Letters .

the northwestern margin where contact is made with the infer¬

tile sand soils lying to the northwest. The per acre value is

only about 40 per cent as great as some of the better lands of

the Region.

Human Occupancy

Human occupancy in the General Dairy District has many

similarities to that of the Districts already discussed. Crop

associations are similar but there is a somewhat greater diver¬

sity of crops and they are grown to a greater extent on the

flatter lands. The following table shows the percentage of land

devoted to the various crops in Jefferson County:17

Table II

The general character of the crops, their use, and function are

practically the same as in the Cheese and Tobacco Districts.

Tame hay is supplemented by large quantities of marsh or wild

hay. It has about one fourth the area of tame hay and is used

extensively for winter feed.

The following table shows the chief sources of farm income

in Jefferson County for 1927, which is typical of the District.18

Table III

The outstanding dominance of animal products is apparent

at once. The minor importance of farm crops for cash income

17 Bulletin No. 90, op cit, p. 15.

18 Supplement No. 1 to Bulletin No. 90, op cit, p. 21.

Lathr op— Geography of Rock River Valley.

21

emphasizes their utilization as feed for the dairy and farm ani¬

mals. Milk constitutes a large element in the food of hogs and

poultry.

The various markets for milk are shown in the following

table, the figures indicating the per cent of milk sold to each

type of market in Jefferson county in 1927.19

Table IV

The wide variety of markets for milk gives the basis for the

regional classification of this section as the General Dairy Dis¬

trict.

The condensery market is limited chiefly to the southeastern

half, including Jefferson and the margins of the adjacent coun¬

ties in the Rock River Valley Region. Six large condenseries

draw milk from this part of the District, where almost ideal

conditions exist for this phase of the dairy industry. An old

well-established dairy industry of great intensity and produc¬

ing large and dependable quantities of milk insures a regular

and continuous milk supply to the expensive condensery plants ;

adequate railway transportation moves the canned product to

the city market or the seaboard if it is to be exported ; and the

fluid-milk market, which is the only competitor that tends to

supplant the condensery is only beginning to make itself felt.

Jefferson County sends 13 per cent of its milk to city mar¬

kets, a large part of which is shipped out of the state. The

receiving stations at Sun Prairie, Oregon, Milton, and Rich¬

mond all collect milk for the Chicago market, while trucking

service to Milwaukee reaches the Palmyra- Whitewater district.

Creameries are important as a market for milk thruout the

General Dairy District. In many ways they form the backbone

of the dairy industry and decline only where there is severe

competition for the other markets. They have their greatest

18 Bulletin No. 90, op cit, p. 74.

22 Wisconsin Academy of Sciences , Arts , cmd Letters .

intensity in the western half of Jefferson County and the row

of townships along the eastern side of Dane County. This is

an extensive rural section away from cities and in which no

condensery is located. A similar area exists along the western

margin of the Region in central Columbia County and extends

southward into northern Dane County, where it joins with the

first district in the northeast corner of this county (Fig. 12).

The cheese industry was important at one time over much of

the District. Competition of other markets for milk has pushed

this industry northward until now it is important only along

the northern margin. There is every indication that its retreat

will continue as the condensery and fluid milk markets extend

their areas.

This District has the same types of ‘other cattle' as indicated

in the Districts discussed previously. In the southeastern part

considerable attention is given to the production of blooded

stock because in this section the dairy industry is old and pre¬

sents more mature aspects. Some beef cattle are produced along

the southeast margin because of the proximity to the corn and

beef region to the south. Hogs are grown extensively thruout

the District but are especially numerous in the creamery sec¬

tions, in the grain producing areas in northern Dane and south¬

ern Columbia counties, and along the southern margin near to

the hog region of the corn belt. Sheep are produced to some

extent on rougher lands thruout the District but are most im¬

portant in a small area in northern Dane County.

The Northern Dairy District

The area lying between the Cheese District and the northern

boundary of the Rock River Valley Region has general char¬

acteristics similar to those of the non-drumlin section of the

General Dairy District. The chief reason for separating it from

the latter is that the two are not contiguous, being divided by

a northwestward projection of the Cheese District. The two

General Dairy Districts have similarity, not only in human

occupance but their natural characteristics show many resem¬

blances. The northern District is a small area and includes

minor parts of Green Lake, Fond du Lac, and Winnebago

Counties (Fig. 1).

Lathrop — Geography of Rock River Valley .

23

Much of the western half of the District is an undulating to

gently rolling upland. It contains the second largest area of

prairie soils in the Upper Rock River Region. This prairie area

extends in a rough semi-circle westward from Waupun, curves

northeast to the south of Green Lake, and reaches northward

past Ripon to the vicinity of Rush Lake, with one prong pro¬

jecting on either side of this lake. Favorable topography, con¬

genial climate, and the black, fertile, silt loams, 12 to 14 inches

deep, make this one of the finest agricultural areas in the

Region. This is emphasized by the various crop and animal

maps. To the west and northwest of the prairie belt is a strip

of timber-land soils lighter in color and having a somewhat

rougher topography. These merge at the western boundary of

the Region into a country of decidedly stronger relief and dom¬

inantly sandy soils.

The eastern half of the Northern Dairy District is made up

chiefly of a series of low, roughly concentric, recessional

moraines, interspersed with extensive areas of marshlands

lying between the low morainal ridges. The morainal areas and

the adjacent intermediate slopes are fair agricultural lands

where they are not too gravelly and sandy, but in general they

are much inferior in their agricultural adaptations to the rich

prairie lands lying contiguous to them on the west. The exten¬

sive marshlands, which are conspicuously absent in the western

half of the District, have been drained only to a limited extent

and are used chiefly for permanent pastures and marsh hay as

similar lands are in the southeastern part of the Rock River

Region. To the north a low moraine, about 50 feet in elevation

and a mile wide, extending from northwest to southeast, marks

the southward limit of a temporary re-advance of the Green

Bay Glacial Lobe and the outer limits of glacial Lake Oshkosh.

To the south and west extensive marshlands border the moraine

while to the north the nearly flat surface, mantled with red

glacial clays, slopes gently to Lake Winnebago. This moraine

forms the northern limit of the Rock River Region. From the

northeastern extremity of Horicon Marsh, the Niagara Escarp¬

ment extending in a broad semi-circle toward the northeast,

rises abruptly 50 to 150 feet above the old lake plain lying to

the west and marks the beginning of rougher topography to

the east. This escarpment forms the eastern boundary of the

Region. The recessional moraines noted above die out to the

24 Wisconsin Academy of Sciences , Arts , Letters .

east and the flat plain adjacent to the Niagara Escarpment,

representing the old basin of Lake Winnebago, is excellent farm

land. There are but small areas of marsh but large areas of

the Clyde soils require drainage.20 The Poygan clays and the

Clyde soils, when properly drained, rank high in natural fer¬

tility.

Crops

The same major crop associations are found in this north¬

ernmost of the Districts as in the remainder of the Region. Cli¬

matic conditions and natural factors are essentially the same

as elsewhere. Ripon, lying near the center of the District, has

a growing season of 158 days which is approximately the same

as that for stations situated farther south.21

Peas for canning are grown extensively thruout the District

and show their greatest intensity of production in Green Lake

and Mackford townships in the southeastern part of Green

Lake County (Fig. 6). This county derives 7 per cent of its

gross farm income from canning peas and ranks first in the

state in this respect. Markesan, a small town in the heart of

the area of intensive pea cultivation, has three large canning

factories handling peas exclusively. In this section the viners,

at frequent intervals along the main highways, are a distin¬

guishing feature of the landscape, while in late summer the

odor from the numerous stacks of fermenting vines is a notice¬

able characteristic of the District.

Hemp attained considerable importance in this section of

the state during the World War, but with the return of cheaper

cotton after the war, hemp prices dropped so low as to make

the returns from it unsatisfactory and its production declined.

It is still grown to some extent and is marketed at hemp mills

located at Waupun, Markesan, and Fairwater. It is used chiefly

for making various types of cord and cheap ropes. There is no

indication that its production will reach larger proportions than

at present.

Animals and Animal Industries

The animal population shows the usual characteristics

observed over the major Region. All types of farm animals are

20Gieb, W. J. and Others, Soil Survey of Fond du Lac County , United States

Department of Agriculture, 1913, pp. 32, 37.

21 United States Weather Bureau, Section 60, Eastern Wisconsin.

Lathrop — Geography of Rock River Valley . 25

grown and, with the exception of sheep, show an almost even

distribution over the District. In general they have about the

same density and importance as elsewhere.

Cattle and the dairy industry continue to assume a role of

first importance. The density of dairy cattle is slightly less

than in Dodge and Jefferson Counties but is about the average

for the Region. Holsteins are the leading type of cattle pro¬

duced. The following table shows the various breeds and their

relative importance as reported by the Wisconsin Livestock

Breeders Association for Green Lake County.22

Table V

Milk is sold to all of the four types of market, which justifies

the designation, the General Dairy District. The creamery is

the most important market but it is confined chiefly to the west¬

ern two thirds of the District (Fig. 12). This area represents

one of the most intensively developed creamery markets of the

entire Region, 97 per cent of the milk of Alto township being

marketed at creameries. They are unimportant in the eastern

part of the District due to the southwestward extension of a

cheese area from Lake Winnebago. Over 60 per cent of the

farmers in Eldorado township dispose of their milk to cheese

factories. South of the cheese area the milk market is dom¬

inated by condenseries located at Ripon and Berlin. Fluid milk

for cities is the least important of the major milk markets.

Marginal areas to the southwest, the southeast, and the north¬

west patronize this type of market. The localized character of

this market is shown on Figure 14.

Considerable attention is given to the production of beef cat¬

tle, which are about half as numerous as dairy cattle. Some

farmers raise beef cattle while others import steers from the

ranges and fatten them for the market. Swine are important in

22 Soil Survey of Green Lake County, op cit, p. 1805.

26 Wisconsin Academy of Sciences , Arts , cmd Letters.

the creamery and prairie sections because of an abundance of

food in the form of grain and skim milk. This District contains

the largest and most intensive sheep producing area in the

entire Region. Their production is concentrated most strik¬

ingly in the vicinity of Green Lake. The utilization of the

rough topography for pasture lands and the abundance of pea

vines for pen feeding may help to explain this distribution.

Transportation

The Region has excellent rail and highway transportation

facilities. Its location near to the two important railroad centers

of Chicago and Milwaukee cause many railroads to cross the

Region as they converge upon these two main centers. The

location within the Region of Madison, a railroad center of sec¬

ondary importance, further increases the network of railways.

Railroads, connecting lake shore cities of Wisconsin with the

Twin Cities and the West, are crossed by lines linking the indus¬

trial Fox-Winnebago Valley and Duluth-Superior and the north¬

west with Chicago. This crossing of railroad lines gives almost

a surplus of railroad transport. All cities and most of the vil¬

lages are located upon railroads and no part of the Region is

handicapped by distance from rail lines.

The highway system of the Region is characterized by the

same well-developed network of roads as the railways. The

locational factors affecting railway development have had a

similar influence upon highways. Many of the concrete roads

parallel the railroads. The intensity of dairy development

requires adequate transportation. Hence roads are numerous

and well kept. Even minor cross roads are usually surfaced

with gravel, an abundance of which is found in glacial deposits

covering the Region. In many cases the gravel pit is located in

the drumlins, eskers, and moraines adjacent to the road being

surfaced. Many county highways are oiled and snow plows

keep them passable in winter because milk must reach the mar¬

ket daily.

Cities

Madison is the only large urban center in the Region.23 Water-

town is the only other city reaching a population of 10,000,

23 Madison is not discussed in this paper. A full treatment is given in a Master’s

dissertation by Cyril Stout, 1931, University of Wisconsin.

Lathrop — Geography of Rock River Valley . 27

altho Beaver Dam falls but slightly under that figure.24 All of

the other cities except Fort Atkinson are under 5,000. Thus,

in general, the Region is characterized by a considerable num¬

ber of towns and small cities from 1000 to 5000 population and

numerous villages and cross-road hamlets. There is little tend¬

ency for any of the cities except Madison to show any marked

growth. Most of them show a small increase in population

during the last census decade. The smaller towns and villages

present a static or declining population and are chiefly impor¬

tant as trade centers for groceries and such staples as are

required by the immediate rural communities. Even this impor¬

tance is threatened as better roads and automobile facilities

enable farmers to reach larger shopping centers in a few min¬

utes. This influence is also affecting the growth and importance

of the larger towns and cities as shoppers can reach the urban

centers such as Madison and Milwaukee in one to two hours.

Without doubt, this is the most important factor in the slow

growth of most of the cities of the Region.

Inland waters have played an important part in the location

and development of the cities. Most of them are located on the

Rock River and its tributaries or upon lakes. Water navigation

has had practically no influence upon the growth of the cities

of the Region, altho there is record of a steamboat ascending

the Rock River as far as Jefferson. The river is too shallow

and has too many rapids and falls to be of value commercially.

Early attempts to connect it with Lake Michigan or to canalize

it failed. Most of the cities grew up around water power sites.

These water powers are all small but they served the early com¬

munities for grinding flour and turned sawmills to saw lumber

for the farm buildings. Several of them are now used to fur¬

nish part of the power for modern factories, but most of them

are used to grind feed. This importance should not be mini¬

mized in a dairy community which uses large quantities of

ground feeds. The streams or lakes have been an important

factor in the growth and development of the cities. Water for

industrial plants, disposal of sewage and industrial wastes,

natural ice supply, bathing beaches, and sites for tourist camps

are some of the more important services of inland waters to the

people of the Region.

24 United States Census Bureau Bulletin, Population of Wisconsin, 1930.

28 Wisconsin Academy of Sciences , Arts , and Letters .

Industries

The industrial development within the Region is exception¬

ally high for a district of dominantly rural characteristics.

Many of the industries are the outgrowth of basic raw mate¬

rials for manufacture or are a response to the demands of the

community for specific types of manufactured goods. Because

of this quality they will be referred to as “community indus¬

tries” to distinguish them from those that have no basis in raw

materials or markets in the community in which they are

located.

Community Industries

Since dairying is the key agricultural interest of the Region

it is natural to find that many of the industries are closely

related to it. Cheese factories, creameries, condenseries, and

ice cream factories are found throughout the Region, and as

indicated above are more or less concentrated in the several

Districts of the Region. Many of them are small, but taken as

a group they constitute the most significant manufacturing

industry of the Region. All of the cities, every village of impor¬

tance, and almost every cross-road hamlet has one or more milk¬

using industries. Rarely can one travel more than five to ten

miles across the Region in any direction without seeing one

or more factories of this type; and in this distance several of

the smaller types such as creameries and cheese factories may

be observed. Practically all the cities have several milk-using

factories. Some of these, as in the case of condenseries, are

large, involve the investment of large sums of money, have a

large pay-roll, and give a market for milk within a radius of

ten to fifteen miles. Practically all of the condensed milk is

shipped out of the state, much of it being exported to foreign

countries.

Several factories are located in the Region for the purpose

of supplying the demands of the dairy industry. The James

Manufacturing Company at Fort Atkinson specializes in the

manufacture of barn equipment and is said to be the “the larg¬

est in the world” producing this type of goods. The factory

has two units with 13 acres of floor space and employs 700 to

800 laborers. Their annual business runs into the millions of

dollars, of which, about 10 per cent represents exports. The

Lathrop — Geography of Rock River Valley .

29

Creamery Package Company with headquarters in Chicago and

with plants at Fort Atkinson and Lake Mills manufactures

equipment for the milk-using factories. The heavy bulky char¬

acter of the product makes it desirable to have the factory

located as near the market as possible. Some other small fac¬

tories manufacturing similar farm and dairy equipment are

found in various cities. Large can manufacturing plants at

Waupun and Oconomowoc supply condenseries and canning

companies with tin cans for their product.

Feed mills are numerous and well distributed thruout the

Region. Practically every village and cross-roads community

has one and several are found in the larger cities. Many of

them have inherited the water power sites from the grist or

saw mill of earlier days. In general, the individual plants are

small but the total volume of business is large.

The canning industry in the northern two thirds of the

Region is second only to the dairy industry. Canning factories

are located in practically every hamlet and village in that part

of the Region. Wisconsin produces about one half of the can¬

ning peas of the United States, and approximately a quarter

of Wisconsin's production, or almost one eighth of the total

for the United States, is produced in this small Region. Some

corn is packed in the factories in the southern half of the Region

and small quantities of fresh vegetables are canned in a few

of the factories. It is peas, however, that give volume and sta¬

bility to the canning industry. Small plants supplying canning

factories with equipment are located at Columbus.

A few mineral industries based upon local raw material are

found. An abandoned iron and steel plant at Mayville was the

largest. Plants producing lime and various types of limestone

products are located along the Niagara escarpment near May¬

ville, at Madison, and at various other points in the Region.

Brick and tile for local use and based upon glacial clays are

produced at Whitewater, Jefferson, and Watertown. Sand and

gravel are obtained in many places and the landscape is mar¬

red by the ugly scars of the abandoned pits. Core sand for

foundries is produced extensively near Berlin.

At several places in the Region small wood-working estab¬

lishments present relict forms of an earlier industrial adjust¬

ment. Such types are the table slide factories at Watertown,

the small wood-working establishments at Jefferson and Stough-

30 Wisconsin Academy of Sciences, Arts, and Letters .

ton, and wagon and carriage factories which have recently-

closed down. Local timber at one time supplied these or similar

parent industries but the raw material is now shipped in.

Commercial Industries

This type of industry ships in the raw material and markets

its production largely outside of the Region. Most of them are

located in the Region due to the interest of local capital or are

units of large factories located outside the near-by urban cen¬

ters to take advantage of cheaper rents and of lower wages in

an unorganized labor market. The group producing iron and

steel products is the most important. This includes the farm

seeders at Horicon, the Western Malleables Company at Beaver

Dam, the Malleable Iron Range Company of the same city, the

Otto Biefield Company of Watertown, producers of boilers and

concentrators, the Kissel Motor Company at Hartford,25 and

the Highway Trailer Company of Edgerton. Some of these

companies employ several hundred men, and have an annual

business which reaches into the millions of dollars. The size and

variety of this type of product is surprising.

Seven shoe factories and six knitting mills are found in the

Region. Some of these plants are large, employing 400 to 500

laborers. Most of them represent units of large industries with

headquarters at Milwaukee or Chicago. They are located in the

Region to take advantage of cheaper rents and lower labor costs.

The proximity of the Region to the large urban centers and the

excellent transportation available make the small cities a good

location for these units.26

25 The latest information indicates that this plant is going out of business.

26 A number of other small factories of a miscellaneous nature are omitted from

this survey because space does not permit adequate description.

This paper is an abridgment of a dissertation submitted to the faculty of the

University of Wisconsin in candidacy for the degree of Doctor of Philosophy. Ab¬

breviation of the paper has necessitated the omission of much data, many pic¬

tures, most of the maps, many footnotes, and the bibliography.

THE TWO CREEKS FOREST RED,

MANITOWOC COUNTY, WISCONSIN

L. R. Wilson

Introduction

After the theory of continental glaciation was accepted, it

was not until more extensive study of glacial deposits was made

that the theory of multiple glaciation was advanced (see review

in Thwaites, 1927). The occurrence of weathered soils, loess,

certain lake deposits, and organic remains between glacial tills

are evidences of intervals of deglaciation between ice advances.

During the periods of deglaciation, several of which were of

considerable length, plants and animals spread northward over

the ice-free areas. In many places exposed by the retreat of the

glaciers, forests established themselves and thrived until the

glaciers again advanced and destroyed them. Many deposits of

organic materials buried by till have been found in various

parts of the world, and where these remains are of trees some

of which are in place, the deposit is spoken of as a “forest bed”.

The Two Creeks Forest Red is such a deposit. It is exposed

on the shore of Lake Michigan two miles east of Two Creeks,

Manitowoc County, Wisconsin, in Sections 11 and 13, Town¬

ship 21 North, Range 25 East. Exposures extend along the

shore for about a half mile. The same forest bed is also exposed

three miles to the north on the lake shore, and in a ravine about

a quarter of a mile to the west in .Section 35, Township 22

North, Range 24 East, Kewaunee County. Other exposures

doubtless exist, but time has not allowed further field work.

The deposit studied in detail by the writer is at the first-

named locality. It is several inches thick and approximately one

hundred feet long. The forest bed lies between the varved clays

deposited during the retreat of the Middle Wisconsin or “Gray

ice” and the till of the Late Wisconsin or “Red ice”.

Previously no extensive study has been made of the Two

Creeks Forest Red, though a notice was published by Goldth-

wait (1907) who discovered the exposure. From 1922 to 1930

F. T. Thwaites visited the area several times, but his examina-

32 Wisconsin Academy of Sciences , Arts , and Letters .

tions were brief. In 1922 he submitted some wood to the Forest

Products Laboratory at Madison, and one species of tree was

identified. In 1926 he sent some mollusks to F. C. Baker at

Urbana, who identified three species. Conditions for examina¬

tion were poor during most of this time on account of the low

level of the lake, which checked wave work.

Other forest beds in Wisconsin have been noted in well drill¬

ings, and occur most abundantly in the southeastern part of

the state (Alden, 1918. pp. 177-179). Lawson (1902) published

a preliminary report on the forest beds of the lower Fox River

valley in which he described extensive deposits. These appar¬

ently contained much valuable material, but the exposures have

nearly all been destroyed.

Geology

Descriptive Geology

The clay banks in which the Two Creeks Forest Bed is

exposed show the following geological formations :

The lowest layer exposed is till deposited by the Gray ice,

presumably of “Middle” Wisconsin age. The drift that is com¬

monly called “Middle” Wisconsin in this state is Leverett’s

third substage of the Wisconsin Stage of glaciation (Leverett,

1929).

The Gray till is overlain by a glacial lake deposit of poorly

varved red and gray clays, interbedded with lenses of sand

and silt. The deposit is about twelve feet thick, and the contact

of the varved clay and the Gray till is shown in Fig. 2. It was

not possible to count the varves because of their imperfect

structure and the amount of disturbance of the strata.

The forest bed is found on top of the varved clays, and above

that are several inches of silty sediment. Red till, presumably

of “Late” Wisconsin age (Leverett’s “substage four”), overlies

the thin lake sediments. At the forest bed the till is about eight

feet thick. On top of the Red till are local varved clays of Lake

Algonquin.

In an unpublished report on the glacial geology of part of

northeastern Wisconsin, F. T. Thwaites says : “Although large

numbers of exposures of Red and Gray tills have been exam¬

ined, no evidence of interglacial weathering or soil development

has been observed except at the forest bed near Two Creeks.

Wilson — Two Creeks Forest Bed .

33

The recent rise of the lake level has shown definitely that the

organic deposit at that place rests upon imperfectly varved silty

clays, which have been much disturbed by the work of the Red

Fig. 1. General view of the Two Creeks Forest Bed exposed on the shore

of Lake Michigan. (Photograph by F. T. Thwaites.)

Fig. 2. Contact of the Gray Till and the silty varved clay. (Photo¬

graph by F. T. Thwaites.)

ice. These sediments are seven to twenty feet thick and rest

upon Gray clayey till, which may be presumed to be of Middle

Wisconsin age.”

34 Wisconsin Academy of Sciences , Arts, and Letters .

Historical Geology

The oldest formation exposed at the Two Creeks Forest Bed

is the till of the Gray ice. This till was probably deposited dur¬

ing the Middle Wisconsin Substage during which the ice

extended to the lower end of Lake Michigan.

With the recession of the Gray ice the water in the Lake

Michigan basin rose to a height of about sixty feet above the

present lake level, forming what is known as the Glenwood

Stage of Glacial Lake Chicago (F. T. Thwaites, unpublished).

The lake then drained through the Des Plaines River at Chi¬

cago ; sediments of this lake were assorted into separate layers

known as “varves”. Of these, the coarser silty material was

deposited during the summer and the fine clays, mostly red,

were laid down when the lake was frozen over in winter. They

are aptly called “the annual rings of the earth”. The varves

of Lake Chicago occur above the Gray till and are best defined

near the bottom of the deposit. Near the top the varves are

indistinct and show that the lake was gradually becoming

warmer as the ice receded toward the north.

The Gray ice retreated northward until the Straits of Mack¬

inac were freed and there was a fall of the lake to a level lower

than the present, flow much farther north the margin of the

Gray ice retreated is not known. The Two Creeks Forest was

developed on the land thus formed.

The interval in which the forest bed developed was at least

eighty-two years in length, as shown by the growth rings of

the oldest log, and the time necessary for the establishment of

the forest would undoubtedly increase this figure several times.

The forest bed interval came to a close with the advance of the

Red ice.

Before the destruction of the Two Creeks Forest by the Red

ice, there was a flooding of the forest floor and a deposition of

several inches of sediment on top of the organic remains. The

water that flooded the forest came either from the rising lake

caused by the Red ice blocking the outlet of Lake Michigan at

the Straits of Mackinac or by streams and ponds beside the

advancing glacier.

The Red ice overrode the forest bed and moved as far south

as Milwaukee, where the outer margin of its drift is found.

The water rose at Chicago to a height about forty feet above

the present level of Lake Michigan, and the outlet was again

Wilson — Two Creeks Forest Bed.

35

through the Des Plaines River. This condition is known as the

Calumet Stage of Lake Chicago.

The striking difference in color between the tills deposited by

the two glaciers gave rise to what was long an unsolved prob¬

lem. Alden (1918. pp. 314-315) suggested that it is due to the

ice overriding red clays deposited in Lake Chicago. These clays

were colored by waters from the iron regions of northern Michi¬

gan. However, the clays deposited during the retreat of the

Gray ice are not particularly red. Possibly a more sound sug¬

gestion has been advanced recently by F. T. Thwaites, namely,

that the Red ice came from a more westerly center than did

the Gray. With an advance from this direction several of the

iron ranges of northern Michigan were passed over and this

gave the red color directly to the till. The ice of the later sub¬

stage reached the forest bed by crossing the low northern part

of Door County, then spreading out to the southwest over the

Lake Michigan lowland.

The retreat of the Red ice left the forest bed buried under

eight to twelve feet of red clay till, and the exposure of the

organic remains has been effected by post glacial cliff forma¬

tion. The clay banks are rapidly receding whenever the level

of Lake Michigan is high enough for effective wave erosion.

Organic Remains

Wood

All the wood thus far examined has been of one, or possibly

two, species of spruce Picea mariana (Mill.) BSP. and P.

canadensis (Mill.) BSP. It is interesting to note that all

interglacial wood found in Wisconsin to date has proved on

critical examination to be either spruce or hemlock. Of the

latter species only one record is known ; this is a specimen taken

from the excavation for the Schroeder Hotel at Milwaukee, and

reported to the writer by Mr. D. Costello of Marquette Uni¬

versity. The geological formations which overlay this log are not

known. Other kinds of trees recorded from the Chicago region

(Baker, 1920 p. 5) are extinct species of spruce and oak.

Like much other wood of interglacial age, the Two Creeks

Forest Bed material is soft and easily broken. It checks and

breaks into short sections on drying. The tissues, however, are

not destroyed, and microscopic sections have been made of them.

36 Wisconsin Academy of Sciences , Arts, and Letters .

Where wood and peat have been in contact with the Red till,

there is a zone in the clay a few inches wide of greenish gray

color due to deoxidation.

Fig. 3 Fig. 4

Fig. 3. Spruce log exposed from sediment layer, showing the usual di¬

rection in which the logs are found pointing. (Photograph by F. T.

Thwaites.)

Fig. 4. Spruce stump in situ with its log exposed on the left, also show¬

ing the root on which the fossil bracket fungus was attached. (Photograph

by F. T. Thwaites.)

The logs occur most frequently in the layer of sediment above

the forest bed, where they evidently fell after being broken

from their stumps by the glacial ice. Most of the logs are found

pointing toward the southwest (Fig. 3), and it is thought that

here the last ice moved in that direction.

Wilson — Two Creeks Forest Bed.

37

One stump was found in situ (Fig. 4), with the butt of the

broken-off log almost attached. The roots of this stump extend

along the forest bed peat and on a portion of the root that had

been exposed above the ground during the interglacial period

was found a bracket fungus. It is a Polyporus , but the species

has not been determined. Goldthwait (1907. p. 61) also observed

a stump in situ when he discovered the Two Creeks Forest Bed.

All the logs that have not been broken by subsequent hand¬

ling show ragged splintering ends in consequence of glacial

action upon the trees. This condition indicates a violent twisting

and bending of live trees before they were felled.

The growth rings were studied in sections of six logs. The

greatest number of rings shown in one section is eighty-two;

the average is about sixty. Five of the logs showed by the width

of successive rings a marked decrease in the rate of growth

in the last twelve years of the Two Creeks Interval. One log

taken from the Red till, which is above the forest bed, showed

but little decrease until the last year of its growth. This parti¬

cular log may be white spruce, Picea canadensis (Mill.) BSP.

The other logs studied have been taken to represent the grow¬

ing conditions in the forest bed, whereas the log taken from the

till above is considered as having been transported by ice from

a different environment farther north. When the log taken

from the Red till is compared with the others, an extreme dif¬

ference in size and growth rate is noticeable. This log is twice

the diameter of any of the others, though it has only the aver¬

age number of growth rings. The width of the rings does not

agree with that of the forest bed trees. There is no growth

decrease until the last year of the tree’s life, which fact alone

suggests conditions unlike those at the Two Creeks Forest.

The growth rings cannot be compared exactly with reference

to particular years, for it is not known whether all the trees

were destroyed in the same year or whether they were all alive

at the time of the ice advance ; however, it is considered prob¬

able that the largest log, having been transported by the ice,

was felled several years before the Two Creeks Forest trees.

It is not probable that more time elapsed, for in that case the

log would probably show more evidence of crushing.

Close study of wood sections has shown that certain small

growth rings of the forest bed trees occur at years approxi-

38 Wisconsin Academy of Sciences , Arts , and Letters .

mately corresponding to those in which wide growth rings

occur in the log from the Red till, and vice versa. If excessive

moisture (Zon and Averell, 1929) was one of the primary fac¬

tors for small growth rings in the forest bed trees, as is sug¬

gested by the character of the flora and fauna, then trees grow¬

ing on higher ground would not have been similarly affected

and probably throve better in wet years. In dry years the forest

bed trees, having more favorable conditions, would grow more

rapidly while those on the upland would be adversely affected,

and consequently the rings formed in those years would be nar¬

rower. Other factors, such as temperature and wind, may also

have affected ring growth, but the primary factor here con¬

cerned seems to have been moisture.

Mosses

The moss flora of the Forest Bed comprises the most exten¬

sive group of plants found in the remains. The moss material

was submitted to Mr. L. S. Cheney, who has identified nineteen

species. Mr. Cheney (1930) has published a notice of these

mosses in which he reports eight species. The additional eleven

species that he has since recognized came from other parts of

the same exposure.

All the mosses recognized are of existing species, and are in

general more northern in their modern distribution than the

Two Creeks Forest Bed. Nearly all are found in northern Wis¬

consin, but the present southern limits of a few are in Canada.

The northern limit of all the species is generally in northern

Canada and northern Europe. Several species range from the

Bering Straits to Spitzbergen and Siberia. One species, Swartzia

montana (Lamk.) Lindb., is found from the Arctic to the Ant¬

arctic, although in lower latitudes it is restricted entirely to

alpine regions.

There are two ecological horizons shown by the mosses, and

it is interesting to note how well they agree with the other

organic remains in these horizons. The problem of succession

however, will be considered under the subject of ecological

history.

Peat

The peat of the forest bed is poorly formed and at some parts

of the exposure is wanting entirely. It is evident from this con-

Wilson — -Two Creeks Forest Bed .

89

dition as well as from other organic remains that the Two

Creeks Forest Bed was not exactly a lowland forest, but rather

a dry forest, at one stage of its existence. There are places

along the exposure where the mosses and other plant remains

have accumulated as a silty peat, such as can be found in any

spruce forest today. It is from this peat that the microfossils

were secured, though their number was comparatively small.

The usual technique used by paleo-ecologists was employed

in the study of the pollen and spore fossils (Ertman, 1931).

Identification of pollen grains and spores was checked by refer¬

ence to prepared slides of recent material of the same species.

Mollusks

Seven species of mollusks have been recognized from the

forest bed by Mr. F. C. Baker, to whom specimens were sub¬

mitted. These were from three levels in the forest bed, and

agree ecologically with other organic remains from their respec¬

tive horizons. One Pleistocene form was reported; this came

from the clay immediately beneath the forest bed. Other indi¬

viduals found in higher levels represent existing species.

Ecological History

The ecological history of the Two Creeks Forest Bed is

remarkably well defined in the thin layer of organic remains

exposed.

As already described, the forest bed rests upon a deposit of

varved clay, and it is in this clay that the first traces of plant

and animal remains are found. Fragments of plants are found

in the varved clay to a considerable depth, but none of these

can be identified. Near the top of the varved clay deposit three

species of mollusks were found. One is identified by Baker as

Fossaria dalli (Baker) ; its habit is wet mud above water.

Because of its large size, he considers this mollusk a Pleistocene

race (in litt. to F. T. Thwaites, 1926). The other two species

are Pupilla mus corum (Linn.) and Succinea avaru Say., both

forest forms might have been common at the edge of a flood

plain. These mollusks seem to date the period of the forest bed

with the earliest exposure of land above the glacial lake that

preceded it. There probably appeared at this time a few

grasses and mosses, for the peat fossils suggest such a develop-

40 Wisconsin Academy of Sciences , Arts , and Letters .

ment. There was probably a brief period when only grasses

and mosses covered the clay, but a spruce forest very soon

became established, for almost directly on the surface of the

clay there occur spruce cones, needles, and forest mosses. Mixed

with these mosses are shells of land mollusks Succinea avara

and Vertigo ventricosa (Morse) One moss is peculiarly

restricted to the lowest level of the forest bed. This is Bryum

cyclophyllum (Schwaegr.) B. and S., a forest form, and it

seems to have been the first moss to have become established

on the Two Creeks Forest floor. Other plants that appear in

this horizon are grasses, heaths, birch, jack pine (Finns Bank -

siana Lamb.) and a species of Asplenium . These are repre¬

sented only by pollen grains and spores. The rare occurrence

of the pollen grains of the birch and pine suggest that they

were probably blown into the forest bed from a short distance.

Fungi were abundant, for the spores are numerous in the peat.

Some of these appear to be of lichens ; the others are all repre¬

sentative of the dark spored Dematieae.

The spruce forest appears to have thriven for at least sixty-

two years before a gradual decrease was observed in the growth

rings of the trees. The decrease in growth began roughly

twenty years before the forest was destroyed by the advancing

ice and water.

Little weathering of the subsoil was accomplished, for it is

all calcareous.

Bark beetle excavations have been found, which may belong

to the period of forest deterioration. Two genera are repre¬

sented, as shown by the patterns of the excavations.

The advance of the Red ice upon the Two Creeks Forest must

have been gradual. Water flowed into the forest floor, for there

is a marked change in the flora and fauna, that would suggest

such a condition. If the growth rings of the black spruce are

reliable indicators of growing conditions, they show conditions

unfavorable to secondary thickening in the Two Creeks Forest

during the last twenty years of the interval. An examination

of the uppermost plant and animal remains shows them to be

entirely wet land and aquatic species. This suggests excess

moisture as a reason for the narrow growth rings of the spruce

and the beginning of a third period in the ecological history of

the interglacial forest.

Wilson — Two Creeks Forest Bed .

41

Apparently the only plants that throve during the last years

of this interval were the following aquatic and subaquatic

mosses: Bryum bimum Schreb., B. pseudo-triquetrum (Hedw.)

Schwaeg., Calliergon cordifolium Kindb., C. stmmineum

(Dicks.) Kindb., C. turgescens (Jens.) Kindb., Camptothecium

nitens Schp., Campylium stellatum (Schreb.) Bryhn., Dicranella

sp., Ditrichum flexicaule Hampe, close to var. brevifolium

Kindb., Drepanocladus aduncus Moenkem., var. typicum

(Hedw.) Ren., D. aduncus Moenkem., var. pseudo fluitans Sanio.,

D. revolvens (Sw.) Warnst., D. Sendtneri (Schrp.) Warnst., D.

vernicosus (Limb.) Warnst., D . Wilsoni (Schpr.) Roth., Scor -

pidium scorpiodes (L.) Limpr., Tortella fragilis (Drumm.)

Limpr.

Mollusks found at the same level as the above named mosses

were the following aquatic species: Gyraulus circumstriatus

(Tryon), Fossaria parva (Lea.), Pisidium sp.

The mosses show several interesting ecological conditions and

furnish also some evidence of the manner in which the forest

was destroyed. The uppermost mosses are upright in position

and have finely divided sediment infiltrated about their stems.

A similar observation was made by Cooper (1923) at Glacier

Bay, Alaska. From this fact it seems evident that the forest

was flooded with silt bearing water before being overridden by

the Red ice. The mosses also show by thin scraggly branches, in

the last year of growth, that a losing struggle for existence

was under way just before they were buried. Mr. Cheney has

pointed out to the writer that some of the leaves of the upper

branches contain the remnants of chloroplasts. That chi or op-

lasts should be preserved is remarkable, and it is strong evi¬

dence that the mosses were living at the time of burial.

Three or four inches of sediment cover the water mosses and

mollusks. In the sediment one noteworthy moss was recorded,

namely Swartzia montana (Lambk.) Lindb., which grows in an

alpine or rocky habitat. As conditions of this type did not occur

in the forest bed, it is most probable that this moss was trans¬

ported from a more northern habitat with the sediment.

The glacial till above the sediment contains fragments of

wood and logs, many of which were flattened by the weight of

the ice. It was from the Red till that the largest log was taken,

and as already noted there is some reason to conclude that this

42 Wisconsin Academy of Sciences, Arts, and Letters .

log was transported by the glacier from a more northern and

upland forest a possibility which would harmonize with the

occurrence of Swartzia montana.

Origin of the Two Creeks Forest Bed Life

The study of many Pleistocene organic deposits has revealed

new varieties and forms of both plants and animals (Williams,

1930), (Baker, 1930), but from the Two Creeks Forest Bed no

new forms with the exception of one Pleistocene molluscan

race, has been recognized. This fact may lead us to conclude

that the Two Creeks Forest was little different from present

forests or habitats in which the recognized species now live,

and the forest did not, of course, exist long enough with pecul¬

iar conditions to produce new or characteristic forms. The ab¬

sence of particular Pleistocene forms in the forest bed proper al¬

so suggests that the Two Creeks Forest flora and fauna are rep¬

resentative of a general distribution, rather than of an isolated

region where local forms and races had been evolved. The

forest was evidently in the path of plant and animal distribu¬

tion, but sources from which the plants and animals arrived

can only be suggested at present.

The top thirty-five inches of the clay underlying the forest

bed contains miscellaneous tissues. The occurrence of vegetal

tissues in the upper varved clays raises a question as to their

source, but the writer has not attempted a solution of this geo¬

logical problem with the present data. It would appear, how¬

ever, that Glacial Lake Chicago existed long enough to allow

plants to establish themselves on nearby land, and that these

were then washed into the lake. The plant tissues in the varved

clays are too fragmentary to be identified, but may be from the

ancestors of the earliest flora of the forest bed. This may

explain the origin of some of the Two Creeks Forest Bed

plants, but it appears that there must also have been a coloni¬

zation from the south or west, as indicated by the pollens of

jack pine and birch, fern spores, and the great quantity of

mollusks. It does not seem probable that such plants and ani¬

mals could have long survived glacial conditions.

The plants that were not closely associated with the glacier

front may have entered the region of the forest bed either from

the southeastern or southwestern part of Wisconsin. This is

Wilson— Two Creeks Forest Bed.

43

suggested by the many records of interglacial materials in

those parts of the state. Several well records west of Baraboo

and North Freedom, Sauk County (Alden, 1918. p. 226) show

successive layers of vegetable remains separated by aqueous

deposits presumably of glacial derivation, and are very sug¬

gestive as evidence of the persistence of plants in the Driftless

Area of Wisconsin during the Glacial Period. At present only

one organic deposit from Baraboo has been examined. The

material was secured by F. T. Thwaites and submitted to the

writer in March, 1931. It was found under 130 feet of glacial

deposits, and according to Mr. Thwaites the deposit may be

of Sangamon age. Leaves of several dicotyledonous plants and

one species of moss were separated from the sandy material.

The leaves are too fragmentary to be identified, except a num¬

ber which belong to a species of Vaccinium resembling most

nearly V. microcarpon J. D. Hook., a species of northern

Europe. The resemblance of the Baraboo specimens is not close

enough to consider them as belonging to this species, and it is

possible that they represent an extinct species of Vaccinium.

The moss was identified by Mr. Cheney as Campylium stellatum

(Schreb.) Bryhn. This deposit will be considered in a later

paper.

Baker (1920, pp. 71-73) has described deposits at the south¬

ern end of Lake Michigan which appear to be contemporaneous

with the Two Creeks Forest Bed, but further correlation of

materials has not yet been possible.

The plants and animals may have spread either northward

into Manitowoc County along the shore of Lake Michigan, or

eastward from the Driftless Area and other, at that time,

unglaciated areas that existed outside the limits of the Gray

ice. There are many records of other forest beds south and

west of Two Creeks (Alden, 1918. p. 179) that probably belong

to the same interval, but none of these have been studied. The

forest beds of the lower Fox River valley evidently were exten¬

sively exposed when Lawson (1902) did his work there, but

his records of the plant remains cannot be considered accurate,

for as he notes, they were never critically examined. It is doubt¬

ful whether logs of pine, cedar, black ash, and tamarack, which

he records, could be definitely identified in the field, for only

after a microsopic examination of the material is it possible

to identify interglacial wood with certainty, and even then to

44 Wisconsin Academy of Sciences, Arts, and Letters .

distinguish between closely related species is impossible. The

exposures which Lawson visited have nearly all been destroyed,

and it has not been possible to secure material from them except

one spruce log (Picea sp.) from Menasha, Winnebago county.

According to Lawson the forest beds of the Fox River valley

cover about five hundred square miles and extend into four

counties. They probably belong to the same interval as the Two

Creeks Forest Bed, and were connected with it. It is evident

that the Two Creeks Interval was of long enough duration to

allow of the establishment of extensive forests over the eastern

part of Wisconsin.

Probable Climate of the Two Creeks Interval

It is impossible to give any definite statement regarding the

climate which prevailed during the Two Creeks Interval because

there are two conflicting lines of evidence. These are (a) the

present known ranges of the plants, and (b) the present known

ranges of the mollusks. Nevertheless, an average range has

been computed and taken as an indicator of the probable cli¬

matic conditions.

To consider first the ranges of the recognized species of

plants, it has been seen that these species are now largely con¬

fined to the northern part of North America and northern

Eurasia. The greatest abundance of individuals of the species

in question seems from a study of literature (Atlas of Canada,

1915; Fernald, 1919; Herzog, 1926) and from the writer's

observations, to be in that region somewhat north of Minnesota

in Ontario. In that region a flora of a northern type is dom¬

inant, and there all the plant species recognized in the forest

bed occur, a statement which, so far as their present ranges

are known, does not hold for any more southern location.

The mollusks show more southern distributions, which accord¬

ing to Mr. F. C. Baker {in litt. 1931) appear to fit into the cli¬

matic conditions of northern Wisconsin. The northern distri¬

butions of these mollusks are not as well known as are those

of the plants, for the reason that they are all small forms and

are not as often collected by naturalists.

If an intermediate range is chosen between the general plant

range and the general molluscan range, then the climatic con¬

ditions of northern Minnesota may be taken as representing

Wilson — Two Creeks Forest Bed .

45

approximately the climate of the Two Creeks Interval at the

time of the climax forest, which was composed of spruce and

other woodland species.

Summary

1. There was an interglacial interval of considerable duration

between the Red (Late) and Gray (Middle) Substages of the

Wisconsin Stage of glaciation. This is shown by the position

of the Two Creeks Forest Bed between tills of characteristic

lithological nature.

2. For the Two Creeks Forest Bed to have been established,

the Gray ice must have receded north far enough to clear the

Straits of Mackinac thus furnishing an outlet to Lake Chicago

and allowing its water level to drop to a level near to, or lower

than the present.

8. The interval was long enough in duration to allow the

growth of a forest whose age was at least eighty-two years.

4. Both plant and animal remains are well preserved and

can be definitely identified as present day forms, with the excep¬

tion of one mollusk (Fossaria dalli), which belongs to a pecul¬

iar Pleistocene race differing in size from existing forms.

5. The plants and animals represented in the Two Creeks

Forest Bed are today arctic, subarctic, and northern in distri¬

bution.

6. There were three periods in the history of the forest bed,

characterized: first, by aquatic and semi-aquatic mollusks on

top of the varved clay of early Lake Chicago ; second, by moist

to dry woodland mosses, mollusks, mites, fungi, and trees mark¬

ing an interval of dry land ; and third, by water mosses, water

mollusks, and sediment caused by the advance of the Red ice

blocking the drainage.

7. The climate of the Two Creeks Interval was probably like

that of northern Minnesota, or somewhat colder, as shown by

the prevailing northern ranges of the identified plants and ani¬

mals. The complete deglaciation of Canada is not suggested

by the evidence at Two Creeks, Wisconsin.

Specimens of wood, peat, and mollusks are to be found in

the Geology Museum of the University of Wisconsin, while the

46 Wisconsin Academy of Sciences , Arts , and Letters .

mosses and bracket fungus from the Two Creeks Forest Bed

are in the collections of the Herbarium of the same university.

The author wishes to express his appreciation to Dr. N. C.

Fassett for his encouragement and suggestions, to Mr. F. T.

Thwaites for his geological advice and interest that have made

this paper possible, to Mr. L. S. Cheney and Mr. F. C. Baker

for identification of specimens, and to Dr. C. E. Allen for his

helpful criticism.

References

Alden, W. C. 1918. Quarternary Geology of southeastern Wisconsin. U. S.

Geol. Survey Prof. Paper 106.

Atlas of Canada 1915. Dept, of Interior, pp. 19-20.

Baker, F. C. 1920. Life of the Pleistocene or Glacial Period. III. Univ.

Bull, Vol. 17, No. 41.

1930. Influence of the Glacial Period in changing the character

of the molluscan fauna of North America. Ecology 11 : 469-480.

Cheney, L. S. 1930. Wisconsin fossil mosses. Bryologist 33 : 66-68.

Cooper, W. S. 1923. The interglacial forests of Glacial Bay, Alaska.

Ecology 4 : 93-128.

Ertman, G. 1931. Pollen-statistics: A new research method in paleo-

ecology. Science N. S. 73 : 399-401.

Fernald, L. M. 1919. Lithological factors limiting the ranges of Pinus

Banksiana and Thuja occidentalis. Rhodora 21 : 42-67.

Goldthwait, J. W. 1907. The abandoned shore-lines of eastern Wisconsin.

Wis. Geol. Nat. His. Surv. Bull. 17 : 61-62.

Herzog, Th. 1926. Geographic der Moose.

Lawson, P. V. 1902. Preliminary reports of the forest beds of the lower

Fox, Wis. Nat. Hist. Soc. Bull. 2 : 170-173.

Leverett, Frank 1929. Morains and shore lines of the Lake Superior

basin. U. S. Geol. Surv. Prof. Paper. 154 A. p. 19.

Thwaites, F. T. 1927. The development of the theory of multiple glacia¬

tion in North America. Trans. Wis. Acad. 22 : 40-164.

Williams, R. S. 1930. Pleistocene mosses from Minneapolis, Minnesota.

Bryologist 33 : 33.

Zon, R. and Averell, L. J. 1929. Drainage of swamps and forest growth.

Univ. Wis. Agric. Exp. Sta. Res. Bull. 89.

VARVED CLAYS OF WISCONSIN

Elmer W. Ellsworth

Introduction

During 1928-1929 the writer, in conjunction with W. L.

Wilgus, made a thorough study of the Waupaca, Wisconsin,

varved clay deposit, and presented the results of this work in

a printed report.1 The present study is an extension of this

work and covers all of the known varved clay deposits in the

state. The writer visited all of the exposures and procured

samples for chemical and petrographic analysis. The clays

studied represent the basin deposits of at least four known

glacial lakes : Glacial Lake Chicago, Glacial Lake Oshkosh, Gla¬

cial Lake Wisconsin, and the glacial lake of northwestern Wis¬

consin.

This study has been made with a definite purpose in view:

(1) To determine whether or not samples of varved clay taken

from different exposures of the same lake bed possess similar

chemical and petrographic properties; (2) To determine to

what extent the chemical and mineralogical properties of the

varved clays of the different glacial lake beds vary throughout

the state; and (3) To determine the value of such analyses in

correlating scattered exposures as belonging to one or another

former glacial lake basin.

The complete results of the many analyses of the clay sam¬

ples are included in this report, as well as the conclusions

deduced from their interpretation.

The writer wishes to thank Mr. H. R. Aldrich and Mr. J. M.

Hansell of the Wisconsin Geological Survey for their coopera¬

tion in the work of examining many of the deposits in the field.

The work was done in the Sedimentation Laboratory of the

University of Wisconsin under the direction of Professor W.

H. Twenhofel.

Location of the Varved Clay Deposits

The geographic areas under consideration are shown upon

the accompanying map of the state, Fig. 1. This map also indi-

1 Ellsworth, E. W., and Wilgus, W. L. : The Varved Clay Deposit at Waupaca,

Wisconsin, Trans. Wis. Academy of Science, Arts and Letters, Vol. XXV.

48 Wisconsin Academy of Sciences, Arts, and Letters.

NORTHWEST AREA

OCONTO

WISCONSIN

OSHKOSH

cates the probable position and extent of the several glacial

lakes at their maximum extent.

Two Rivers. Along the shore of Lake Michigan, a short dis¬

tance north of the town of Two Rivers, is an exposure of

Pleistocene deposits comprising glacial till, a buried forest bed,

and varved clay. The varves of the latter are poorly defined,

and are deformed by folding. Representative samples were col¬

lected here during the spring of 1929.

Fig. 1. Key Map. Areas outlined in black indicate the probable position

and boundaries (within the state) of the several glacial lakes at their

maximum extent. The names on the map are the names of these respective

glacial lakes ; the numbers denote the locations of the varved clay ex¬

posures by geographical units : (1) Two Rivers ; (2) Manitowoc; (3) New

London ; (4) Waupaca; (5) Friendship; (6) Northwest Area.

CHICAGO

Ellsworth — Varved Clays of Wisconsin.

49

Manitowoc. Along the southern banks of the Manitowoc

River at Manitowoc, varved clay is exposed in the clay pit

operated in conjunction with the Leach Company brick yard.

The exposed face is fifteen to twenty-five feet high, and the

varves are fairly well developed. For the most part they are

in a normal horizontal position, though slight folding of some

of the beds is noticeable. A large specimen, containing a dozen

varves, was brought to Madison during the spring of 1929.

New London. The deposit here has been opened by a local

brick concern, and a ten to fifteen foot exposure created. The

varves are fairly well defined and appeared to be in a per¬

fectly normal position. Limy concretions, or “clay dogs”, were

found in the varves at several places. Samples of clay repre¬

sentative of the deposit were taken for analysis.

Friendship. The varved clays here are exposed along the

banks of a small stream, and the exposure visited in the fall of

1929 was four to six feet in height. The varves are well defined,

but have an average thickness of less than one inch. A small

section was cut from the face of the exposure and taken to

Madison for examination.

Northwest area. This area includes parts of Burnett and

Polk Counties, and contains numerous exposures of varved clay.

Those along the Trade, Wood, Clam and St. Croix rivers were

examined in the field during the summer of 1929, and repre¬

sentative samples taken from each exposure. The varves are

excellently developed in all exposures visited, and are very

similar to those studied at Waupaca during the previous year.

At three locations the complete sections exposed were measured.

From these measurements, which included over five hundred

varves, curves showing the variation in varve thickness were

constructed, to be later used in the correlation of these deposits.

Samples Analyzed in this Study

In general, at least three complete varves from each exposure

were sampled for analysis. Care was taken to select representa¬

tive varves and to include one from the bottom, middle and top

of each exposure. In the laboratory the samples were divided

into summer and winter components. The resulting series of

samples numbered sixty-five, and the list of these samples and

their field locations are presented in Table I.

50 Wisconsin Academy of Sciences , Arts, and Letters.

Table I

*Where the winter and summer components are from the same varve, the letters W and S are

followed by a common letter, as Wa and Sa.

fOver one hundred nearly complete carbonate analyses were accidentally destroyed. Samples

were replaced wherever possible, and the analyses repeated. The missing percentages represent

samples which could not be replaced.

Ellsworth — Varved Clays of Wisconsin.

51

Chemical Analysis of the Clay Samples

In Table I are presented the results of chemical analyses of

the clay samples. The Fe203 content was determined by the

permanganate method of titration, and it was found necessary

to filter the sample in order to secure a proper end-point. The

percentage of C02 was determined and this was divided by .459

to secure the approximate total percentage of the common car¬

bonate minerals (assuming the samples contained 50% calcite

and 50% dolomite, their total percentage is derived from the

percentage of C02 by dividing the latter figure by .459). The

silicates and insoluble were determined by difference. Duplicate

samples were analyzed, and the percentages given below express

the average percentages. The average difference between dup¬

licate samples was .3%

From the analyses given in Table I it will be noted that:

(1) In every case the Fe203 content of the winter compo¬

nents is higher than that of the summer components of samples

from the same exposure.

(2) In every case the silicate content of the winter compo¬

nents is lower than most of the summer components of samples

from the Northwest, New London, and Friendship areas.

(3) The silicate content of the varve components from the

other geographic areas appears to be about the same.

(4) The carbonate content of the winter components is

higher than that of the summer components of samples from

the Northwest, Friendship and New London (one slight excep¬

tion) areas.

(5) The carbonate content of the winter components is lower

than that of the summer components of samples from the Mani¬

towoc exposure.

(6) The carbonate content of the varve components from

the Two Rivers exposure is about the same.

It will also be noted that :

(1) The percentage of Fe203 in the samples of varved clay

from the Northwest area is nearly double the percentage of

Fe203 in samples of clay from each of the other areas. Samples

of varves from the Northwest area have an average Fe203 per¬

centage of 9%, whereas this figure for samples from each of

the other areas is either 5 or 6%.

52 Wisconsin Academy of Sciences , Arts , and Letters .

(2) The percentage of carbonate in the samples of varved

clay from the Northwest area is approximately one-third the

percentage of carbonates in samples of clay from each of the

other areas. Sample of varves from the Northwest area have

an average carbonate percentage of 12%, whereas this figure

for samples from each of the other areas varies from 29 to

39%, averaging 33% total carbonates.

Summarizing the above, and interpreting the results in terms

of geographic units, it appears that :

(1) Varves of the Northwest area are characterized by:

(a) A silicate content of the winter components which is

lower than that of the summer components.

(b) A carbonate content of the winter components which

is higher than that of the summer components.

(c) A percentage of Fe203 which is nearly double that of

varves from each of the other areas. Percentage = 9%.

(d) A percentage of carbonates which is approximately

one-third that of varves from each of the other areas. Per¬

centage = 12%.

(e) An Fe203 content of the winter components which is

higher than that of the summer components.

(2) Varves of the New London areas are characterized by:

(a) A silicate content of the winter components which is

lower than that of the summer components.

(b) A carbonate content of the winter components which

is higher than that of the summer components.

(c) A percentage of Fe203 which is one half that of

varves of the Northwest area. Percentage = 5%.

(d) A percentage of carbonates which is approximately

three times that of varves of the Northwest area. Percentage

= 29%.

(e) An Fe203 content of the winter components which is

higher than that of the summer components.

(3) Varves of the Manitowoc area are characterized by:

(a) A silicate content of the varve components which is

approximately the same.

(b) A carbonate content of the winter components which

is lower than that of the summer components.

Ellsworth — Varved Clays of Wisconsin .

53

(c) A percentage of Fe203 which is about one half that

of varves of the Northwest area. Percentage = 6%.

(d) A percentage of carbonates which is approximately

three times that of varves of the Northwest area. Percentage

= 37%.

(e) An Fe203 content of the winter components which is

higher than that of the summer components.

(4) Varves of the Two Rivers area are characterized by:

(a) A silicate content of the varve components which is

approximately the same.

(b) A carbonate content of the winter components which

is higher than that of the summer components.

(c) A percentage of Fe203 which is approximately one-

half that of varves of the Northwest area. Percentage = 5%.

(d) A percentage of carbonates which is approximately

three times that of varves of the Northwest area. Percent¬

age — 39%.

(e) An Fe203 content of the winter components which is

higher than that of the summer components.

(5) Varves of the Friendship area are characterized by:

(a) A silicate content of the winter components which is

lower than that of the summer components.

(b) A carbonate content of the winter components which

is higher than that of the summer components.

(c) A percentage of Fe203 which is one-half that of the

varves of the Northwest area. Percentage = 5%.

(d) A percentage of carbonates which is more than twice

that of varves of the Northwest area. Percentage = 29%.

(e) An Fe203 content of the winter components which is

higher than that of the summer components.

(6) Varves of the Waupaca area2 are characterized by:

(a) A silicate content of the winter components which is

lower than that of the summer components.

(b) A carbonate content of the winter components which

is higher than that of the summer components.

(c) A percentage of Fe203 which is one half that of varves

of the Northwest area. Percentage = 4%.

2 Taken from B. A. Thesis, 1929. University of Wisconsin.

54 Wisconsin Academy of Sciences, Arts, and Letters,

Fig. 2. Results of the petrographic analysis.

Ellsworth — Varved Clays of Wisconsin.

55

(d) A percentage of carbonates which is more than double

that of varves of the Northwest area. Percentage = 27%.

(e) An Fe203 content of the winter components which is

higher than that of the summer components.

From the above data it is clear that the varved clays from

each geographic area possess definite chemical characteristics.

The varves of the New London, Waupaca, and Friendship areas

are very similar with respect to their chemical composition.

The varves of the Northwest, Manitowoc, and Two Rivers areas

are not only distinctly different from those of the other areas,

but each of these possesses varves which are different in chemi¬

cal composition from those of each of the other areas.

Petrographic Analysis op the Clay Samples

The same series3 of samples which was analyzed chemically

was made the subject of a petrographic analysis.

The samples were very thoroughly washed and rewashed (by

decantation) in an ammoniacal solution, and the heavy mineral

grains separated from the light grains by use of tetrabromo-

ethane (specific gravity 2.9). The heavy minerals were then

mounted on glass slides, with the use of Canada balsam.

A table presenting the results of the petrographic analysis

of the heavy minerals is presented in Fig. 2. A study of this

Table II. The average percentages of the common minerals found in

the clays of each area.

* See Ellsworth, E. W. and Wilgus, W. L. The Varved Clay Deposit at

Waupaca, Wisconsin. Trans. Wis. Academy. Vol. XXV. 1929.

3 The winter components were not analyzed for their mineral content.

56 Wisconsin Academy of Sciences, Arts, and Letters,

table reveals that there are wide differences in the analyses of

samples from the same geographic areas, and even from the

same field exposure. It is very apparent that if the clays of

a single area possess common mineral characteristics, these

will be revealed only through the average of many petrographic

analyses. Even then the individual differences appear to be so

great, in some instances, that the value of such averages is very

doubtful.

Table II presents the average mineral composition of clays

of each of the areas studied. Only the important (abundant)

minerals are included. A careful comparison of these analyses

brings to light a grouping of the clays of these areas into two

divisions: (1) The clays of the Northwest, Two Rivers, and

New London areas, and (2) the clays of the Manitowoc, Wau¬

paca, and Friendship areas. The average mineral composition

of the clays of each of the divisions is presented in Table III.

Table III. The average mineral composition of the clays of (1) the

Northwest , Two Rivers and London areas; and (2) the Manitowoc, Wau¬

paca and Friendship areas.

While the table shows that clays of each of these divisions

are quite different, the writer is inclined to attach no great

importance to this grouping of the clays. If the common source

material of the clays is indicated by their common petrographic

characteristics, it is difficult to see how the clays of the North¬

west, New London, and Two Rivers areas, could be derived

from the same terranes, and different from those which were

the source terranes for the clays of an overlapping area (divi¬

sion 2).

Ellsworth — Varved Clays of Wisconsin. 57

Correlation of the Varved Clay Deposits

At three localities, Benson, Clam River and Wood River, the

complete sections exposed were carefully measured in the field.

Over five hundred varves were measured, and curves showing

the variations in varve thickness were constructed from the

field measurements. - Copies of these curves were sent to Baron

Gerard De Geer, at Stockholm, for correlation with the solar

curve. Regarding the correlation which he made, De Geer

states4 :

4*_„__„_a close comparison has made it possible to identify

almost all of the varves on your diagrams. The similarity is so

conclusive that your measurements will afford a valuable con¬

tribution to the material which I am collecting for a universal

solar curve.”

By a detailed comparison you will find that the simi¬

larity of the curves does not leave any doubt as to their identi¬

fication.”

This correlation dates the varves of the Northwest area as

having been deposited several hundred years after those at

Waupaca and Manitowoc.

Summary and Conclusions

(1) This study has shown that samples of varved clay taken

from different exposures of the same glacial lake bed in Wis¬

consin present similar chemical characteristics but their petro¬

graphic analyses show a wide range in variation.

(2) The chemical properties of the varved clays of the Lon¬

don, Waupaca, and Friendship areas have been shown to be

similar, while those of the Northwest, Manitowoc, and Two

Rivers areas are different from the three just mentioned. Thus

the deposits of glacial lakes Oshkosh and Wisconsin are shown

to be similar chemically, and were probably derived from the

same source materials. The deposits in Glacial Lake Chicago

and the glacial lake of Northwestern Wisconsin are different

chemically, and each in turn is quite different from those of

glacial lakes Oshkosh and Wisconsin.

(3) This study would tend to indicate that the chemical

analysis of varved clays of scattered exposures is of value in

4 Letter of May 3, 1930.

58 Wisconsin Academy of Sciences , Arts, and Letters.

correlating them with one or another glacial lake basin in Wis¬

consin. The value of a petrographic analysis in this connection

is very doubtful, due to the wide range in the variations of the

mineral content of varves of even the same exposure.

(4) De Geer’s correlation of the varves of the Northwest

area assigns to them an age which is several hundred years

younger than those of Glacial Lake Oshkosh exposed at Wau¬

paca.

THE DISTRIBUTION OF CLOUDINESS IN WISCONSIN

Eric R, Miller

The six charts of this paper show by isonephs (lines of equal

cloudiness) the bimonthly average percentage of sky covered

with clouds during the ten years, 1920 to 1929 inclusive.

The most obvious feature of these maps is a region of maxi¬

mum cloudiness in northwestern and north central Wisconsin,

apparently derived from the western end of Lake Superior, or

associated with the western front of the northern highlands.

Another area of maximum cloudiness hangs over Lake Michi¬

gan. A belt of less cloudiness follows the valleys of the Missis¬

sippi, lower Wisconsin, Fox and its extension in Green Bay

from September to April. From May to August this is oblit¬

erated by the union of the two regions of maximum cloudiness.

The annual march of cloudiness shows a single period, with a

maximum in November and a minimum in August, differing

from the rainfall periodicity with two maxima in Spring and

Autumn.

The data from which these charts were prepared are the

results of personal, not instrumental, observation. Some inter¬

esting statistical and psychological problems arose in working

them up for charting.

“Instructions for cooperative Observers, Circulars B and C,

Instrument Division, U.S. Weather Bureau, 7th ed.” contains

the following directions : “Par. 67. Character of the day. — The

general character of the day from sunrise to sunset should be

recorded as “clear” when the sky averages three-tenths or less

obscured ; partly cloudy, when from four-tenths to seven-tenths

obscured; and cloudy when more than seven-tenths obscured.

The average cloudiness from sunrise to sunset may be esti¬

mated with considerable accuracy by noting the degree of

cloudiness on the scale given, as near sunrise as possible,

between noon and 1 p. m. and near sunset ; add these and divide

by 3; the quotient will be the average cloudiness.” The fre¬

quency of clear, partly cloudy and cloudy days is the sole form

used in printed tables for the cooperative stations. To reduce

these three figures to one for use in mapping the average cloud-

60 Wisconsin Academy of Sciences , Arts , and Letters .

iness at Madison on the clear, partly cloudy and cloudy days

was taken out (Table I). From these data conversion tables

were prepared for each month. As a check on the use of Madi¬

son data throughout the state, the conversion tables were applied

Table I. Average cloudiness at Madison , Wisconsin on clear , partly

cloudy and cloudy days.

Fig. 1. Cloudiness, J anuary-February, per cent of sky covered with

clouds, 10-year average, 1920-1929.

Miller — Cloudiness in Wisconsin.

61

Fig. 2. Cloudiness, March-April, per cent of sky covered with clouds,

10-year average, 1920-1929.

to data on “Character of Day” from the regular Weather

Bureau offices, where the average cloudiness is also recorded.

The results of these comparisons appear in Table II. The aver¬

age difference for the whole 60 pairs is -.07 or a little more

than one per cent.

The records from the observing stations were first plotted

individually, but were so inconsistent that it was necessary to

smooth by averaging them five at a time and plotting the results

at median points.

The results also averaged lower than the data from the regu¬

lar observing stations of the Weather Bureau. The regular

62 Wisconsin Academy of Sciences , Arts , anc? Letters ,

Table II. Comparison of monthly cloudiness , estimated from number of

clear , partly cloudy and cloudy days , with observed data.

Table III. Cloudiness at regular and cooperative weather stations.

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

Duluth .

12 coop, obsrs. . . .

Difference .

Cornucopia, Wis. .

Danbury, Wis. . . .

Wausau .

20 coop, obsrs. . . .

Difference .

Prentice .

Florence .

Green Bay .

11 coop, obsrs. ...

Difference .

Marinette .

Menasha .

Milwaukee .

5 coop, obsrs .

Difference .

Plymouth .

Fond du Lac .

Madison .

1 1 coop, obsrs. . . .

Difference . . .

Prairie du Sac . . .

Brodhead .

La Crosse .

11 coop, obsrs. ...

Difference .

Meadow Valley . . .

Hillsboro .

Minneapolis .

St. Paul .

11 coop, obsrs. ...

Downing .

Mora, Minn .

Miller — Cloudiness in Wisconsin,

63

Fig. 3. Cloudiness, May-June, per cent of sky covered with clouds, 10-

year average, 1920-1929.

observers are required to take account of all kinds of clouds in

estimating cloudiness, so that a day with the sky covered with

cirro-stratus cloud would be recorded cloudy although the sun

were shining brightly all day, while the cooperative observer

would doubtless record it clear. In order to show the character¬

istic difference between regular and cooperative observers in

recording cloudiness, the data from each of the regular weather

observing stations in or near the borders of Wisconsin is tabu¬

lated in Table III, with the average cloudiness at the coopera¬

tive stations in the vicinity, and records from one cooperative

64 Wisconsin Academy of Sciences , Arts , and Letters .

Fig. 4. Cloudiness, July-August, per cent of sky covered with clouds,

10-year average, 1920-1929.

observer with a high average and another with a low average.

These data are for the same period, 1920 to 1929 inclusive.

The larger difference between Madison and the surrounding

cooperative observers may be due to the pyrheliometric observa¬

tions at Madison. These make the observer conscious of every

faint cloud. Such observations are not made at the other regu¬

lar stations. The less amount of cloud at St., Paul than at Min¬

neapolis may be due to more smoke, since St. Paul is the east¬

erly of the Twin Cities, and smoke is not included in the count

of tenths of sky obscured by cloud.

Miller — Cloudiness in Wisconsin.

65

Fig. 5. Cloudiness, September-October, per cent of sky covered with

clouds, 10-year average, 1920-1929.

All of the regular Weather Bureau stations, except Duluth,

in and around Wisconsin are supplied with the Marvin ther¬

mometric sunshine recorder. This instrument nominally records

the duration of bright sunshine. Actually its sensitiveness

varies with the intensity of sunshine. One difficulty that has

not been solved is that it requires readjustment to the intensity

of sunlight as the latter varies from season to season. No quan¬

titative method of doing this has been devised, hence the records

of the instrument are not comparable one station with another,

nor one time with another.

66 Wisconsin Academy of Sciences, Arts, and Letters .

Fig. 6. Cloudiness, November-December, per cent of sky covered with

clouds, 10-year average, 1920-1929.

In order to measure the relation of the records from the sun¬

shine recorder to the cloudiness during daylight hours the coeffi¬

cient of correlation between the cloudiness and the percentage

of duration of sunshine in each month for 26 years at Madison,

and for January and July at a number of other stations, has

been calculated and set down in Table IV.

In the case of Charles City, the sunshine is recorded as vary¬

ing with the cloudiness, in July, in 9 of the first 13 years of

the record. Since then the two elements have varied in the

opposite sense 11 years in 13, so that the coefficient would be

much larger for the last 13 years.

Miller — Cloudiness in Wisconsin . 67

i a n & a &

9E LA & J £*

Table IV. Coefficient of correlation between monthly cloudiness and

percentage of sunshine. (26 years.)

The diurnal march of cloudiness at Madison is shown in Table

V.

The midday maximum in the warmer months would be accen¬

tuated if the convectional types of cloud only were included in

Table V. Average cloudiness from bihourly observations, Madison,

Wis. 1920-1929.

the count. For comparison the average cloudiness at 7 a. m.,

noon and 7 p. m. at a number of cities is given in Table VI.

Observations are made at Minneapolis at noon and night

only, at St. Paul morning and noon only, and as we have seen

that there is a systematic difference it will be interesting to

compare the data for the hour when records are made at both

places (see bottom of Table VI.)

The average percentage of possible sunshine for each hour,

at Madison for the 20 years 1911-1930, in Table VII, when

compared with the preceding tables, shows the essentially dif-

68 Wisconsin Academy of Sciences , Arts , and Letters .

Table VI. Average cloudiness at 7 am, noon and 7 pm, 1920-1929 inc.

ferent character of the records from the thermometric sun¬

shine recorder. The high percentages in the middle of the day

in summer directly contradict the observations of cloudiness.

The November maximum of cloudiness does not appear.

In closing, it may be remarked that the chief cloud-making

process, the cyclonic storm, has been eliminated by the method

Table VII. Percentage of possible sunshine at Madison, Wis., 1911-1930,

in the different hours of the day.

Miller — Cloudiness in Wisconsin .

69

of tabulating the data. The maps of average distribution show

the influence of the Great Lakes as nearby sources of moist air,

the tables bring out the annual and daily cycles. It has been

necessary to devote much space to the discussion of methods

of observation because the observations are personal estimates,

which could be better standardized only at an impossible

expense.

THE DEVELOPMENT OF THE ICE CREAM FREEZER

H. A. SCHUETTE AND FRANCIS J. ROBINSON

Contribution from the Laboratory of Foods and Sanitation,

Department of Chemistry, University of Wisconsin

The ice cream industry was one of the laggards in the

mechanization process of the industrial revolution for reasons

which are probably twofold. On the one hand it was slow to

attain a full measure of growth — the first wholesale ice cream

manufacturing plant was established by Jacob Fussel as late

as 1851 — and on the other the vicissitudes in the demand for

the product had made an extensive early localization in fac¬

tories economically infeasible. Not until the latter part of the

past century did ice cream-making, firmly intrenched as it was,

leave the small shop of the confectioner and the kitchen of the

housewife for the factory of commerce. This move brought

notable results for ice cream became the product of a vast and

well regulated manufacturing industry, a standard article of

commerce within the reach of every one, a food as well as a

confection. Such changes cannot rightly be accounted for by

any single factor; yet without the development of the modern

freezer, which has enforced the advantages of large scale pro¬

duction, few of these changes would have occurred and ice

cream might still be the simple “cream ice”, the frozen cream

of past years. It should prove worth while, therefore, to look

into the history of ice cream freezers and thus gain an appreci¬

ation of an important phase in the development of the science

of ice cream making.

Parkinson (1), a “practical confectioner of Chestnut Street,

Philadelphia” in 1849 prepared a statement of what was prob¬

ably the typical equipment of American ice cream makers until

the middle of the nineteenth century. The utensils requisite in

that period for making this food, and descriptive comments

thereon, are reproduced in his own words, to wit :

“1. Pewter pots of various sizes suitable to the quantity of

mixture intended to be frozen. Tin or zinc will not answer

the purpose as it congeals the mixture too quickly without al-

72 Wisconsin Academy of Sciences, Arts, and Letters .

lowing it a sufficient time to become properly incorporated,

and forms it in lumps like hailstones.

2. Moulds.

3. Ice pails.

4. The spatula. This is an instrument somewhat resem¬

bling a gardener’s spade; it should be made of stout copper

and tinned, the blade being about four inches long by three in

width, round at the end, and having a socket to receive a

wooden handle; this is for scraping the ice cream, etc., from

the sides of the pot as it freezes and for mixing it.

5. Either a large mortar or pestle, or a strong box and

mallet for pounding the ice.

6. A spade wherewith to mix the ice and salt together,

fixing your pails, etc.

7. A tin case or box — for keeping the ices in form of fruits

after they are finished.”

It has been reported (2) that a Nancy Johnson, the wife of

a young naval officer, invented the ice cream freezer shortly

after Dolly Madison had “officially” introduced this confection

to Washington society. Be that as it may, however, letters

patent on a similar piece of apparatus which was a combination

of revolving freezer and beater were granted for the first time

in the United States in 1848 (3). The patentee, Young, before

proceeding to describe his invention says in part,

“Many devices have been resorted to for expeditiously

freezing ice cream, but all have been found to be defective.

The best now in use is that known as ‘Johnson’s’, which is,

like the ordinary freezer, with a revolving shaft inside it, on

which are two curved wings that move round and cause the

cream to revolve in the freezer and be thrown to the outside.

I find that the operation is greatly facilitated by causing the

freezer itself to move rapidly as well as the cream inside.”

Describing his invention he says,

“ — there is a beater, full of holes, for the purpose of mov¬

ing the cream inside, while by turning the freezer in the ice

the ice is brought into close contact with it, and the cream is

so put in motion as to bring all of it rapidly into contact with

the cold sides of the freezer, which cannot be done by stirring

alone, while, by the aid of the beater, the cream is lightened

and the air allowed to come between the particles as effectu¬

ally as by any other mode of stirring, and by their united

operation the cream is more perfectly and speedily frozen

and well beaten than by either of the modes now used.”

Schuette and Robinson— -The Ice Cream Freezer . 73

Two other patents on freezers were issued that same year.

One of these (U) was for a device for congealing the cream in

the annular space between two concentric cylinders when the

smaller one was filled with ice and salt and the larger one was

!. 6. Young

Patent No. 6501

May SO, 1848

A. H. Austin

Patent No. 5775

Sept. 19, 1848

surrounded by this refrigerant. The cream was agitated with

a specially constructed plunger which served also to scrape the

frozen product from the walls. The other (5) covered a contriv¬

ance which suggests in a sense the modern household freezer.

In describing his invention, the patentee explains,

“My invention consists in constructing a freezer so as to

revolve the dasher within it and also at the same time to turn

the freezer in an opposite direction within the ice box, and I

form the spindle in the dasher large and hollow for the pur¬

pose of containing ice.”

During the next year two more freezers were invented. The

first (6) one embodied a minor structural feature for making

more convenient the use of a central freezing core, but the sec¬

ond (7) one apparently did not suffer from a lack of novelty.

The inventor proposed to freeze the cream by forcing through

it a current of cold air for he says,

“The nature of my invention consists in causing a blast of

chilled air to permeate, be diffused through and disturb the

liquids and materials of which ice cream is made. I chill the

74 Wisconsin Academy of Sciences , Arts , and Letters .

blast by drawing it from the atmosphere into a receptacle

which is made to surround the sides and bottom of the vessel

containing the ice or refrigerating mass. Within this vessel

the can containing the liquids and materials of which the ice

cream is to be formed is placed, and the interval between the

two packed with ice or the freezing compound. The air may

be drawn off at a central opening in the bottom of the air

chamber. A section of elastic hose is fastened in any usual

way to the opening and similarly attached at its other end

to an ordinary double bellows, mounted on a suitable frame.

. . . To the nozzle of this bellows I append a tube which

passes down through the middle of the ice cream tub and sep¬

arates into four or more horizontal branches open at their ends

at the bottom of the same. . . The chilled air blast being

forced through the horizontal branch tubes, bubbles up

through out the whole body of the liquids and materials in¬

tended for ice cream and besides abstracting caloric from

them by its own immensely extended contact therewith, it

thoroughly disturbs them and brings every portion of the

same into continually repeated contact with the refrigerating

surfaces . . .”.

Coffeen, the inventor of this device, was perhaps ahead of

the times and public taste with his air bubbling freezer that

would turn out ice cream with a scandalously high over-run —

providing, of course, that the contrivance would function as

claimed. And granting that it did work, the product made with

it would probably not have enjoyed any large measure of popu¬

larity. Until almost the end of that century people generally

had looked upon ice cream as a luxury rather than as a readily

ingested food, as an out-of-the-ordinary dessert whose sweetness

should remain upon the palate as long as possible. The turn of

the century still found a popular preference for a “heavy” ice

cream as the following comment (8), which incidentally gives

us a hint of the then accepted and genteel mode of eating this

dessert, testifies:

“Ice cream made from cream containing but 16 or 20 per

cent of fat will lack body or character; when put into the

mouth it immediately vanishes which is disappointing to the

lover of good ice cream.”

France, already rich in ice cream lore, is also in a position

to claim the distinction of having granted early patents, for it

is recorded that such were issued to Gamier (1829), to Koy-

mans (1833), and to Columbin (1837) (9) for various types

Schuette and Robinson — The Ice Cream Freezer . 75

of freezers. Such early inventive activity was no doubt stimu¬

lated by the perfection and popularity which ice cream had

attained in its capital for the Parisians of that day were already

well acquainted with this dish through the ice cream “parlors”

operated by the Italians Velloni and Tortoni.1 From Paris, too,

comes what is probably one of our earliest printed recipes

(1768) for making this confection (10).

Like the simplicity-loving Yankees, the British did not develop

the niceties in manufacturing this table delicacy until some

years later yet they did anticipate American invention in this

field by letters patent granted a Thomas Masters in 1843 (11)

for a device whose name suggests that the uses to which it was

adapted were many. It consisted essentially of a pewter can

containing a three-bladed revolving “spatula” and surrounded

by a “frigorific material” such as ice, new-fallen snow, or mix¬

tures of the two and common salt, sal ammoniac, salt petre,

ammonium nitrate, or calcium chloride. Lacking any of the

latter the operator might also use, mirabile dictu, diluted sul¬

phuric, concentrated muriatic or sulphurous acids in conjunc¬

tion with the snow ! The inventor summarized his claims in the

following words :

“Firstly, freezing, cooling, churning, and ice-preserving

may be conjointly and simultaneously effected.

Secondly, the solution of which ice creams and water ices

are made may be beaten up while in the act of freezing.

Thirdly, the apparatus may be applied occasionally to some

of the said purposes only and occasionally to others, and

Fourthly, each of the several parts of the apparatus may

be used with advantage either in combination or separately.”

Master’s invention of a freezer, or “churn”, was soon followed

by the publication of his “Ice Book” (1844) which was “a com¬

pendious and concise history of everything connected with ice

from its first introduction into Europe as an article of luxury”

up to the time of the appearance of this treatise. Besides which

it was “a valuable collection of the most approved recipes for

making superior water ices and ice creams at a few minutes

1 In 1798 Velloni established a magnificent place, outfitted with lounges, mirrors

and marble top tables, for the serving of ice creams, etc., at 10 Boulevard des

Italiens and similar branches throughout the city. He failed in this enterprise

and turned it over to Tortoni, an employee, who made a success of the business

while poor Velloni committed suicide. Tortoni retired in 1825 with an annual in¬

come of 100,000 francs.

76 Wisconsin Academy of Sciences , Arts , and Letters.

notice.” The publication of this book, which is deemed to be

the first one devoted exclusively to water ices and ice cream,

was marked by the simultaneous appearance of newspaper

advertisements circulated in the hope of educating the public

in the use of his freezers.

Since the evolution of the modern ice cream freezer is due

in a large measure to improvements in refrigeration processes,

a brief history of the latter will reveal some of the paths of

that evolution. In 1755 (12) a Dr. William Cullen is reported to

have developed an apparatus for freezing water by evaporation

in a partial vacuum. It was not until the middle of the follow¬

ing century, however, that mechanical refrigeration began to

assume any practical value for then Perkins (1834), Twin¬

ing (1850) and Harrison (1857) devised refrigerators that

employed the principle of evaporating a highly volatile liquid

under diminished pressure. Certain inherent defects in the

construction of the machinery necessary in operating with the

liquids in question apparently led to the development of the

ammonia refrigerator by Carre (1859). His apparatus con¬

sisted of two strong vessels, a boiler containing a concentrated

ammonia solution and an evaporating chamber, joined together

with a tube. In appearance and operation it was not unlike the

small domestic refrigeration unit which was developed several

years ago for use in those communities where electricity is not

available. To operate this machine, one raised the temperature

of the saturated ammonia solution solution in the boiler to

130-150°C. whereupon the liberated ammonia, driven over under

high pressure into the water-cooled refrigerator, condensed to

a liquid. The boiler was then placed in cold water, the effect of

which was a fall in temperature which was accompanied by a

reduction of the pressure in the apparatus, a rapid vaporization

of the liquid ammonia in the refrigerator and the production

of an intense cold. Reece (1869) improved upon this machine

in that he used brine flowing through a coil within the refriger¬

ator.

German brewers are said to have employed artificial refriger¬

ation as early as 1867. Its use in sugar refineries, meat-curing

houses and for cold storage also began at a rather early date,

but the only really extensive application which these machines

found was in the manufacture of artificial ice. And in this con-

Schuette and Robinson — The Ice Cream Freezer . 77

nection it may not be untimely to add the thought that it

requires no stretch of the imagination to picture a union of the

ice cream and artificial ice industries in that the latter found in

the former a convenient outlet for its surplus or unsaleable ice.

Until the beginning of the twentieth century freezing an ice

cream mix with manually operated equipment was still a com¬

mon procedure as witness the following reminiscence (13) of

the “good old days” by F. D. Hutchinson, one of Iowa's pioneer

manufacturers: “I remember on the Fourth of July of 1890

we shipped out three hundred gallons all frozen by hand power.”

The advent of commercial electricity made it possible, of course,

to utilize a cheaper form of energy in turning the freezer crank.

The precursor of the modern brine freezer and storage tank

is probably of the type said to have been operated by Edward

Walker in 1902 (1U). During the same year the Miller brine

freezer was developed and this was soon followed by the Miller-

Tyson machine. The reception accorded this type of freezer by

industry is reflected rather well in the following conservative

prediction, (15) “It may safely be assumed that the brine

freezer will never put the ordinary ice and salt freezers out of

business. It bids fair to play an important part in the develop¬

ment of the industry from this time on.” Commenting further

on this subject the author says, “There are some manufacturers

who still pin their faith to the steam engine regardless of cost

of installation and operation, the complicated nature of the

plant required and the necessity of employing a licensed engi¬

neer, who, in all likehood, will ever be found too busy to do

more than look after his engine. This is difficult to understand

for surely economy is as necessary in the ice cream business as

elsewhere.”

During the first decade of the present century there were in

use (16) several types of commercial freezers. Among them

were the vertical-batch ice freezer, a very common type which

at that time had already been in use for many years ; a similar

one cooled with brine ; a horizontal brine machine which seems

to have been much preferred because excellent results were

obtainable with it and little trouble experienced in getting

“swell” (17) ; and an open horizontal continuous brine freezer

which was said to possess advantages not shared by the closed

machines, especially in affording the operator an opportunity

78 Wisconsin Academy of Sciences , Arts, and Letters.

Fig. 2. Patents annually granted in the United States during the period

1860-1930.

for a more frequent use of the thermometer. Within the past

fifteen years the temperature of the refrigerants of the batch

type of freezers has been lowered by five to ten degrees, a

change in technic which has resulted in an improved texture of

the product and an increased output per unit of freezer capacity.

A direct expansion or ammonia type of freezer was brought

out in the year 1914 but it was not well received until, some

eleven years later, it had undergone numerous modifications

and improvements. The new machine immediately became

popular. It has certain advantages over the brine type of

freezer for, among other reasons, refrigeration losses were

materially reduced.

One more freezer requires mention for it enjoys a wide popu¬

larity. The machine in question is the so-called horizontal con-

Schuette and Robinson — The Ice Cream Freezer . 79

Fig. 3. Number of certain types of patents pertaining to ice cream is¬

sued in the United States 1848-1930.

tinuous disc freezer which was introduced in 1928 as an

improvement over early machines of this type which were not

entirely satisfactory, although the theory of their operation

was attractive. The ice cream made in them invariably was

coarse and fluffy. Not so, however, the present machines with

Fig. 4. Indexes of wholesale prices and patents and re-issues in the

United States 1860—1930. (Expressed as percentage deviations from the

trend in multiples of the standard deviation.)

80 Wisconsin Academy of Sciences , Arts, and Letters .

which the manufacturer is able to turn out a uniform product

of a fine texture because of a more rapid freezing at a low

temperature.

The degree of perfection which freezers have attained is

reflected in the general high state of development of ice cream

manufacturing processes. A glance at Figure 2 shows that the

annual number of ice cream freezer patents has not exhibited

any appreciable upward long time trend within the past twenty

years, perhaps because the need and the possibility for the

improvement of freezers was not as large as for the improve¬

ment of other ice cream equipment. If we look into this more

closely, we find, indeed, that during the same period the number

of patents for equipment and apparatus designed primarily to

facilitate the distribution and consumption of ice cream has

shown a large increase. From this one is led to believe that ice

cream manufacturers have pursued the general course of others

in shifting the main emphasis from improvements in the meth¬

ods of production to improvements in the methods of distribu¬

tion.

Inventive activity, as exemplified by the number of patents

granted, in the ice cream field seems to follow the course of

general inventive activity except that the amplitude of the cycli-

Fig. 5. Indexes of per capita ice cream consumption and wholesale

prices in the United States 1914-1930.

Schuette and Robinson— -The Ice Cream Freezer . 81

cal fluctuations in the ice cream patent curve is larger (Figure

3). This latter circumstance may be attributed in part to the

small numbers which we are considering-— less than fifty— but

a more definite meaning is suggested when we consider the

large positive degree of correlation existing between the per¬

centage deviation from the trend of patents and of wholesale

commodity prices (Figure 4). Ice cream is a food yet not an

essential one, hence its consumption (Table I) is dependent at

least to some extent upon the purchasing power of the consumer.

Table I. Annual per capita consumption of ice cream in the United States for

the period 1914-1930.

♦Data for the period 1914-1926 from Pirtle : A Handbook of Dairy Sta¬

tistics, Bur. Agric. Economics, U. S. Dept. Agric., 1928, p. 5.

flnterpolated. Data for the period 1927-1929 from Bur. Agric. Eco¬

nomies, U. S. Dept. Agric.

^Yearbook of Agriculture, U. S. Dept. Agric.

§ Based on estimate made by Bureau of Service and Statistics of the

International Association of Ice Cream Manufacturers. Ice Cream Trade

J., 27, No. 4, 56 (1931).

While other commodities show a more ready adjustment to

changes in the price level, the retail price and the composition

of ice cream as fixed by statute, definition or public taste remain

nearly constant over long periods of time. Thus it becomes

apparent that ice cream consumption should be doubly “sensi¬

tive” to changes in general prices or economic conditions.

Figure 5 bears out this assumption.2 Minor fluctuations in ice

cream production could perhaps be attributed to weather con¬

ditions ; but the diversity of climate over the United States fur¬

nishes us with a fairly reliable statistical sample. Thus it is to

be noted that while the weather conditions in 1930 were “good”

2 It must be borne in mind that the manufacture of ice cream was partially

restricted during the period of the World War which accounts for the low level

of production for the years 1916-1919.

82 Wisconsin Academy of Sciences, Arts, and Letters,

Population (in millions) — > w 8 2 S ? S 2 5* #§ «

from the manufacturers* point of view, a sharp decline in pro¬

duction occurred during that year (18),

Turning again to a comparison of the number of patents

granted annually and wholesale commodity prices (Figure 6),

we may conclude that there is a definite relation between varia¬

tions in these two factors. Yet the question of which is the

Schuette and Robinson — The Ice Cream Freezer . 83

forerunner or the cause in initiating the movements of the

cycles can not be answered as readily as would appear at first

sight; for the simplest view that high prices and a high general

level of business activity should result in a larger demand for

inventions does not explain the sharp drop in the number of

patents granted in 1919 and 1928, years just preceding the

“depressions”. As a possible explanation of this it may be sug¬

gested that during the height of a business cycle there is a

diminished incentive for seeking better methods of production ;

businesses almost run themselves and everyone runs a business.

The smaller annual variations in the number of patents granted

probably are an indication of the intimate connection between

the forces operating in the patent market and the forces operat¬

ing in other markets. As a fruitful source for speculation we

can see in these curves a relation between invention and money ;

a graphic illustration of Dean Hoover's view on the utility of

wars in stimulating inventive activity (but certainly not ice

cream inventions) ; and a hint of the extent of future ice cream

freezer development.

Literature Cited

1. Parkinson, “The Complete Confectioner, Pastry Cook and Baker”,

Leary and Getz. Philadelphia, 1854.

2. Winslow, Illustrated World, 25, 650 (1910).

3. W. G. Young, assignor to A. H. Reip, United States Patent 6601,

May 30, 1848.

4. A. H. Austin, United States Patent 5775, Sept. 19, 1848.

5. H. B. Masser, United States Patent 5960, Dec. 12, 1848; Reissue 751,

June 28, 1859.

6. John Decker, United States Patent 6661, Aug. 21, 1849.

7. Goldsmith Coffeen, Jr., United States Patent 6865, Nov. 13, 1849.

8. Hayward, Natl. Stockman and Farmer, reprinted in New York

Produce Rev., 14, 700 (1902).

9. Commissioner of Patents, Subject-Matter Index of Patents for In¬

ventions (Brevets d’lnvention) granted in France from 1791 to 1876 in¬

clusive, Washington, 1883, p. 394.

10. Emy, “L’art de bien faire les glaces d’office; ou, Les vrais prin-

cipes pour congeler tous les rafraichissemens”, Chez Le Clerc Paris, 1768.

11. Thos. Masters, British Patent 9825, July 6, 1843.

12. Encyclopaedia Britannica, Cambridge, 1910, 11th ed., Vol. 23, p. 31.

13. Mortensen, Proc. World's Dairy Congress 1923, 1, 468.

14. Schantz, Ice Cream Trade J., 8, No. 2, 46 (1912).

15. Anon., Ice Cream Trade J., 26, No. 1, 48 (1930).

16. Washburn, Vt. Expt. Sta. But., 155, 54-7 (1910).

17. Baer, Wise. Exp. Sta. Bui., 262, 31 (1916).

18. Hibben, Ice Cream Trade J., 27, No. 4, 56 (1931).

.

-

■

.

,

SHAKESPEARE’S USE OF ENGLISH AND FOREIGN

ELEMENTS IN THE SETTING OF

THE TWO GENTLEMEN OF VERONA

Julia Grace Wales

I

Introduction

Professor 0. J. Campbell has given us an enlightening study

of the influence of Italian comedy in The Two Gentlemen of

Verona

“A search for time-worn commonplaces of Italian comedy in

this drama,” he says, . . has convinced me that practi¬

cally all its important structural elements are patterned after

recurrent features of ‘Italian comedy’.” Among the features

having prototypes in Italian plays or scenarios he discusses the

conflict of love and friendship, the rescue of the faithless friend

by the faithful, the repentance of the former, the double wed¬

ding at the end ; the balcony scene, the lady disguised as a page,

attached to one of the amorosi ; the pathetic irony of scenes in

which the supposed page talks with or of her master, an errant

lover; the Petrarchan conceits, discourses on love, etc., the in¬

termezzi of clowns and “the contrast between the quick-witted

rogue and the slow-witted rustic”, the appearance of the dog

on the stage.

Professor Campbell finds “Shakespeare’s contribution to the

growth of romantic comedy, not in new forms of dramatic

ingenuity, but in the emotional deepening of elements taken

bodily from a drama which was at once comedy of intrigue and

highly complicated farce”.

The present study of The Two Gentlemen of Verona may be

said to deal with this “emotional deepening”, and especially

with one of the factors through which it seems to come about

— namely, Shakespeare’s treatment of the setting.

i Two Gentlemen of Verona and Italian Comedy”, in Studies in Shake¬

speare , Milton, and Donne , by members of the English Department of the Uni¬

versity of Michigan. New York, 1925.

86 Wisconsin Academy of Sciences, Arts, and Letters .

Recent historical studies of Shakespeare, Professor Camp¬

bell’s and others, serve to reawaken in our minds a number of

related questions. How are the near and the distant, the native

and the foreign, the particular and the universal related in

Shakespeare’s art? How far did the audience imagine a foreign

scene, how far an English scene? And whether the scene was

in fancy English or foreign, how did Shakespeare combine and

utilize the elements of concrete reality out of which he fashioned

it? What imaginative end did these elements serve? And

wherein does a study of them deepen and vivify our sense of

the action as a whole? These are general and more or less

speculative questions, to which precise answers can perhaps

never be given, since it is only in flashes that we can hope to

get back imaginatively into the mind of Elizabethan playwright

or audience. Yet it may be worth while to bring together again

the data we have, in the hope that brooding over them may

here and there give some new insight.

In his edition of 1921, Sir Arthur Quiller-Couch thus sum¬

marizes the facts about the play :

So far as we know, this play first achieved print in the folio of

1623, where it follows The Tempest. But it stands first on the list

of six comedies mentioned by Meres in 1598, and all internal tests,

of craftsmanship and versification, point to a date considerably

earlier yet. It is indeed, by general consent, a youthful production :

and we may safely place it somewhere near the threshold of

Shakespeare’s dramatic career.

Yet in this play Shakespeare was already combining English

and foreign elements in a shot fabric of dazzling weave. Here

where the workmanship is comparatively obvious, it is worth

while to seek hints of his later process.

II

Elements of the Setting

The Travel Motif . In the first scene of The Two Gentlemen

of Verona Valentine’s defense of travel, though it looks toward

Italy, is obviously English color. Valentine is going far away

‘To see the wonders of the world abroad”. Though, as far as

we know, Shakespeare never left England, it is clear that he

knew well the feelings, the reactions and counter-reactions, of

going from the inner to the outer world, from country to town,

and back to country and back again to town. Now he may in

Wales — The Two Gentlemen of Verona.

87

his mind have linked these transitions with the greater ones

of going from one’s own land to foreign ones and coming home

again. Hence it may not be too fanciful to seek references in

the plays to transitions from country to town as throwing light

on the wider experiences of his imaginary travellers.

“Home-keeping youths have ever homely wits”.2 It is inter¬

esting to find the words in a play which intervenes between

Love's Labour's Lost and As You Like It. The former play may

very well be an outcome of the surface fascination of town and

court life for the country boy and his first reaction against

their shallowness. A student of Love's Labour's Lost has com¬

mented on a “certain tiredness in the tone”3 of the play, but it

is the healthy tiredness of energetic youth, which will throw

off a momentary obsession. In The Two Gentlemen of Verona ,

a spirit of energy and adventure predominates. In As You Like

It, later on, Shakespeare has a rollicking holiday in the country

and comes back to town ready for the year’s work.

T. F. Ordish shows that such a holiday was an easy matter :

A young man from a provincial town, used to rural sights and

sounds, endowed with the love of nature, would not pine for the

green fields at home; he would take a walk into the country. He

would find a forest of Arden on the heights of Hampstead and

Highgate; he could take part in a sheep-shearing celebration at

even less distance. As he walked through the city on business

bent, a flock of wild duck or teal might wing over his head with

outstretched neck, taking flight from the marshes on the north

of the city, to the river or the marshes on the south between Paris

Garden and Lambeth.4 5

The link between The Two Gentlemen of Verona and As

You Like It is found again in Valentine’s soliloquy in the forest,

which foreshadows the meditations of the banished Duke :

How use doth breed a habit in a man !

This shadowy desert, unfrequented woods,

I better brook than flourishing people’d towns:

Here can I sit alone, unseen of any,

And to the nightingale’s complaining notes

Tune my distresses and record my woes.6

2 The Two Gentlemen of Verona. I. i. 2.

3 A Girton student, in a class discussion.

4 T. F. Ordish, Shakespeare’s London, 1897, pp. 7-8.

5 The Two Gentlemen of Verona. V. 4. 1-6.

88 Wisconsin Academy of Sciences , Arts, and Letters .

Thus Valentine, making a virtue of necessity, finds charm and

wholesomeness in the simple life. Of the simple and conven¬

tional life much has been written in connection with As You

Like It Yet in re-reading the play with the present question

in mind — Do Shakespeare’s English journeys throw light on

the continental journeys of his characters? — one is struck

afresh by the number of passages dealing with town and coun¬

try, life at home, and life abroad. It will serve our purpose to

turn aside and examine these for a moment.

In the first scene of As You Like It, Orlando is chafing

because, while his brother is kept at school,6 he himself is “kept

rustically at home”,7 must “feed with hinds”,8 is trained like a

peasant,9 and is not allowed “such exercises as become a gentle¬

man”.10 He wishes to go disguised to the place of wrestling* I 11

and “try the strength of” his “youth”.12

Among the older men who are associated with Orlando’s

youthful adventures, are three who have seen life, each in his

own way: (1) Adam, (2) the banished Duke, and (3) Jaques,

who, whatever his years, is constitutionally an “old gentle¬

man”,13 as Audrey calls him. Old Adam’s age is “as a lusty

winter, frosty but kindly”,14 because he has disciplined himself

by the standards of “the antique world”,

When service sweat for duty, not for meed,15

not according to “the fashion of these times,”

Where none will sweat but for promotion;

And, having that, do choke their service up

Even with the having.16

He offers Orlando what he has saved for his own time of need,17

for the old and poor the height of sacrifice. To change his place

is for Adam a painful uprooting:

0 As You Like It. I. 1. 5.

I As You Like It. I. 1. 7.

8 Ibid. I. 1. 20.

9 Ibid. 1. 1. 71.

™Ibid. 1. 1. 75.

II Ibid. I. L 131

™Ibid. I. ii. 182.

« As You Like It. V. 1. 4.

14 Ibid. II. iii. 52, 53.

™Ibid. II. iii. 57, 58.

40 Ibid. II. iii. 60-62.

” Ibid . II. iii. 30-45.

Wales — The Two Gentlemen of Verona.

89

From seventeen years till now almost fourscore

Here lived I, but now live here no more.

At seventeen years many their fortune seek;

But at fourscore it is too late a week.18

He has never grown reconciled to the condition of the world

Where what is comely

Envenoms him that bears it.19

“To most men/' says Einstein, “the court spelled misery and

disappointment. . . . The poetic tradition of satire against court

life revived from Alexandrian example was in part convention¬

al, in part caused by disappointment”.20 We may compare the

similar comment of E. K. Chambers: “It is precisely this dis¬

content of the finer spirits with the condition of the court life

that gives its burden to the pastoral comedy of As You Like It”.21

The conclusions of the banished Duke are similar. What¬

ever his inward regrets, he has found the woods “more free

from peril than the envious court”,22 and “the churlish chiding

of the winter’s wind”23 a better counsellor than the flatterers of

prosperous days.24 As for Jaques, he has carried into the wil¬

derness a “complex” of resentment against the world. The

song sung by the courtly foresters as they lay the cloth25 for

a picnic dinner voices the optimism of the Duke’s view of the

simple life:

Who doth ambition shun

And loves to sit in the sun,

Seeking the food he eats

And pleas’d with what he gets,

Come hither, come hither, come hither.

Here shall he see

No enemy

But winter and rough weather.28

Jaques’ addition is more cynical,27 and the song of Amiens gives

the key of their melancholy :

18 Ibid. II. iii. 71-74.

19 Ibid. II. iii. 14-15.

20 Lewis Einstein, Tudor Ideals, New York, 1921, p. 45-7.

21 E. K. Chambers, Shakespeare’s England, 1916, I. iii, p. 82.

22 As You Like It. II. i. 4.

28 Ibid. II. i. 7.

24 Ibid. II. i. 10-11.

™Ibid. II. v. 31.

26 Ibid. II. v. 40-47.

21 Ibid. II. v. 52-59.

90 Wisconsin Academy of Sciences , Arts , and Letters .

Blow, blow, thou winter wind,

Thou art not so unkind

As man’s ingratitude .

Most friendship is feigning, most loving mere folly :

Then, heigh-ho, the holly!

This life is most jolly.

Freeze, freeze, thou bitter sky,

Thou dost not bite so nigh

As benefits forgot:

Though thou the waters warp,

Thy sting is not so sharp

As friend remembered not.28

Orlando has been touched with the same infection, though his

disposition has escaped remarkably well.

Why should this a desert be?

For it is unpeopled? No:

Tongues I’ll hang on every tree

That shall civil sayings show.

Some how brief the life of man

Runs his erring pilgrimage,

That the stretching of a span

Buckles in his sum of age;

Some of violated vows

’Twixt the souls of friend and friend.29

The country atmosphere of As You Like It is of course ro¬

manticized, partly conformed to the conventions of the pas¬

toral ; yet there are flashes of realism. Orlando who is “inland

bred”,30 having made the city man’s little mistake about the

country, hastens to apologize :

Pardon me, I pray you:

I thought that all things had been savage here;

And therefore I put on the countenance

Of stern commandment.81

Touchstone gets into similar difficulties, but Rosalind is more

tactful :

Touchstone. Holla, you clown!

Rosalind. Peace, fool: he’s not thy kinsman.

Corin. Who calls?

Touchstone. Your betters, sir.

28 As You Like It. II. vii. 174-6, 181-189.

29 Ibid. III. ii. 133-143.

80 Ibid. II. vii. 106-109.

81 Ibid. II. vii. 96

Wales — The Two Gentlemen of Verona .

91

Corin. Else are they very wretched.

Rosalind. Peace, I say. — Good even to you, friend.

Corin. And to you, gentle sir, and to you all.32

Later in the play, Corin and Touchstone discuss the manners

of shepherds and courtiers for some seventy-five lines.33 Corin,

though literal-minded, is no fool. He has the wit to point out

that ‘Those that are good manners at the court are as ridiculous

in the country as the behaviour of the country is mockable at

the court.'' And Touchstone, while pretending to mock at

Gorin's rusticity, helps him to prove further that rustic man¬

ners can be intrinsically at least as good as courtly manners,

that the latter are often artificial and not as fastidious as they

seem.

The conclusion of the whole matter — country versus town,

the simple life versus the court, home versus the great world

— is left somewhat obscure. Several pairs of our foresters

wrangle over some side of it. Jaques easily talks down the

optimism of the Duke and is as easily worsted by the common

sense of Rosalind.

Jaques .... but it is a melancholy of mine own, compounded of

many simples, extracted from many objects, and indeed the sun¬

dry contemplation of my travels, which, by often rumination,

wraps me in a most humorous sadness.

Rosalind. A traveller! By my faith, you have great reason to be

sad. I fear you have sold your own lands to see other men's ; then,

to have seen much and to have nothing, is to have rich eyes and

poor hands.

Jaques. Yes, I have gained my experience.

Rosalind. And your experience makes you sad. I had rather have

a fool to make me merry than experience to make me sad ; and to

travel for it too ! 84

In the end all hie themselves cheerfully back to public life

— with two exceptions. The comic, impulsive, suspicious, wor¬

ried, highly suggestible little tyrant, Frederick, has, quite in

character,

put on religious life,

And thrown into neglect the pompous court.*5

82 As You Like It. II. Iv. 66-70.

83 Ibid. III. ii. 11-90.

34 As You Like It. IV. i. 15-29.

35 Ibid. V. iv, 187-188.

92 Wisconsin Academy of Sciences, Arts, and Letters.

And Jaques, his intellectual curiosity piqued by this conver¬

sion, will stay to consider matters further at the Duke’s aban¬

doned cave.

These ideas are of course commonplaces of the pastoral.36

But commonplaces themselves reveal contemporary reality, and

may betray the mind of a man whose nature it was to assimi¬

late the imaginative elements of his age. Einstein’s comment

on this point is interesting:

English country life became established on its modern basis dur¬

ing the Sixteenth Century. The novel conditions of stability and

order, the diffusion of luxury and the rise of a new propertied

class, were all circumstances which made for its appreciation and

which in turn were to be productive of a domesticated idea of na¬

ture. Another cause contributing to this result was entirely in¬

tellectual. The revival of classical antiquity and its lesson both di¬

rect and indirect, through text, translation, and continental influ¬

ence, added to this feeling, and by an odd paradox, dusty manu¬

scripts helped to arouse the sense of nature. Men loved the coun¬

try more after they had read Theocritus and Virgil, while so arti¬

ficial a growth as the pastoral comedy could by devious paths

throw back to nature.37

Let us return now to the play which deals directly with

travel. Valentine feels that a love affair is the only excuse for

not setting forth to see “the wonders of the world abroad”.38

Proteus also takes seriously the privilege of seeing “rare and

noteworthy objects”39 in travel, and in his soliloquy commends

his friend’s course rather than his own:

He after honour hunts; I after love:

He leaves his friends to dignify them more;

Thou, Julia, thou has metamorphosed me —

Made me neglect my studies, lose my time,

War with good counsel, set the world at nought.40

Similarly in a later scene, the father of Proteus is reproached

for suffering his son to spend his youth at home:

36 For the vogue of the pastoral in England see E. K. Chambers, English Pas¬

torals, London, 1906 ; A. H. Thorndike, “The Pastoral Element in the English

Drama before 1605”, Mod. Lang. Notes, XIV (1899), 4, p. 227; W. W. Greg,

Pastoral Poetry and Pastoral Drama, London, 1906 ; C. F. Tucker Brooke, The

Tudor Drama, 1911, Ch. viii & Bibliography.

37 Einstein, Tudor Ideals, New York, 1921, p. 269.

38 The Two Gentlemen of Verona, I. i. 6.

83 Ibid. I. i. 13. Cf. Bacon : Of Travel.

40 Ibid. I. i. 64-65, 67-69.

Wales— The Two Gentlemen of Verona .

93

While other men of slender reputation,

Put forth their sons to seek preferment out ;

Some to the wars to try their fortunes there;

Some to discover islands far away;

Some to the studious universities.

For any or for all these exercises

He said that Proteus your son was meet,

And did request me to importune you

To let him spend his time no more at home,

Which would be great impeachment to his age

In having known no travel in his youth.41

W. J. Rolfe’s note on this passage is illuminating. He quotes

Gifford, Memoirs of Ben Jons on:

The nobility, who had been nursed in domestic turbulence, for

which there was now no place, and the more active spirits among

the gentry, for whom entertainment could no longer be found in

feudal grandeur and hospitality, took advantage of the diversity

of employment happily opened, and spread themselves in every

direction.

Rolfe also quotes Knight:

Here in three lines we have a recital of the great principles that

whether separately, or more frequently in combination, gave their

impulses to the ambition of an Essex, a Sidney, a Raleigh, and a

Drake.42

Antonio has already been hammering on the same notion:

I have considered well his loss of time

And how he cannot be a perfect man,

Not being tried and tutored in the world;

Experience is by industry achieved,

Then, tell me, whither were I best to send him?13

In these two passages there is an interesting fusion of two

appeals to the English imagination — that of the New World

and that of the Italian Renaissance, the appeal on the one hand

of the undiscovered and untried, and on the other of a mature

and traditional civilization. I do not here mean to imply that

the effect of the Italian Renaissance in England was conserva¬

tive — it was, of course, in the main, far other — but only that

Englishmen turning to the continent of Europe, came under

sophisticated, not primitive, influences. Part of the apparently

41 Ibid. I. iii. 6-16

42 See pages 130-32 and pag-e 11.

43 The Two Gentlemen of Verona. I. iii. 19-24.

94 Wisconsin Academy of Sciences, Arts, and Letters .

magical power of the sixteenth century in literature is due to

the fusion of these two forms of appeal. Neither alone could

be such a source of power. Emphasis upon the old, emphasis

upon the new, will lead equally to decadence. It is the impact

of new upon old that is the creative force. In our own period

of conscious intellectualization, conservative and radical ten¬

dencies too often eye each other across a gulf of controversy.

The more intuitive Elizabethan instinctively combined the two.

There is a certain contrast between the wholehearted belief

in the value of travel as expressed in these passages and the

Puritan fear of the evil influences abroad expressed by Ascham

and others.44 Yet in both there is the same tendency, peren¬

nially English, to consider practical and ethical values. The

traveller went forth to improve himself, to study and report

systematically, and to do a service for his nation.

To quote a contemporary writer on travel :

Wherefore both in these days and in all ages heretofore the best

and wisest, the chiefe and noblest men, have alwaie travelled, as

by example might be proved, were it not tedious to entreat of a

matter so presumptuous ... so to profite, and inrich themselves

with experience and true wisdom, and especially to benefite their

owne proper and natural countrie, they traversed over and trav¬

elled into other countries.45

The young man is warned against pleasure-seeking and ad¬

monished to take his opportunities seriously.

In The Traveller of Jerome Turler we find similar senti¬

ments : Travel should help a man

to discern what is good and bad in his owne countrey ... He

shall also have more skyll how to entertain strangers, and under¬

stand the maners of men more perfectly, and according to his

affayres and dealinges with them, applie himself unto them ac¬

cording as the circumstances of time and place shall require.46

No less striking than this motive of the conscientious tourist

is that of the adventurous pioneer. In the sixteenth century

44 See Einstein, The Italian Renaissance in England, New York, 1902, Chap¬

ter IV, “The Italian Danger”. Einstein quotes Ascham, Sandys, Dallington, Nashe,

Gascoigne, Harrison, Mulcaster, and others. Cf. Clare Howard, English Trav¬

ellers of the Renaissance, London and New York, 1914, Chapter III, “Some

Cynical Aspersions upon the Benefits of Travel”, pp. 50-70.

45 A Direction for Travellers. Taken out of Justus Lipsius and enlarged for

the behoofe of the right honourable Lord, the young Earle of Bedford, being

now ready to travell. London, 1592. (Brit. Mus. 10024. Acc. 5) p. 3.

48 The Traveller of Jerome Turler, London, 1575. See also, W. B. Rye, Eng¬

land As Seen by Foreigners, London, 1865, pp. xix-xxviii, quotations from several

early English works in praise and censure of foreign travel.

Wales— The Two Gentlemen of Verona .

95

the pioneer was still discoverer rather than “settler”. And

yet even at this time are discernible the two qualities which

have together made the “settler” — the love of home mixed

with the love of adventure, the home basis of contentment itself

giving depth and vitality to the wandering spirit — the tend¬

ency to go and return, or if return is not feasible, to build

stable New Englands overseas. This double quality is re¬

marked by a nineteenth century student of English life.

That Englishmen are such hardy explorers, such persistent set¬

tlers of the waste places of the earth, attests their love of home.

They go, not because they wish to go, but because they hope to

return with enough to establish a home in England . To

have a house and bit of garden of one’s own, an Englishman or

woman will submit to the utmost economy of expenditure, and the

most rigorously accurate system of accounts. It may be a social

prejudice or an ingrained habit of the British stamp of mind, but

whatever it is, there can be no doubt, that the Englishman’s ideal

of life is to be a free man and master of the castle of his own

house.47

The quality is of course not merely English but a charac¬

teristic contribution of the Northern races to human develop¬

ment.

The Elizabethan attested his belief in the value of travel by

willingness to face the dangers it entailed.

If dangers do environ thee,

says Proteus,

Commend thy grievance to my holy prayers,

For I will be thy beadsman, Valentine.

These dangers were considerable in England.

Travellers were exposed to a variety of risks .... Apart from

the dangers incident to the state of the roads and bridges, there

was the possibility of encountering highway robbers .... Or¬

ganized gangs infested exposed places .... Travellers who car¬

ried no great valuables were as liable to attack as the richly-

laden. But highwaymen were commonly credited with merciful

treatment of the very poor.48

The dangers were at least as great in Italy.

47 Price Collier, England and the English , 1909, p. 173.

48 Shakespeare’s England, London, 1916, Vol. I, Chap, vii, “Land Travel”, by

Charles Hughes, pp. 207-8. See also bibliography for the chapter — Harrison,

Moryson, etc.

96 Wisconsin Academy of Sciences , Arts, and Letters .

The long series of invasions rendered many parts of Italy almost

impassable to women, and since the larger and better policed

states of the sixteenth century still bestowed their mauvais sujets

on their neighbours, who no longer welcomed the exiles with open

arms, the banished men either hired themselves as secret agents

to do murder that could not now be openly contrived, or took to

the hills and preyed on the wayfarer.49

Perhaps even greater dangers were encountered in journeying

from England to Italy. On the dangers to travellers Clare

Howard in English Travellers of the Renaissance50 cites many

contemporary references showing that travellers meet with

dangers not a few by sea and land, from highwaymen and

other robbers, to disease and exposure. Which of these kinds

of journeys had Shakespeare uppermost in mind when he

thought of the dangers of travel? We can perhaps safely con¬

jecture that in his imagination there was a mixture of all three.

But is there any evidence within the play that, whatever the

casual suggestions from which his picture of travel was made,

he tried to imagine and meant his audience to imagine not a

journey up to London from Stratford, nor a voyage from Eng¬

land to Italy, but a journey literally from Verona to Milan?

That to all intents and purposes Valentine sailed from Lon¬

don and landed at some Italian port, seems at first glance, to

find support in allusions to distance,51 dangers,52 letters,53 ab¬

sent friends,54 remittances of money,55 and especially in the fact

that the journey is by water.56 This last, however, is less sig¬

nificant than it seems. For Sullivan, in his researches on the

waterways of Northern Italy in the sixteenth century57 has

shown that the facts do not contradict anything that Shake¬

speare has implied as to a possible river route from Verona

to Milan. The Adige was the main highway from Verona to

49 wm. Boulting, Women in Italy, London, 1910, p. 209.

1914, p. 47.

81 The Two Gentlemen of Verona, I. iii. Proteus is to travel in company.

52 Ibid. I. i. 16.

82 Ibid. I. i. 57.

54 Ibid. II. iv. 59.

88 Ibid. I. iii. 58-59.

86 The Two Gentlemen of Verona, II. iii.

57 See Edward Sullivan, “Shakespeare and the Waterways of Northern Italy,”

the Nineteenth Century, 1908, August, 64, p. 215. Sullivan cites numerous pass¬

ages from Italian works of the fifteenth and sixteenth centuries and from the

works of travellers regarding these river routes. See also M. P. Tilly, “Shake¬

speare and Italian Geography,” Jour. Eng. and Germ. Phil., 1916, p. 454. Tilly

quotes The Pilgrimage of Sir R. Guylforde, Knight to the Holy Land, A. D. 1506,

which describes just such a journey by river and canal.

Wales— The Two Gentlemen of Verona .

97

many Italian cities. Similarly Sarrazin finds realistic allusions

in Saint Gregory’s Well, the forest near Milan, “the rising of

the mountain foot that leads toward Mantua.”58 While these

allusions are perhaps not definite and precise enough to justify

the conclusion that Shakespeare had at this time much detailed

knowledge of Italy, they are at least convincing evidence that

he meant to give a degree of local color to the play.

Thus, however we came here, we are, as we enter into the

action of the play, to feel ourselves in Italy ; and it has not been

stage scenery, nor even any considerable mass of detailed allu¬

sions,59 such as Shakespeare uses in later plays, but chiefly the

general talk of journeys by sea and land, that has wrought the

magical transition and wafted us hither. This, then, for the

purposes of the play itself, is the significance of so marked an

emphasis on the glamor of foreign scenes. It has none the

less a wider significance : it reflects an ever present element in

the Elizabethan consciousness and in Shakespeare’s own ima¬

gination, — the absorbing and many-sided interest, individual

and national, materialistic, romantic, and cultural, of travel and

adventure in the New World and the Old. Nor is this spirit

inconsistent with homelier passages in As You Like It and

other plays. Shakespeare’s imagination found everywhere

more than enough to keep it busy, and it is easy to believe that

he was well content to spend his days in his own country. But

had circumstances favored his going abroad no man would

have enjoyed seeing the world more than he. And no doubt it

delighted him to pick up knowledge of distant places and to

imagine the scenes of his plays in foreign lands.

Minor Romantic Machinery . A Shakespearean scholar has

casually expressed the opinion that Shakespeare uses Italian

color chiefly as romantic background,60 much as the Balkan

states have been used in popular English novels, as a region

88 G. Sarrazin, “Neue Italienische Skizzen zu Shakespeare,” D. S. G. Jahrbuch,

Vol. XXXIX (1903), pp. 62-68. See also E. Koeppel, Jahrbuch, XLIII, p. 247.

Koeppel attempts to demolish Sarrazin’s theory of St. Gregory’s Well as an in¬

dication that Shakespeare knew Italy. He points out that in passages cited by

Sarrazin there is no actual mention of a well, and that as the hospital was for

the plague, there probably would not be a famous spring in such a dangerous

neighborhood. See also a full note on this passage in the Arden Edition, p. 81.

69 See Sir Edward Sullivan’s discussion of the names in the play — not all

Italian. He notes that almost no Italian words or phrases occur. See “Shake¬

speare in Italy”, Nineteenth Century, LXXXIII, p. 146.

60 Miss Janet Spens in conversation. I quote with permission.

98 Wisconsin Academy of Sciences, Arts, and Letters .

far away, not too precisely known, favorable to all kinds of

adventure. The present play, especially, lends color to this

theory through the large proportion of details of purely roman¬

tic machinery as compared with more realistic furnishing: a

disguise for a lady,61 including doublet and jerkin,62 her hair to

be knit up with silken strings,63 a chamber window,64 a lady’s

picture,65 two rings,66 a sealed sonnet,67 banishment,68 a sere¬

nade,69 a thicket where a nightingale sings,70 a lost glove,71 a

corded ladder provided with anchoring hooks,72 an upper tower

under lock and key,73

And built so shelving that one cannot climb it

Without apparent hazard of his life,74

a long cloak,75 a friar’s cell,76 an abbey, a postern by the abbey

wall,77 an emperor’s court,78 an out-law infested forest,79 a rob¬

ber’s cave.80

These may seem homogeneous elements, but they have a cer¬

tain variety. They include many of the stock properties of the

romantic drama. In the unscalable tower and the dungeon we

have further accessories of mediaeval romance. It is to do the

same romantic service that the monastic elements come in.

These, like the “corded stair”, are suggested by Brooke’s

Romeus and Juliet*1 Silvia goes to Friar Patrick’s cell as Juliet

to Friar Laurence’s. But the Friar Laurence of this play, wan¬

dering through the forest in the garb of his order, muttering

his penitential prayers, is more alert and effectual than the

61 The Two Gentlemen of Verona. II. vii. 39

62 Ibid. II. vii. 49.

63 Ibid. II. vii. 45.

64 Ibid. IV. ii.

66 Ibid. Also IV. iv. 122.

68 Ibid. IV. iv. 137 ; V. iv. 96.

67 Ibid. III. i. 140-150.

88 Ibid. IV. i. 170-187.

89 Ibid. IV. ii.

78 Ibid. V. iv. 1-6.

71 Ibid. II. i. 3, 4.

72 Ibid. III. i. 117-118.

73 Ibid. II. i. 36.

74 The Two Gentlemen of Verona. III. i. 115-116.

26 Ibid. III. i. 131.

78 Ibid. V. ii. 42.

77 Ibid. V. ii. 9.

78 Ibid. I. iii. 27.

79 Act IV. Scene i.

80 Act IV. iii. 12.

81 Arthur Brooke, Romeus and Juliet, Ed. J. J. Munro, London, 1908, 11.813-816.

Wales— The Two Gentlemen of Verona .

99

benevolent priest of the later play. No doubt he was none the

less suggested by Brooke's Franciscan Friar Laurence.

This barefoot friar girt with cord his grayish weed,

For he of Francis’ order was, a friar, as I rede.82

Brooke has no mention of the abbey. We possibly have the hint

for that in Montemayor's Diana: “My brother and I were

brought up in a nunnerie, where an aunt of ours was abbesse.”83

“The postern by the abbey wall” no doubt suggested the paral¬

lel phrase “behind the abbey wall” in Romeo and Juliet . Ordish

says : “ ‘Out at the postern, by the abbey wall’ was a line in

all probability suggested by a London locality.”84 Eglamour

has been associated with these ecclesiastics because of his vow

of chastity and because his helping Silvia to escape is a remote

parallel to Laurence's intended aid to Juliet. But he is really

a very different element and an interesting one. Quiller-Couch

and Wilson85 believe the “adapter”, not Shakespeare, to be

responsible for his disappearance from the play.

Now these romantic elements themselves, especially window

scenes and disguises, were, as Professor Campbell has so fully

established, material of Italian plays and stories, and were asso¬

ciated in the English mind with an Italian background. Also

the window, and in some cases the disguise,86 actually played

an important part in the life of the Italian woman of the six¬

teenth century. Boulting speaks of the balcony as a means of

flirting.87 Similarly the opportunity that her religious duties

gave the young girl to get away from the imprisonment of her

father's home into the thrill of the outer world,88 was a privilege

dearly cherished and often abused. And, of course, in a general

sense, Roman Catholic color is proper to Italy. Thus on the

whole, it is safe to say that in this play, at least to some extent,

Shakespeare uses romantic machinery for Italy as well as Italy

for romance.

82 Ibid. 11.565-6.

83 Hazlitt’s Shakespeare’s Library, Vol. I. p. 278.

84 Op. cit., p. 172.

85 See p. 115-6, below. Cf. Op. cit., p. xvi-xvii.

88 See Victor Owen Freeburg, Disguise Plots in Elizabethan Drama, New York,

1915.

87 Boulting, Woman in Italy, pp. 117-118.

88 See Boulting, p. 119. “Church was the accepted covert for a woman to

wait in ambush, make eyes and display her charms.”

100 Wisconsin Academy of Sciences, Arts, and Letters.

The Emperor's Court. But two elements of the romantic

machinery are less certain in their geographical suggestion:

the Emperor's Court, and the Wood.

Before we turn to these let us pause to note a view put for¬

ward by T. F. Ordish in Shakespeare's London:

The fact that Shakespeare took his plots and stories from foreign

sources, retaining the names and locales of the originals, has

proved a more effective disguise to posterity than it was to the

Elizabethans. With them the device was taken for granted— it

was the mode of the time. It constituted part of the “play,” with

the audience, to detect the reality beneath the mask, the actual

beneath the fictitious.80

Thus Ordish would perhaps say of the Emperor's Court that

it is in reality only the court of Elizabeth, where the would-be

courtier

shall practice tilts and tournaments,

Hear sweet discourse, converse with noblemen,

And be in eye of every exercise

Worthy his youth and nobleness of birth.

Tilts and tournaments are mentioned in Diana.

When he had, therefore, by sundrie signes, as by Tylt and Tour-

neyes, and by prauncing up and down upon his proude jennet be¬

fore my windowes, made it manifest that he was in love with me

(for at the first I did not so well perceive it) he determined in the

end to write a letter unto me.80

Shakespeare no doubt gathered most of his knowledge of

courts and courtiers from his chance acquaintance with men of

the world. There was a literature of the subject open to him,

however. That Shakespeare knew Castiglione's Book of the

Courtier finds evidence in Much Ado.91 T. F. Crane in Italian

Social Customs of the Sixteenth Century gives us an idea of the

extensive influence of The Book of the Courtier in Europe. He

comments on Johnson’s high opinion of The Courtier, and adds

a quotation which might well be a commentary on the general

situation in the present play :

“I observed” says Boswell, “that at some courts in Germany,

there were academies for the pages, who are the sons of gentle-

89 T. Fairman Ordish, Shakespeare’s London , 1897, p. 140.

90 Shakespeare’s Library , 1875, I, p. 279.

91 See M. A. Scott, “The Book of the Courtier : a possible Benedick and Bea¬

trice.” P. M. L . A., V. 16 (1901), pp. 475-502.

Wales— The Two Gentlemen of Verona .

101

men, and receive their education without any expense to their

parents.” Dr. Johnson said, that manners were best learned at

these courts. “You are admitted with great facility to the

prince’s company, and yet must treat him with respect. At a

great court, you are at such a distance that you get no good.” 92

Einstein says :

Halfway between the learning of the scholar, and the practice in

sport and arms of the knight, was the new Renaissance idea of

a gentleman’s education. Borrowed largely from the Italian and

French writers, though also from Spanish and even Polish, a

body of opinion grew up for the training of those who aimed to

fit themselves to be of service to their country. They were ad¬

vised to study laws and treaties, civil policy and moral sciences.

. For the rich, travel became the approved meth¬

od, and even those of small means like Sidney went from country

to country in search of experience. The system was at best hap¬

hazard.98

He goes on to speak of Humphrey Gilbert’s plan of an academy.

The programme of studies embraced civil government and fi¬

nances, martial exercises, and surgery. This was the real human¬

ist ideal applied to life with its vision of a New England ventur¬

ing into distant lands. Gilbert had realized the promise of what

the colonies were to mean for British expansion, but his sugges¬

tion met with no response from the most parsimonious of

rulers.94

In our play other gentlemen of good estimation have jour¬

neyed to the court to salute the sovereign. Quiller-Couch and

Wilson quote Stevens:95

Shakespeare has been guilty of no mistake in placing the em¬

peror’s court at Milan. Several of the German emperors held

their courts there occasionally.

»2 t. F. Crane, Italian Social Customs of the Sixteenth Century (1920), Chap.

IV, pp. 205-7. See also (referred to by Crane, p. 205) Sir Walter Raleigh’s in¬

troduction to Hoby’s translation of Castiglione, London, 1900, passim. See also

M. A. Scott, Elizabethan Translations from the Italian, Boston and New York,

1916, descriptions of some twenty-one books on manners and morals, translated

from the Italian into English and published in England between the years 1561

and 1607.

93 Einstein, Tudor Ideals, 1921, pp. 54-5. A footnote refers us to Grimaldus

Goslicius : The Counsellor.

94 Tudor Ideals, p. 166. On the education of a gentleman see also for ex¬

ample, Sir Philip Sidney’s letters to his brother Robert, ( The Miscellaneous

Works of Sir Philip Sidney, London, 1893, pp. 328-339) for advice on travel,

horsemanship, fencing, etc.

93 See their edition of The Two Gentlemen of Verona, 1921, p. 88.

102 Wisconsin Academy of Sciences , Arts , and Letters .

They think “emperor” was the Shakespearean style and “Duke”

that of the theater and the abridger. Professor Young says,

however,96

We need not infer that Shakespeare had in mind the occasional

sojourn of Charles V (Emperor 1519-1556) at Milan. ‘Emperor’

here probably refers only to the Duke of Milan.

The court is a place of set love-making, fanciful compliments,

artificially witty conversation. There is an extensive literature

not only of worthy ideals and accomplishments in general but

of conversation in particular. In reading Crane's chapter on

Parlor Games in Italy,97 we realize how highly conventionalized

the give and take of social intercourse had become abroad. In

the England of Shakespeare’s plays (springing out of the ordi¬

nary life of cruder people) the talk is of course more demo¬

cratic and less patterned than in the Italian books of social

usage, but we are not surprised to find behind any pun, riddle,

question, or debate something harking back to Italy. When

Valentine says, “I have dined,” he is far from original. Com¬

pare a dialogue from Lyly:98

Manes . . . which I have not done these three days.

Pyllus. What is that?

Manes. Dined ....

Pyllus .... How many have so fed their eyes with their mistress

picture that they never desired to take food, being glutted with

the delight in their favours."

In a delightful chapter on “The High Purpose of the Eliza¬

bethan Traveller,” Clare Howard summarizes the directions of

the manuals for travellers:

They have in common the tendency to rationalize the activities of

man which was so marked a feature of the Renaissance. The sim¬

ple errant impulse that Chaucer noted as belonging with the

96 K. Young, The Two Gentlemen of Verona, New Haven, 1924, p. 85.

97 See Crane, Italian Social Customs in the Sixteenth Century (1920) passim,

especially Chapter VI.

98 Alexander and Campaspe, I. ii. 14, 65-7. Ed. R. Warwick Bond, V. II. p. 320.

98 Compare Geoffrey Fenton, Certain Tragical Discourses of Bandello (1567),

Discourse XIII, pp. 248-9 : “And in place to performe the expectation of his hos¬

tess in tasting the sondrie delicate meates she prepared for hym, he fed only

upon the dishes of love ; and contenting hymselfe with the dyot of his eyes, who

. . . imparted their norriture to the hearte” etc. I do not know whether any one

else has remarked these rather obvious and very insignificant parallels. For a

discussion of this general type of indebtedness, see R. Warwick Bond, The Com¬

plete Works of John Lyly, 1902, Vol. II. pp. 473-86 : “Note on Italian Influence.”

Wales — The Two Gentlemen of Verona .

103

songs of birds and coming of spring, is dignified into a philosophy

of travel . . . .10°

In short, the perils and discomforts of travel made a mild pre¬

lude to the real life into which a young man must presently fight

his way. Only experience could teach him how to be cunning,

wary, and bold; how he might hold his own, at court or at sea,

among Elizabeth’s adventurers.101

In another passage the same writer says :

Underneath worldly ambition was the old curiosity to see the

world and know all sorts of men — -to be tried and tested. More

powerful than any theory of education was the yearning for far-

off, foreign things, and the magic of the sea.102

We may compare with this a passage from Shakespeare's

England:103

Thus the court was a place where high prizes were to be won.

To the lads of England it offered in anticipation a romantic ad¬

venture and in retrospect too often a memory of sordid intrigue.

So confesses Sir John Harrington, a godson of the Queen, who

fluttered in the wake of Essex, and just escaped being entangled

in his fall.

‘I have spente my time, my fortune, and almost my honestie,

to buy false hopes, false friends, and shallow praise; — and be it

remembered that he who casteth up this reckoning of a courtlie

minion, will sette his summe like a foole at the ende, for not be-

inge a knave at the beginninge.’

At the court, we find in our play, one meets foreign visitors

with commendations from great potentates, tyrannical fathers,

caddish rivals, ladies versed in the ways of the world — all new

to Valentine. Silvia is a great lady, not a little spoiled, used to

having her own way, an accomplished flirt. Her admirers are

her slaves, and she means to keep them all in an obedient humor.

When Valentine's temper and bluntness get the better of him,

she welcomes interruptions, murmuring sweetly,

No more, no more, gentlemen— -Here comes my father.104

When Proteus arrives at the court, she instantly enrolls him

100 Howard, English Travellers of the Renaissance , London and New York, 1894.

Chapter II, p. 29.

101 Howard, English Travellers of the Renaissance, London and New York, 1894,

p. 30. See also the bibliography.

102 Ibid. p. 19.

10Z Shakespeare’s England, 1916, I. iii. pp. 81-82.

104 II. iv. 47.

104 Wisconsin Academy of Sciences, Arts, and Letters .

among her servitors, tossing down challenges which he knows

only too well how to take up.105

Such details, it is true, serve equally well to fill in a picture

of court life far away or near at hand. Yet, as Professor Pyre

has pointed out106, we can say that the court itself, that a soph¬

isticated society itself, was associated in the popular mind with

continental and above all Italian cities, that the English little

Italy of the sixteenth century was found in the higher circles

of London life. The cosmopolite of the sixteenth century was

Italianate. W. B. Rye quotes from Cardan (1585) :107

In dress they are like Italians .... I wondered much especially

when I was in England and rode about on horseback in the neigh¬

borhood of London, for I seemed to be in Italy. When I looked

among those groups of English sitting together, I completely

thought myself to be among Italians; they were like, as I saw, in

figure, manners, dress, gesture, colour; but when they opened

their mouths I could not understand so much as a word, and won¬

dered at them as if they were my countrymen gone mad and

raving.

An Elizabethan traveller says:108

... it is growen into a proverbe amonge the Italians, Thedesco

Italianato, Diaholo incarnato : that is to saye, a Dutchman become

in maners lyke an Italian putteth on the nature of the Devill,

and is apt unto all kinds of wickednesse.

The Wood . The wood presents a similar problem. Ordish

says of it : “The scene in the forest of the frontiers of Mantua

could also be realized without the aid of scenic art. All play¬

goers, all Londoners, knew the forest which almost encompassed

London on the Middlesex side”.109 Sarrazin, though he admits

discrepancies, finds a real forest “upon the rising of the moun¬

tain foot that leads toward Mantua”.110 Miss Spens emphasizes

the romantic and traditional element and finds that the wood,

like the forest of Arden and like Sherwood in Munday’s Robert

105 II. iv. 102-115.

106 In lecturing at the University of Wisconsin.

1OT W. B. Rye, England as Seen by Foreigners , London, 1865, p. 7.

108 Turler, The Traveller of Jerome Turler , London, 1575, p. 66.

109 ordish, Shakespeare’s London, 1897, pp. 112-3.

no The researches of Sir Edward Sullivan make it further possible to realize

the mountain foot in the realm of the actual. “I now find there were two high¬

ways between Milan and Mantua, and that the N. E. road passes ‘the mountain-

foot’ when nearing Palazzolo — (See Schotti Itinerarium Italiae, De Brixia)”

“Shakespeare and Italy”, the Nineteenth Century, LXXXIII, p. 339.

Wales — The Two Gentlemen of Verona .

105

Earl of Huntington is the woodland scene of the Robin Hooa

stories, and that the outlaws are our ancient playfellows, the

Merry Men of English oral tradition :

Robin Hood seems to have been something of an obsession

with Monday, and it may therefore not be without significance

that Shakespeare twice uses the outlaw motive in his comedies:

in The Two Gentlemen of Verona (which we have seen is prob¬

ably otherwise indebted to Munday) and in As You Like It. The

name of Robin Hood is mentioned, however, merely in passing.

What Shakespeare gets is the free life of the forest. The effect of

this natural background in The Two Gentlemen of Verona is al¬

most magical.111

Certain it is that the outlaws “practice an honorable kind of

thieving”,112 like a manly fight, are chivalrous “to silly women

and poor passengers”,113 welcome a brave leader, are capable of

being reformed, “civil, full of good, and fit for great employ¬

ment”.114

Though Robin Hood is English, the pastoral romance is of

wide continental connections. Sidney’s Arcadia, itself a region

where anything can happen, tells of far, far away and draws

on continental sources. As You Like It, moreover, comes as

much from Lodge’s Rosalynd as from oral traditions of Robin

Hood; and a striking analogue of Rosalynd (and possibly a

source) is to be found in Certain Tragical Discourses of Ban -

dello.115 Here we find — located in Spain some of the flora and

fauna of Arden, much of its free spirit, a young man and his

servant making themselves at home with Crusoe-like ingenuity

in the wilderness,115 the young man disguised as a pilgrim and

relieving his feelings in an unfortunate love affair by carving

verses on rocks and trees. In our effort to locate “the rising of

the mountain foot” it may be worth pausing to note that three

or four days’ and nights’ wandering on the part of Bandello’s

111 Janet Spens, Shakespeare and Tradition, Oxford, 1916, p. 33.

112 IV. i. 40.

113 IV. i. 72.

114 V. iv. 156-7.

115 There his man made provision accordynge to the condicion of their state

and necessity of the place, dyggynge for his firste indevor certeine soddes and

lomppes of claye, wherewith he entrenched and rampierd their felden shopp, to

defende theym aganste the furye of wilde beastes, who other wayes myghte op-

presse theym in the nyghte. He made, also, two beddes, or lytle couches, of

softe mosse, wyth a testure and sides of wodde, which he hewde in no less fyne

proporcion then yf the skill of the carpenter had assisted the worke.” — Certain

Tragical Discourses of Bandello, translated by Geoffrey Fenton, Discourse XIII,

pp. 273-4. ( Tudor Translations, Vol. XX.)

106 Wisconsin Academy of Sciences , Arts , and Letters .

travellers “broughte them at last to the foot of a large moun-

taine, inhabited only with savage beasts and creatures unrea¬

sonable”.116 Truly if we are Elizabethans we have been in this

wood before, but not necessarily near London; just as often in

our dreams of parts unknown.

But it is not necessary to turn to pure romance in order to

find outlaws some of whom in the tyranny of petty govern¬

ments and municipal feuds, may have been of noble descent

and even kept some noble qualities. As we have seen, the

regions between the Italian cities were infested by adventurers

and exiles of various classes.

English Elements of the Setting. We have considered as ele¬

ments in the background the motive of travel, the obviously

Italian allusions, the non-geographical romantic elements. It

remains to discuss the more definitely English elements. In

these we find not a little support for Ordish’s theory. Country

and every-day life are suggested through allusions chiefly sup¬

plied by the low-comedy characters: the life of the shepherd

and the ways of sheep, fodder, wages, pasture, pound and pin¬

fold;117 the robin redbreast singing on the branch, the cock in

the barnyard,118 the alehouse,119 the weather-cock on the stee¬

ple,120 the melancholy gait of lions in the tower,121 the country

housewife who can milk, brew, sew, knit, wash and scour and

spin.122 And Speed gives us a series of pictures, struck out

boldly in a line apiece, of familiar types in characteristic action :

“to walk alone, like one that hath the pestilence; to sigh, like

a school-boy that hath lost his ABC ; to weep like a young wench

that hath buried her grandam ; to fast, like one that takes diet ;

to watch, like one that fears robbery; to speak puling, like a

beggar at Hallowmas.”123 We have a hint or two of a northern

116 Certain Tragical Discourses of Bandello, translated by Geoffrey Fenton.

Discourse XIII, p. 272.

117 1. i. 72-99.

118 II. i. 21, 28.

110 II. v. 54.

120 II. i. 142.

121 II. i. 29.

122 III. i.

123 II. vii. 19

Rolfe in his edition of The Two Gentlemen of Verona, 1905, p. 138, quotes

Knight on this passage : “The beggar not only spake ‘puling’ at Hallowmas, but

his importunities or his threats were heard at all seasons. The disease of the

country was vagrancy ; and to this deep-rooted evil there were only applied

the surface remedies to which Launee alludes, ‘the stocks’ and ‘the pillory’.”

Wales— The Two Gentlemen of Verona .

107

climate ; the folly of trying to kindle fire with snow,124 “the un¬

certain glory of an April day”.125 And the passage on the brook

need not take us far from home :

The current that with gentle murmur glides,

Thou knowest, being stopped, impatiently doth rage ;

But when his fair course is not hindered,

He makes sweet music on the enamelled stones,

Giving a gentle kiss to every sedge

He overtaketh in his pilgrimage;

And so by many winding nooks he strays

With willing sport to the wild ocean/28

These elements for the most part, however, English in origin

though they undoubtedly are, are not such as definitely to break

an Italian illusion. There were shepherds, housewives, school

boys, beggars, in Italy as well as England; and untravelled

spectators could think of these details in other lands as well as

their own.

Such is the background of the play. Exceedingly little of

this concrete detail can be traced to the sources.127 We have

already quoted a few suggestive passages from Diana Enamo-

mda . To these may be added a few others as calling up fairly

vivid scenes.

Don Felix is sent by his father to the Princess Augusta

Caesarina’s court. Felismena follows him disguised as a man.

The journey takes twenty days.

“I took up mine Inne in a street less frequented with concurse of

people . But midnight being a little past, mine host called

at my chamber doore, and tolde me if I was desirious to heare

some braue musicke, I should arise quickly, and open a window

towards the street.”

She hears Don Felix say to Fabius, the page,

(The rest of the note is also of interest.) Cf. Shakespeare’s England, Oxford,

1916, II, Chapter XXVIII, “Rogues and Vagabonds/’ by Charles Whibley. The

problem of poverty had been aggravated by the dissolution of the monasteries

(p. 486)— “The three-fold object of Elisabeth’s poor-laws was to feed the hungry,

to punish the evildoer, and to exact payment from the unwilling rich.” (p. 490)

— “The severest punishments meted out to the scoundrel, the heaviest toles levied

upon the wealthy, neither cured nor discouraged the crime of vagabondage.” (p.

491.)

124 II. vii. 19.

125 1, iii. 85.

126 II. vii. 25-32.

121 As far as we know what these are. Here as elsewhere there is, of course,

the possibility of an intervening play.

108 Wisconsin Academy of Sciences , Arts, and Letters .

“ ‘Now it is time, my masters, bicause the Lady is in her gallerie

ouer her garden taking the fresh aire of the coole night.’ He had

no sooner saide so, but they began to winde three Cornets and a

Sackbot, with such skill and sweetnesse, that it seemed celestiall

musicke ....

“After they had first, with a concert of musicke, sung this song,

two plaied, the one upon a Lute, the other upon a siluer sounding

Harpe, being accompanied with the sweete voice of my Don

Felix ....

“The sonnet being ended, they paused awhile, playing on fower

Lutes togither, and on a paire of Virginals, with such heauenly

melodie, that the whole worlde (I thinke) could not affoord sweet¬

er musick to the eare ....

“About dawning of the day the musicke ended, and I did what

I could to espie out my Don Felix, but the darknes of the night

was mine enimie therein.”

The next day she goes to the palace.

“Comminge therefore to a faire broad court before the pallace

gate, I viewed the windowe and galleries, where I sawe such store

of blazing beauties, and gallant Ladies, that I am not able now to

recount, nor then to do any more but wonder at their graces,

their gorgeous attyre, their jewels, their brauve fashions of ap-

parell, and ornaments wherewith they were so richly set up.

Up and downe this place, before the windowes roade many lords

and braue gentlemen in rich and sumptuous habits, and mounted

upon proud Jennets, euery one casting his eie to that part where

his thoughts were secretly placed.”

As she stands at the palace gate, she sees Don Felix. A highly

detailed passage follows:

. I sawe him comming along with a great traine of fol¬

lowers attending on his person, all of them being brauely ap¬

parelled in a liuerie of watchet silke, garded with yellow veluet,

and stitched on either side with threedes of twisted siluer, wear¬

ing likewise blew, yellow, and white feathers in their hats. But

my Lorde Don Felix had on a paire of ash colour hose, embrod-

ered and drawen foorth with watchet tissue; his dublet was of

white saten, embrodered with knots of golde, and likewise an em-

brodered jerkin of the same coloured veluet; and his short cape

cloke was of blacke veluet, edged with gold lace, and hung full of

buttons of pearle and gold, and lined with razed watchet satten:

by his side he ware, at a paire of embrodered hangers, a rapier

and dagger, with engrauen hilts and pommell of beaten golde. On

his head, a hat beset full of golden stars, in the mids of euerie

which a rich orient pearle was enchased, and his feather was

likewise blew, yellow, and white. Mounted he came upon a faire

Wales — The Two Gentlemen of Verona .

109

dapple graie Jennet, with a rich furniture of blew, embrodered

with golde and seede pearle.128

Don Felix dismounts and goes up “a paire of staires into the

chamber of presence”.

Felismena learns from Fabius that Don Felix serves Celia

“and therefore weares and giues for his liuerie an azure blew,

which is the colour of the skie, and white and yellow, which

are the colours of his Lady and mistresse”.

With the exception of the above passages the background of

the story is vague and conventional.

Munday’s Two Italian Gentlemen , which presents some resem¬

blances to The Two Gentlemen of Verona in situation, has street

scene, window, lute, and letter. But, as is true of Diana , the

detailed passages have little in common with our play, though

Medusa's box of enchantments — an egg of a black hen, a quill

plucked from a crow, the heart of a black cat, the blood of a

bat, a goat’s brain, the liver of a purple dove, a cock’s eye,

capon’s spur, the left leg of a quail, goose’s bill, gander’s tongue,

mounting eagle’s tail, etc. — bring to mind the ingredients of

the witches’ caldron in Macbeth . Riche’s Apolonius and Silla,

in which the resemblances of situation are stronger, has very

little concrete detail of setting. It does present, however, a

general background of Mediterranean travel.

In our survey of the elements of the setting of The Two Gen¬

tlemen of Verona , we have glanced at several theories: that of

Sarrazin, that Shakespeare is using Italian detail to fill in a

realistic Italian background, that suggested by Miss Spens, that

he uses Italy for an effect, not of realism, but of indefinite dis¬

tance and romance, that of Ordish, that the Italian setting is a

thin disguise for pictures of English life.129 The various theories

may be less conflicting than they seem. At all events, degrees

and distinctions are to be observed, from play to play, in the

nature and use of the foreign scene. Before attempting to draw

further conclusions, however, let us turn to the characters and

setting of the present play, which the background is intended

to throw into relief.

128 Hazlitt, Shakespeare’s library. Vol. I, p. 290.

128 Ordish, Shakespeare’s London, p. 161. “Ephesus, perhaps, stood for Lon¬

don, and Syracuse for Antwerp.” Quiller-Couch and Wilson apparently tend to

Ordish’s view. Op. cit. 1921, pp. ix, x. Cf. the similar view stated by Jacobs, in

the introduction to Painter’s Palace of Pleasure, 1890.

110 Wisconsin Academy of Sciences , Arts, and Letters .

Ill

Characters and Action

The story seems to be Spanish and Italian in origin. Among

the possible sources are Montemayor’s Diana Enamorda, Riche’s

History of Apolonius and Silla,130 Brooke’s Romeus and Juliet,

and Munday’s The Two Italian Gentlemen .

Two points are brought out by the study of the sources: (a)

the stock nature of many of the incidents,131 (b) the relation

of the play to later plays, and especially to Romeo and Juliet

and to Twelfth Night . The resemblance to later plays,132 again,

is chiefly interesting as bringing out the difference in treat¬

ment, the Two Gentlemen of Verona having less depth of real¬

ism, less clearness of characterization, and less unity of effect.

To judge by the relation of the play to its possible sources

and to the later plays similar in source, the elements that chiefly

interested Shakespeare were (a) the adventures of travel,133

(b) the story of intrigue, disguise,134 and escape, (c) the pathos

of Julia’s situation, a close parallel to Viola’s, (d) a friendship

130 For an admirable summary of the parallels between Two Gentlemen of

Verona and these sources see Introduction to the edition of the play in the Arden

Shakespeare, ed. R. Warwick Bond, London, 1906.

131 Cornelia C. Coulter, “The Plautine Tradition in Shakespeare,” Journal of

English and German Philology , v. xix, p. 78.

“The pater familias of a Latin comedy was useful chiefly because he furnished

(albeit unwillingly) the necessary funds for his son’s romance. Sometimes the

memory of his own wild oats made him tolerant of the young man’s misdemean¬

ors ; more often he took an uncompromising stand as a censor of morals and

laudator temporis acti. In four plays of Plautus ( Asinaria , Bacchides, Casina,

Mercator) the old men cast lustful eyes at their sons’ mistresses; in the Aulu-

laria, the rich old bachelor Megadorus makes an honorable request for the hand

of the miser’s daughter, without dowry. Italian dramatists took over these

figures, ' and, by exaggerating their ridiculous aspects, developed the Pantaloon

and the Pedant or Doctor, the former, as a rule, the father of hero or heroine,

and the latter often a suitor for the lady’s hand. Both were unattractive fig¬

ures, stupid, avaricious, amorous, and easily duped by the young people in the

play. Shakespeare’s treatment is much more kindly, but we can still recognize

the traits of the classical senex in the stern decrees of Antonio ( The Two Gen¬

tlemen of Verona, I. 3.) and Baptista ( Taming of the Shrew I. 1.), in Capu-

let’s reminiscences of by-gone days ( Romeo and Juliet I. 5.), and in the ‘wise

saws’ of Polonius to Laertes ( Hamlet I. 3.). Silvia’s father traps Valentine by

the story of a coy lady whom his ‘aged eloquence’ had failed to move ( Two

Gentlemen of Verona III. 1. 76-136), and ‘old Signior Gremio’ offers plate and

gold, Tyrian tapestry and arras counterpoints as dower for the fair Bianca

( Taming of the Shrew II. i. 347-364).”

i82 por a summary of resemblances to later plays see Furnivall’s introduction

to “The Two Gentlemen of Verona” in The Century Shakespeare , 1908. See also

the Edition of Quiller-Couch and Wilson, 1921, pp. ix-xv.

133 Cf. Merchant of Venice.

134 See Freeburg, Disguise Plots in Elizabethan Drama, New York, 1915.

Wales — The Two Gentlemen of Verona .

Ill

complicated by differences of temperament,135 (e) an enchanted

place and a happy ending.

Professor Campbell has shown, however, that even these are

so essentially stock appeals to popular interest that we cannot

assume anything more than that Shakespeare knew how to use

them as such. Yet if, in the comedies, as Professor Stoll con¬

tends,136 a swift theatrical appeal, without deeper dramatic

implications, is the norm, Shakespeare often departs widely

from it. Given a story of a certain degree of remoteness from

real life, he sometimes gets a detail out of focus, brings it too

close to the camera, over-realizes it. Shakespeare is stronger

in his sense both of tragic unity and of romantic unity than

in his sense of comic unity. In Love's Labour's Lost , in The

Comedy of Errors he encounters this difficulty.

In Love's Labour's Lost the scene suddenly “begins to cloud”.

The players doff their fantastic disguises, their artificial speech,

and their courtly self-deception, and step forth human beings

who feel pain as we do.

Honest plain words best pierce the ear of grief.m

The bubble of the comic illusion has broken when the King is

told to

go with speed

To some forlorn and naked hermitage

Remote from all the pleasures of the world,188

and Biron to

Visit the speechless sick and still converse

With groaning wretches.139

Biron replies

To move wild laughter in the throat of death?

It cannot be; it is impossible;

Mirth cannot move a soul in agony.140

386 See Quiller-Couch’s introduction. The Two Gentlemen of Verona, New York,

1921.

138 Cf. on this point the general position of “the skeptics”. See Karl Young,

“The Shakespeare Skeptics”, North American Review, March, 1922. Also Mr.

Young’s review of Schftcking, Philological Quarterly, I, 3, July 1922, pp. 228-234.

See especially note 5, page 229 (bibliography).

337 V. ii, 763.

388 Love’s Labour’s Lost, V. ii, 804-6.

383 Ibid. 861-2.

340 Ibid. 865-7. , .

112 Wisconsin Academy of Sciences , Arts , and Letters .

Yet because the play must end as a comedy, there is an attempt

at the end, not altogether unsuccessful, to persuade us that the

grief of the princess-natural and expected rather than tragic

— is not one of the storms of life, but slips over the grass of

the park like the shadow of a drifting cloud on a summer after¬

noon. Nevertheless we have been strangely reminded that there

are such things as storms and winter.

In The Comedy of Errors, similar realities break through,

out of keeping with what Miss Spens has called “the hard,

unlovely type” of the Latin comedy.141 The grief of Aegeon

acquires a certain vitality, likewise the romance of Antipholus

and Luciana, the religious atmosphere of the abbey, the reunion

of the family. In the Merchant of Venice and Much Ado About

Nothing tragedy comes through in a way disastrous to the

comic unity.142

In The Two Gentlemen of Verona are there any characters

or situations which tend to emerge into stronger reality than

the general plane of the action leads us to expect?143

The character of Julia changes somewhat and becomes more

real as the story goes on. In the first act she is coy, self-

centered, unreasonable, and unrefined. The scene has a fairly

close parallel in Yonge. One has only to compare it cursorily

with a later parallel, Portia's discussion of her suitors with

Nerissa, to see how inferior to the latter it is in dignity. In

Act II, Scene vii, there is more liveliness and sincerity as well

as more lyric beauty in Julia's speeches. In Act IV, Scene ii,

Shakespeare has grown much more interested in Julia, and her

situation. Her lines are brief and strong; and in the poignant

note of “I would always have one play but one thing,”144 it is al¬

most the voice of Viola that we hear. In Act IV, Scene ii, Julia's

experience is still further developed. Silvia's character is ex¬

ternally presented; we know little of her experience from the

inside; but Julia, like Viola, utters words that in their double

meaning have an aspect of soliloquy.145

141 Shakespeare and Tradition, Oxford, 1916, p. 7.

142 Commented on by many critics. See, for example, Heinrich Bulhaupt, Vario¬

rum edition of Much Ado About Nothing, p. 379.

143 Quiller-Couch and Wilson account for discrepancies by the interesting theory

that the play does not stand as Shakespeare wrote it but has been “tightened”

and revised for acting purposes by another hand. See pp. 115-6 below.

144 IV. ii. 72.

145 IV. iv.

Wales — The Two Gentlemen of Verona.

113

In the course of the play there is some attempt to show the

mind of Proteus, also, from the inside. There are some solilo¬

quies — one on his love affair with Julia — in which he ex¬

presses contempt for himself for wasting his time;146 one ex¬

plaining the lack of courage which results in his going

abroad;147 one in which he reveals his infatuation for Silvia

and tries to account for the complete change in his own state

of mind:

She’s fair, and so is Julia that I love, —

That I did love, for now my love is thawed,

Which, like a waxen image ’gainst a tire,

Bears no impression of the thing it was.

Methinks my love to Valentine is cold,

And that I love him not as I was wont:

0, but I love his lady too too much

And that’s the reason I love him so little.148

There is a fourth soliloquy of about forty lines, which is ap¬

parently meant to display some degree of moral struggle. It

sounds like a forlorn effort on Shakespeare’s part to make a

case for Proteus. Proteus realizes the ignominy of what he is

doing :

To leave my Julia, shall I be forsworn;

To love fair Silvia, shall I be forsworn;

To wrong my friend, I shall be much forsworn.149

But he defends his course by a two-fold argument: (1) love

is all-powerful:

At first I did adore a twinkling star,

But now I worship a celestial sun,

the celestial sun being Silvia the great lady, (2) self-realiza¬

tion is the first law of life :

Julia I lose, and Valentine I lose:

If I keep them, I needs must lose myself;

If I lose them, thus find I, by their loss,

For Valentine, myself; for Julia, Silvia.

I to myself am dearer than a friend,

For love is still most precious in itself;

146 I. i. 63-69.

14TI. iii. 78-87.

148 II. iv. 191-214.

148 II. vi. 1-3.

114 Wisconsin Academy of Sciences , Arts, and Letters.

I cannot now prove constant to myself,

Without some treachery used to Valentine.550

Proteus then carries out his plot with the relentlessness of a

man hypnotized by a motive that he never again stops to sur¬

vey, since he knows that it is evil.

Except in a passage at the opening of Act IV, intended chief¬

ly for description and atmosphere, there is no such attempt

to present directly the working of Valentine's mind. Yet he

seems much more alive. Indeed it is because he is more real

that he needs less explanation. He is reflective,151 something

of an idler in his way,152 yet with plenty of energy, lacking the

alertness and finesse of a man of the world, engagingly slow

to understand what is to his advantage,153 blunt of speech, im¬

petuous of temper, capable of vigorous action but incapable of

successful intrigue,154 without experience of treachery and easi¬

ly duped by it, of resourceful imagination, adventurous, yet de¬

manding little of life, and adaptable to adverse circum¬

stances.155

His praise of Proteus shows what he thinks, with more or

less injustice to himself, to be his own defects:

I know him as myself; for from our infancy

We have conversed and spent our hours together;

And though myself have been an idle truant,

Omitting the sweet benefit of time

To clothe mine age with angel-like perfection,

Yet hath Sir Proteus, for that’s his name,

Made use and fair advantage of his days;

His years but young, but his experience old ;

His head unmellow’d, but his judgment ripe:

And, in a word (for far behind his worth

Come all the praises that I now bestow),

He is complete in feature and in mind,

With all good grace to grace a gentleman.158

Because of this comparatively clear characterization of Val¬

entine, we, as we read the play, stand in his shoes in the last

iso This long soliloquy seems to me to be an attempt at psychology. The earlier

soliloquies are no doubt, as Professor Stoll would hold, chiefly external explana¬

tion to the audience.

151 V. iv. 1.

162 II. iv. 64.

“3 II. iv.

III. i.

iso v. iv.

168 II. iv. 62-74.

Wales- — The Two Gentlemen of Verona .

115

act. Through his eyes we are staring at Proteus, trying vainly

to understand what manner of man this false friend can be.

In the final scene, however, the dramatic situation is aban¬

doned as a whole. Silvia to all intents and purposes is forgot¬

ten. If we take the play seriously as it stands, the conclusion

of the whole matter seems to be: “I forgive you; I take you

back — but I don’t understand.” Proteus is put to a final test,

“Here, take the girl if you want her.” But how he responds

to this test we do not know, since Julia at this point re-enters

the action. If we think of this scene as falling below what

we are to expect, instead of thinking of the rest of the play as

rising above it, the denouement is puzzling.

Professor Stoll, and other skeptical critics,157 maintain that

Shakespeare, like his audience, cared more for story than for

drama in the true sense, and easily threw ftside the problem

of character in favor of a mechanically happy ending. It

would be absurd to deny that there is truth in this view.

Shakespeare knew this ending to be theatrically acceptable to

his audience, and was satisfied to give it to them. It would

be easier to take this as the whole truth, if it were not that

again and again in the plays, — - in almost any scene of every

play — Shakespeare shows that, whatever the professional nec¬

essities under which he worked, his own spontaneous interest

is easily captivated by character as character and by the genu¬

ine dramatic values of a situation. Thus the real puzzle of

The Two Gentlemen of Verona is not alone in the unmotivated

happy ending but in the combination of this with more strongly

motivated earlier passages. We must accept the simple expla¬

nation as far as it goes; but in Shakespeare we are dealing

with a personality as well as a condition, and it is well to re¬

member that beneath and beyond the simple fact are more

complex possibilities. Speculation about these may be harm¬

less, may even be wholesome, as protecting us from any temp¬

tation to dogmatism and rigidity, provided and only provided

we never forget that it is speculation merely.

And here the theory of Quiller-Couch and Wilson may help

us out. “It may be then that Shakespeare invented a solution

which at the first performance was found to be ineffective;

that the final scene was partly rewritten — not by Shakespeare

157 See footnote 136, p. Ill above.

116 Wisconsin Academy of Sciences , Arts , and Letters.

— and given its crude and conventional coup de theatre; that

in this mutilated form it remained on the play-copy; and that

so it reached the printer.158 But it will be asked, if we omit

All that was mine in Silvia I give thee

how do we account for Julia’s swoon? Our own answer is that

we do not try to account for it; our hypothesis being that the

swoon and the couplet together are ‘other man’s work’; and

that Shakespeare had another denouement which possibly

proved ineffective on the stage, and that the one we have is a

stage-adapter’s substitute.”159

Sir Arthur Quiller-Couch wisely refrains from telling us

what Shakespeare’s other denouement was. Our guess would

be (a guess, pure and simple) — an effort by Valentine to probe

the mind of Proteus, resulting in a dialogue more interesting

to the dramatist himself than to the Elizabethan play-goer.

Before we leave the subject let us turn once more to the

relation between the present play and Twelfth Night. A curi¬

ous episode in the latter play is that of Antonio and Sebastian.

Antonio is an older man who bears to Sebastian something the

relation of Adam to Orlando, and of the Antonio of the Mer¬

chant of Venice to Bassanio. In As You Like It and The Mer¬

chant of Venice , the younger man proves worthy of the fidelity

of the elder. In Twelfth Night , likewise, he proves worthy.

But there is a scene of almost tragic vividness,160 in which An¬

tonio, thinking himself deceived, takes an attitude much more

mature than Valentine’s:

Will you deny me now?

Is’t possible that my deserts to you

Can lack persuasion? Do not tempt my misery,

Lest that it make me so unsound a man

As to upbraid you with those kindnesses

That I have done for you .

But O how vile an idol proves this god!

Thou hast, Sebastian, done good feature shame.

In nature there’s no blemish but the mind;

None can be call’d deformed but the unkind.

Virtue is beauty .

158 See introduction to Two Gentlemen of Verona, edited by Quiller-Couch and

Wilson, New York, 1921, p. xvi.

159 Ibid. p. xviii.

160 Miss Margaret Ashdown of Westfield College, the University of London, com¬

mented to me (I think in this connection) on the poignancy of this scene.

Wales — The Two Gentlemen of Verona.

117

And again when he has had time to digest his wrongs, Antonio

utters words very unlike Valentine's too easy re-instatement of

the wrong-doer.

A witchcraft drew me hither :

That most ungrateful boy, there by your side,

From the rude sea’s enrag’d and foamy mouth

Did I redeem; a wreck past hope he was.

His life I gave him, and did thereto add

My love, without retention or restraint,

All his in dedication . .

. His false cunning

Not meaning to partake with me in danger,

Taught him to face me out of his acquaintance,

And grew a twenty years removed thing

While one could wink.

Matters are cleared up for Antonio. There are two Sebastians.

Antonio. How have you made division of yourself?

An apple cleft in two is not more twain

Than these two creatures. Which is Sebastian?

Sebastian . A spirit I am indeed

But am in that dimension grossly clad

Which from the womb I did participate.

All is resolved by the simple device of mistaken identity. In

a figure, perhaps, Valentine gets out of his difficulty in some¬

what the same way. Who has not succeeded in reinstating his

friend by reflecting on the notion of man’s Protean double na¬

ture? He did wrong, we say to ourselves, through yielding

to the hypnosis of an evil idea ; his real self, thoroughly aroused,

will repudiate his action, and will learn from the bitterness of

repentance not to lapse from its own integrity again. In The

Two Gentlemen of Verona, as it stands, the injured man’s re¬

sentment, and his forgiveness, are more hastily presented than

any dramatic condensation allows — so hastily, indeed, as to

seem ironical. But in the later play where the situation of the

betrayed friend is only an incident, and an illusion at that,

Shakespeare shows his power to make it almost tragically

real.161

161 Cf. on the subject of friendship vs. love, Max Forster, (reviewing Hans

Kliem, on Sentimentale Freundschaft in der Shakespeare-Epoche, Jena, 1915),

S. G. Jahrbuch, LI (1915), p. 261.

“Dieser schwarmerische Freundschaftkultus stammt nattirlich aus der Antike,

wobei auch diesmal wieder, wie bei sonstiger Renaissancekultur, Frankreich

(namentlich Montaigne) eine gewichtige Vermittlerrolle zufallen mag. Antik

sind vor allem die oft in der damaligen englischen Literatur wiederkehrenden

118 Wisconsin Academy of Sciences , Arts, and Letters .

Thus, however they account for it, the critics feel that the

denouement is below the level of Shakespeare’s usual treatment,

below the level, even, of this play as a whole. Yet there may

be an advantage in treating the weakest point as our point of

departure, in approaching the dramatic problem of the play

as it were from the direction of its lower level. From this

standpoint what we have to try to understand is not the weak

points of the play but its strong points. Wherein lie the out¬

standing differences between this play and its cruder proto¬

types ?

One conspicuous difference lies in the greater variety of its

scene and background. Professor Campbell stresses the point

that “the scene of the action in an Italian comedy, both learned

and professional, was a public place”. In our play both forest

and court scenes are directly presented. Then there is a

strengthening in our sense of the persons, though, as we have

seen, this gain is uneven and varies from incident to incident.

There is great strengthening of the spirit of romance and no

less of the sense of reality; and through these there is a deep¬

ening of the whole dramatic and poetic appeal.

IV

The Setting and the Total Effect

Now does this examination help us at all in framing a theory

of Shakespeare’s use of background in this play? How is the

story related to the background, and does the background help

us to interpret the story?

Before answering this question, let us again turn for a mo¬

ment to that very different type of play, The Comedy of Er¬

rors . It is, of course, not a romantic comedy, but a farce of

the Latin type; the situations are too wildly absurd to admit

of being made probable by any artistic device whatsoever. It

is meant that we should confine ourselves to the realm of farce,

and not ask why or how such things could happen. We must

not reflect upon their strangeness ; we must only laugh at their

Anschauungen, dasz Mannerfreundschaft Hoher sttinde als Frauenliebe und alle

Bande der Familie (Bacon, Spenser, Th. Browne), dasz der Freund ein anderes

Selbst (another himself) sei und dasz echte Freundschaft ‘eine Seele in zwei

Korpern’ bedeute . . . Dasz dabei manche psychologische Unmoglichkeit mit un-

terlauft, verschlug dem damaligen Leser wenig . . . FGr ein Stiick, wie ‘Die

beiden Edelleute von Verona* gewinnt man erst das richtige Verstandnis, wenn

man es in Lichte dieses Freundschaftkultus betrachtet”.

Wales — The Two Gentlemen of Verona .

119

absurdity. Yet oddly enough, reflection upon their strange¬

ness is exactly what Shakespeare demands of us repeatedly in

the following expressions:

They say this town is full of cozenage;

As, nimble jugglers that deceive the eye,

Dark-working sorcerers that change the mind,

Soul-killing witches that deform the body.

This is the fairy land. O spite of spites!

We talk with goblins, owls, and elvish sprites:

If we obey them not, this will ensue,

They’ll suck our breath, or pinch us black and blue.

Against my soul’s pure truth, why labour you

To make it wander in an unknown field?

I’ll stop my ears against the mermaid’s song.

Sure, these are but imaginary wiles,

And Lapland sorcerers do inhabit here.

. Witness

That he is borne about invisible.

I think you all have drunk of Circe’s cup.

Why this is strange — Go call the abbess hither.

I think you all are mated or stark mad.

These phrases and others indicate that reality has begun to

break through the farcical unity so that we are in danger of

testing events by probability; hence the illusion must be recap¬

tured in some other way — in fact by the method of poetry.

In other words, Shakespeare has even here begun an experi¬

ment in romantic unity.162 The same method seems to be car¬

ried further, though not to a wholly successful issue, in The

Two Gentlemen of Verona .

There may thus be a relation between the wood, with its

effect of enchantment, commented on by Miss Spens, and the

162 The Menechmi supplies a faint suggestion of the method :

Either she is a witch, or else she hath dwelt there and knew ye there.

Fie, awake Menechmus, awake ; ye oversleepe yourselfe.

They vex me with strange speeches.

They try to persuade Menechmus that he is mad.

They say I am mad.

This same is either some notable cousening Jugler, or else it is your

brother whom we seeke.

120 Wisconsin Academy of Sciences , Arts, and Letters .

improbability of the story which has to be rendered dramatical¬

ly credible. Shakespeare is employing what we may call the

device of the enchanted place. This device has, of course, been

recognized, at least in implication, by commentators on A Mid¬

summer Night's Dream, The Tempest, and other plays; it is

not confined to Shakespeare, and comes indeed from a univer¬

sal artistic law governing the relation of any event to the at¬

mosphere in which it is represented as taking place. None the

less it is with Shakespeare perhaps even more elaborate than

has been recognized. The mechanism of the device is best

studied in A Midsummer Night's Dream.1™ Suffice it for the

present to say that in A Midsummer Night's Dream, by the use

of the atmosphere of a place, a highly complex action, of hetero¬

geneous elements, is removed from life, set back, brought into

focus, simplified in motivation, and given an illusory credibility

such as could have been obtained in no other way. In The Two

Gentlemen of Verona , Valentine’s forgiveness of Proteus as it

stands164 remains a difficulty which even this magic chemical

fails to dissolve. But it does succeed for the disentanglements

of the love complications and the regeneration of the outlaws.

Now does this device of the enchanted place subserve the

Italian setting, or vice versa? Shall we accept the theory of

Sarrazin, that Shakespeare is aiming at a picture of Italy, or

that of Miss Spens, that he is aiming chiefly at romantic dis¬

tance? The dramatic needs of this play seem to throw the

emphasis with the latter idea. Yet each element contributes to

the other; we have a distant and romantic scene but one dis¬

tinctly Italian.

What, now, is the artistic relation of the English detail to

this Italian romantic background ?

Miss Spens has an illuminating passage on Lyly’s influence

on Shakespeare:

Now Lyly’s plays are essentially masques; that is to say, they

are representative of actual incidents of the time at which they

were performed, translated by the language of allegory and sym¬

bolism into a more radiant plane of existence . .

It was not the business of the poet to create, but to flood the

given facts with a golden light of poetry, which should show all

183 Of which I hope soon to complete a study.

184 But see p. 116 above and footnotes.

Wales— -The Two Gentlemen of Verona.

121

the fairer connections. The masque is, in fact, the direct antithe¬

sis of satire. Now the great comedy of all other literature belongs

to the satirical family-— it exaggerates whatever is ugly in human

action in order to make it ridiculous, and Ben Jonson’s comedy

shows that without Lyly and Shakespeare English comedy would

probably have followed the same lines. The scene of Shake¬

speare’s comedy is laid in a golden world, and the suggestion of

that ethereal atmosphere came from Lyly. Lyly failed because he

does not at the bottom of his heart believe in this golden world.

Shakespeare’s task was to give it truth.165

Taking this sentence of Miss Spens’, “Shakespeare’s task was

to give it truth”, as a text, let us see what are the factors of

Shakespeare’s method as we are able to discern it already in

The Two Gentlemen of Verona.

And even when we use the phrase “Shakespeare’s method”,

we must give ourselves a warning. It is characteristic of

Shakespeare that he never carried any particular mode far.

The moment we begin to define his method we meet trouble;

for he was too wholly a creator to have method in a deliberate

sense. His conscious intellectual process was too far merged

in something more unconscious and vital. Professor Basker-

ville says of Ben Jonson: “It was the life of the man to be

in the midst of things. Let a type of the drama like the masque

become popular and he is almost certain to adopt it and exert

all his powers to excel in it”.166 Now Shakespeare was also

“in the midst of things”, — and he too adopted popular types.167

But he was probably not chiefly concerned with excelling in any

particular type, for he was concerned with something more im¬

portant, with using the type as a means to the end — to delight

his audience, and to find outlet for his own peculiar power of

holding the mirror up to nature. So when we talk of his meth¬

od, we mean only the way he did things, and not the way he

meant to or self-consciously set out to do things. Shakespeare

has in mind not so much the author’s point of view as (a) the

point of view of the actor or group of actors, (b) the point of

view of the spectators. He did not think of personal fame, ap¬

parently, or of showing off learning or cleverness.

Shakespeare’s method, then, if we use the word in the sense

of a partly unconscious process, was to furnish his golden

165 Shakespeare and Tradition, Oxford, 1916, p. 8.

168 C. R. Baskerville, English Elements in Jonson’s Early Comedy , 1911, p. 4.

167 Cf. A. H. Thorndike, The Influences of Beaumont and Fletcher on Shake¬

speare , Worcester, Mass., 1901.

122 Wisconsin Academy of Sciences , Arts , and Letters .

world with the detail of every-day life, not in order to create

an illusion of every-day life, but in order to give reality to the

golden world. We have seen in the present play how thriftily

he uses the circumstantial bits of romantic machinery. Already

the scene has far more substance than a scene as Lyly pre¬

sents it. Yet the detail of the play is thin and scanty com¬

pared with that of any other play of Shakespeare. How elabo¬

rately furnished is the forest of Arden, the wood near Athens,

the Island of the Tempest! The fairy life of the Midsummer

Night's Dream is real because made of the stuff of our hum¬

drum routine, and not only is a place or situation given sub¬

stance in this way, but the mind of a character is similarly

realized. Oberon, the Ghost, Caliban, are alive to us because

they incidentally reveal two elements in their consciousness:

(a) the little objects in the foreground of their attention, (b)

the deposit of concrete recollection of the past. A modern poet

has deliberately made this method characteristically his own.

Fra Lippo Lippi, in a way convincingly incidental to the play

of ideas going on in the area of his immediate consciousness,

reveals sharply detailed pictures from his memories of street

and cloister. Bishop Blougram, conscious of every trivial thing

in his surroundings, even of Gigadibs the literary man,

Who played with spoons, explored his plate’s design,

And ranged the olive stones about its edge,

pours out the riches of a long-stored mind. And in Andrea del

Sarto , “that cartoon, the second from the door”, is only a mod¬

ern and self-conscious version of “the picture that is hanging

in your chamber”,168 of which Proteus, from his place among

the trees, is evidently able to get a glimpse in the candle-light

behind Silvia's head. In this play Shakespeare's most striking

use of detail for characterization is in the treatment of Launce.

He knows all the odds and ends that Launce would happen to

have in his mind: cats and dogs and geese, puddings and

trenchers, Jews, and shoes, and pebble stones, the market-place

and the pillory.

Ordish in his valuable book on Shakespeare's London , dis¬

cusses the vast amount of English concrete detail which occurs

in those plays whose scene is nominally outside England.169

las Two Gentlemen of Verona : Act IV, Sc. ii.

use Ordish, Shakespeare’s London, 1897. See especially Chapter IV.

Wales — The Two Gentlemen of Verona . 123

We have already quoted a passage in which he states the belief

that “beneath the masquerade of foreign manners lay tacitly

the familiar scenes of England and of London”. He speaks of

this as a “device”170 taken for granted, a “disguise” to be pene¬

trated, an “illusion” which “gave zest to the satire”,171 a “mas¬

querade of the remote”.172 In this view there is much truth

that it would be folly to dispute. Yet it is doubtful whether

either Shakespeare or his audience recognized anything two¬

fold in the method. Probably the audience did not sharply dis¬

tinguish English and foreign elements. They probably did not

think of parallel situations, one standing for the other. Local

hits amused and delighted them ; but these were, to a great ex¬

tent, extraneous, inserted just as they are by travelling players

to-day, and not of the essence of the play itself. The story as

story, the people as people, were, in general, the first consid¬

eration of playwright, actor, and spectator.

But while the topical allusions were not essential to the play,

the local details were essential. They are present not primarily

for a satirical purpose — though sometimes Shakespeare is sa¬

tirical — but for a higher creative purpose — namely, to make

the ideal, or in other words, the foreign, the distant, the ro¬

mantic element, become real. When Shakespeare laid the scene

of a play in Italy, he meant the audience to think of Italy. He

himself imagined Italy as far as his knowledge of Italy en¬

abled him to do so. When his knowledge of Italy gave out, he

supplied English detail, not because it was English but because

it was real. He put his Italian story into an Italian setting

so far as his mind happened to have the materials of such a

setting. Possibly when he found something not clear to his

imagination, something that for purposes of realizing the ac¬

tion to himself he needed to get clear, he may have looked up

or enquired about points of Italian geography, history, or con¬

temporary manners. But if he did this at all, it was merely

for the purpose of realization, and not for the purpose of accu¬

racy in the abstract. Thus we are concerned in the plays with

Shakespeare's imaginative Italian equipment, not his scientific

Italian knowledge. Probably his imaginative equipment in

Italian knowledge was greater than that of his audience, for

170 Ibid. p. 14Q.

171 Ibid. p. 211.

172 Ibid. p. 212.

124 Wisconsin Academy of Sciences , Arts, and Letters .

he was a man that knowledge (of any human interest, that is)

had a tendency to stick to. Ben Jonson liked to show more

knowledge than his audience possessed. Shakespeare was too

sensitive to the spectator's point of view to do that if he could

help it. As a rule the color Shakespeare uses is that familiar

not only to himself but to his spectators, part of their imagina¬

tive equipment. Hence a great proportion of detail necessary

in realizing a situation will have to be English. And his meth¬

od of using it implied three factors, none of them exactly de¬

liberate: (1) spontaneous observation, (2) subconscious mem¬

ory, (3) spontaneous selection.

Now at the bottom of every very good mind there is a little

machine — something like a dictagraph — automatic, accu¬

rate, that goes on noting down every sort of impression and

talking to itself in a subterraneous way about these impres¬

sions, — very steadily (even when one is not watching it)

sorting, arranging, interpreting, storing away. Has one not

heard it say, as it made some unusual and unintelligible record,

“Now what was that? What was that?" like an automatic

“line's busy" machine? Something turns out, let us say, as we

did not expect it to turn out. Perhaps a personality proves

quite other than the theory we had made of it. And yet we

find — to our surprise — that we are not surprised. Why?

Because our memory has disclosed its subconscious records.

We remember now the points at which we were puzzled, the

snag that set the little mechanism purring, “Now what was

that? What was that?" The creator — the literary creator

more especially — has a superlatively good subconscious re¬

corder. He may be rather absent-minded on the conscious side ;

for practical purposes he perhaps does not notice things as

he ought to do; but sometime after an experience, when he is

ready to use it for a creative purpose, he finds his material

available, every detail sharpening under the gaze of recollec¬

tion, like (to change the figure) a photograph developing in the

dark room of the subconscious mind.

Shakespeare was accurate also in his selection of the records

of memory for artistic use. One more illustration : some years

ago there was a popular kind of exhibition in America called

a cylorama. One entered a circular building, climbed a wind¬

ing stair and found oneself in a tower apparently overlooking

the circle of a vast plain. The mind knew, of course, that a

Wales— -The Two Gentlemen of Verona .

125

brick wall was somewhere within a few feet of the handrail

— that the far horizon was an impossibility; yet it seemed

real. How was it done? By a combination of painted canvas

in the background, actual objects near at hand. Here is a

well sweep; the rope and bucket are a real rope and bucket;

the well, perhaps, is painted. To any ordinary uncritical eye

the illusion was perfect and unbreakable. A childish spectator

was acutely irritated by its perfection, wanted, quite desperate¬

ly, to hurl a pebble into that deceptive distance, see it strike

against a canvas wall, disclose where the painted scene began.

Just so has Shakespeare baffled our senses, by his distant hori¬

zons and his details of every-day life, his magical combination

of the actual and the imaginary. This is not to say, of course,

that Shakespeare invented the method, only that he was its

supreme master. All artists have used it in one degree or an¬

other in all periods. We find it already in the miracle plays —

where in the story of the first Christmas, the shepherds bring

to the Child gifts of toys and mittens, real woollen mittens

worn on English hills.

We perhaps do not realize how far our sense of the persons

depends on this concreteness of setting. In the details of their

surroundings lies much of the substantiality of their experi¬

ences to themselves and much of their own substantiality to the

reader. As we have seen, the sense of romance is very largely

a matter of an atmosphere subtly created through the manipu¬

lation of detail. And this romance is itself vitalized, made

more powerfully romantic by vividness, individualization, color,

and perspective. Similarly the illusion of reality has been at

critical moments protected by the veil of romance, by a skill¬

fully woven place enchantment that has tended to deepen the

dramatic and poetic appeal of the whole.

COWLEY AS A MAN OF LETTERS

Ruth Wallerstein

Every thoughtful reader who goes even a little way into

Cowley will become increasingly aware of two dominant im¬

pressions forming in his own mind. He will feel a massive¬

ness and intensity of poetry, latent almost everywhere, but not

often realized by Cowley in actual word and image and design.

Though Cowley rarely wrote poetry as great as the Ode on

Hervey, there is an equally high spirit latent in much of his

poetry, as Milton felt, and as Wordsworth was to feel. The

reader will perceive in the second place, as many critics since

Gosse have pointed out, how far he is moving into the world

of Dryden and Pope. What I have to say, taking its start from

these two points of massiveness and rhetoric, should perhaps

be called a footnote to previous work on Cowley. In the light

of them, I want to stress a new emphasis in the interpretation

of his work and to point out certain aspects of his influence

which deserve special attention. For Cowley is not only a poet

but even more a man of letters. Studied from this side he

illustrates both the type and its function to a remarkable de¬

gree. Moreover, the study of his influence as a man of letters

on later poetry gives us significant material for the study of

aesthetics and the laws of the literary imagination.

A cursory glance at his types of work indicates at once the

range of interests and hospitality of taste that should charac¬

terize a man of letters. To remind the reader briefly, there is

the enormous sweep from the childhood poems and schoolboy

play, which look back to the earlier Elizabethan world, to the

Anacreontics ; from these to the love lyrics of The Mistress ,

and again to the Pindarics, which open up a type of reflective

lyric new to England. There are the exuberant boyish Latin

play, and the half satiric Cutter of Coleman Street, written in

college and revised years later, and the long Latin Libri Plana -

torum, written after he had studied medicine in his middle

years, and amazing for the virtuosity and fluency of their

elegiacs and lyric measures. Then there are the familiar es¬

says, the project for an agricultural college and experimental

128 Wisconsin Academy of Sciences , Arts , and Letters .

laboratory, and finally the notes to Davideis and the Pindarics,

which are remarkable for their considered theory and defini¬

tions, as well as for their appreciations.1

If we look at Cowley's themes, we find the same catholicity

and enterprise. Let me only note in passing that he added two

new themes to the then traditional and current subjects of lyric

poetry, — - namely, critical theory and the new science and

philosophy. In style, he began with Spenser and Fletcher,

wrought under Donne and again under Ben Jonson, garnered

much from the classics. Here too he innovated; he both en¬

larged the scope of irregular verse and set the English pattern

for it; he built up the couplet and gave currency to the Alex¬

andrine.

His experimentation is in extraordinary proportion to his

actual accomplishment. Other poets of robust nature try a

number of modes of poetry and a number of styles before they

find their own ; but we feel through all the shifts of their work

the gradual emergence of one poetic personality, of one poetic

vision, and of one style able to hold the impress of that vision.

Cowley's various endeavors, on the other hand, though we feel

one intelligence behind them, have no such continuity of poetic

personality and do not merge into one whole. Cowley never

found himself. He brought none of his endeavors to sustained

excellence, except the Anacreontics , and they are the simplest

and most obvious of his poems, as well as the ones most readily

translatable from the classics. He almost never has steadily

at his command a style as adequate as that of a hundred much

lesser poets though he has moments of greater style than they.

Even the play of ingenuity which pervades all the surface of

his writing, and which is really so characteristic of the man’s

temper, is not an integral style. Why was this? In truth, in¬

telligence more than poetic fury possesses his work. His intel¬

ligence, which coolly used his poetic feeling as a tool, rather

than becoming itself the instrument of that feeling, turned to

one and to another experiment; and his energy went succes¬

sively into exploring the theory of these new modes, so that he

never subdued and absorbed them into creative feeling. Cow¬

ley lived in a poetic age, and he is caught up in its spirit, but

1 Mr. Nethercot has gathered these notes together in an article called Abraham

Cowley’s Discourse on Style, published in The Review of English Studies, Octo¬

ber 1926, Vol. II, pp. 385-404.

Wallerstein — - Cowley as a Man of Letters .

129

in him intelligence was more active than primary poetic or

philosophic imagination. That was one reason indeed why he

accomplished so much.

To explain more fully, Cowley himself felt that he was too

much hindered by circumstance and by the attendant dissipa¬

tion of spirit to drive his work through to its full promise.

But in the same disturbed age Marvell, working in a completed

tradition, and Davenant, as a transitional poet reaching out to

the new feeling and the French influence, are poets equally

hindered who brought their vision and style to completion.

Cowley attempted more and had something deeper to say than

Davenant, and hence his material was far more difficult to

assimilate and create into new poetic life; this, however, does

not fully explain his incompleteness. In detail he does not show

the quickening, the assurance, the high clear light of the Sev¬

enteenth Century. Dryden is an example of the accomplish¬

ment in the absorbing and perfecting of new modes which can

be achieved by the great man of letters who has also a very

great artistic gift. In truth, Cowley's gift, as this comparison

shows more clearly, was primarily intellectual, and only sec¬

ondarily poetic. I do not mean by this that Cowley had not a

deep and fine imagination, and a sensitive personality. But

the range and scope of one's thought, and its sympathetic partic¬

ipation in life, do not necessarily become an immediate part of

one's efficient poetic and philosophical creative power unless

there be special creative gift. Lacking this, Cowley can only

say mediately and in terms of analysis, sometimes dry, what

the poet says immediately. His Ode on the Death of Mr. Wil¬

liam Hervey, in its later stanzas, is an example of this limi¬

tation. True, the spacious and poignant opening stanzas, un¬

der the whip of perhaps the strongest emotion Cowley ever

sought to express, concentrate thought and feeling in imagery

that is immediate and splendid:

It was a dismal, and a fearful night,

Scarce could the Morn drive on th’ unwilling Light,

When Sleep , Death's Image , left my troubled brest,

By something liker Death possest.

My eyes with Tears did uncommanded flow,

And on my Soul hung the dull weight

Of some Intolerable Fate.

What bell was that? Ah me! Too much I know.

130 Wisconsin Academy of Sciences , Arts, and Letters .

This tide of feeling moves onward into the great traditional

elegiac reflection upon the emptiness of the world and the bar¬

renness of a friend left desolate by the death of a noble man.

The traditional imagery and thought, with all their glow, are

recast into terms of Cowley’s and Hervey’s actual experience;

and in piercing simplicity the realization of grief transcends

Lycidas .

My dearest Friend , would I had dy’d for thee!

Life and this World will henceforth tedious bee.

Nor shall I know hereafter what to do

If once my Griefs prove tedious too.

Silent and sad I walk about all day,

As sullen Ghosts stalk speechless by

Where their hid Treasures ly;

Alas, my Treasure’s gone , why do I stay?

Ye fields of Cambridge , our dear Cambridge , say,

Have ye not seen us walking every day?

Was there a Tree about which did not know

The Love betwixt us two?

Henceforth, ye gentle Trees, for ever fade;

Or your sad branches thicker joyn,

And into darksome shades combine,

Dark as the Grove wherein my Friend is laid.

But as the reflection grows more actual, the movement slows

up, and even though it does not lose its nobility, it passes into

the step by step analysis of prose :

His mirth was the pure Spirits of various Wit,

Yet never did his God or Friends forget.

And when deep talk and wisdom came in view,

Retir’d and gave to them their due.

For the rich help of Books he always took,

Though his own searching mind before

Was so with Notions written ore

As if wise Nature had made that her Book.

The effect of the close of this ode is due in part to the nature

of the world Cowley was experimenting in, the world of Hobbes,

Harvey, and the Royal Society. But on the whole, it may be

ascribed to the fact that reason rather than poetic imagina¬

tion is at work in a poetic stream.

Cowley’s intelligence is an interesting one. Curiosity, a

vigorous intellectual grasp of the external world as fact, and

Waller stein— Cowley as a Man of Letters .

131

the free play of interest in this fact and in theories of all sorts

are basic in it* To compare it with Donne's will define it and

will show how really different Cowley is from Donne, and how

the pupil changed what he took. Donne's is the far greater

and more intense intellect. Raymond Alden noted in the love

lyrics of Donne a profound intellectual seriousness not found

in Cowley's.2 All the drive of Donne's character and imagina¬

tion are behind his intellect. All parts of Donne's life and per¬

ception bear on each part. Sheer restless intelligence and

curiosity are strong in him; but what gives his thought its

special quality is that there is nothing he observes or studies

but he strives to make it at once a part of his philosophy and

thence a part of his actual belief and will and feeling. Donne's

is that “concentration of reason in feeling” which can lead to

mysticism. Cowley's intellect, on the other hand, is not phi¬

losophical. It has not the Baconian sheer dry weight and

depth, nor has it, as I have said, the imaginative drive of

Donne's. Yet Cowley has a keen interest in scientific ideas,

and he is a leader in giving them currency in the cultivated

world as the Proposition for the Advancement of Learning , To

the Royal Society , and the influence of his poems show.

A brief glance at the method of reading of Cowley and Donne

reiterates the same truth. Mediaeval thought and logic had a

deep influence in shaping Donne's imagination, as Courthope

has shown.3 But it found its opportunity in the character of

Donne's mind and his way of reading. One can imagine him

plunging into every book he reads as a matter of life and

death. Cowley is far more detached from his intellectual game.

He reads as a scholar and critic. One remembers in all this

that Cowley is forty-five years younger than Donne, and that

Hobbes and other reading of Cowley's maturer years dealt with

ideas less imperatively applicable to the individual conscious¬

ness than was Donne's reading. But even where Cowley han¬

dles the mediaeval reading, in the notes to Davideis , he handles

it as he handles Hobbes, with shrewd critical insight rather

than passion. His temper of a man of letters, if it deny him

certain other gifts, is fortunately suited to the material of the

2 Raymond M. Alden, The Lyrical Conceits of the “Metaphysical Poets”,

Studies in Philology , Vol. XVII. pp. 183-197.

8 W. J. Courthope, A History of English Poetry, Vol. Ill, Chapter VI, pp.

103 ff.

132 Wisconsin Academy of Sciences , Arts , and Letters .

time and to making it current in the literary traffic of the day.

Johnson's word on Cowley’s mind is final: “A mind capacious

by nature and replenished by study . . . always either ingenious

or learned, either acute or profound.”

With these differences in the two minds, as I have suggested,

Donne’s perception finds its characteristic and inevitable ex¬

pression first in the love lyrics and the religious lyrics, poems

in which the whole content of thought is absorbed in the flash

of feeling; and then in the gusto and irony and passionate

anger of question which characterize the Satires and Epistles.

Cowley’s genius is not suited to the love lyric. True, The Mis¬

tress is specifically in the school of Donne, and Cowley, sensi¬

tive as he is, has caught the vision of Donne, wondering at the

complex of personality impinging on the outer world, just as

he caught the sweep of Pindar. But that imaginative intensity

is not the chief note of even The Mistress , as I shall show when

I speak of imagery.

Samuel Johnson could see no ground for what Clarendon

said of the influence of Ben Jonson on Cowley. Yet the ob¬

jective definition of his condition and feeling in the ode on Her-

vey is in the spirit of Ben Jonson’s amplest verse, just as his

grasp of the traditional imagery is in the spirit of Jonson’s

epitaphs, and of Jonson’s general “judgement” and “eloquence”

and “masculine expression”, to use Clarendon’s phrases.* 4 Though

Cowley has no note of sublimity and though in detail he has a

grotesque literalness which pushes into an arabesque of style,

he has a deep sense for noble and comprehensive thought, for

the details of the visible world as material and as instrument

of thought, and for the glow of emotion that spring from such

thought. A splendor of this sort hovers in the Pindarics , in

much of the Davideis, in the bold conception and rapid imagery

of the Hymn to Light. It was these things that Milton must

have felt when he ranked Cowley as a favorite. The final

poetic fusion of this material wanting in the actual words of

Cowley, Milton could himself make, and must have made even

as he read. The range of life and reflection, the sweep of hu¬

man experience, he appreciated. And these, tempered by Cow¬

ley’s personality, give everywhere the sense of a great implicit

poetry so much larger than the actual accomplishment.

_

4 Edward Hyde, Earl of Clarendon: His Life by Himself, Oxford 1857, Vol. I,

p. 28.

Waller stein— Cowley as a Man of Letters .

133

It was another side of this intellectual quality which worked

upon Dryden and Pope, — namely, its power of definition and

generalization. Where Donne contributed amply to the gen¬

eral spirit of Dryden’s rhetoric in the Satires , Cowley gave

much to the detail of its reflection and to its technique and even

more to Dryden in the Odes and to Pope. Consider the Pinda¬

rics, for example, and the moral and sentimental reflections of

Pope’s Homer . The same drawing out of moral reflection and

sentiment as we find in Pope is clearly present, for example, in

Cowley’s Second Olympic of Pindar :

They [Theron’s forefathers] through rough ways, o’re many

stops they past,

Till on the fatal bank at last

They Agrigentum built, the beauteous Eye

Of fair-fac’d Sicilie,

Which does it self F th’ River by

With Pride and Joy espy.

Then chearful Notes their Painted Years did sing,

And Wealth was one, and Honour th’ other Wing.

Their genuine Virtues did more sweet and clear,

In Fortunes graceful dress appear.

To which great Son of Rhea, say

The Firm Word which forbids things to Decay .

For the past sufferings of this noble Race

(Since things once past, and fled out of thine hand,

Hearken no more to thy command)

Let present joys fill up their place,

And with Oblivions silent stroke deface

Of foregone Ills the very trace .

But Death did them from future dangers free,

What God (alas) will Caution be

For Living Mans securitie,

Or will ensure our Vessel in this faithless Sea? 5

Again, for very specific sentiment and definition we may look

at the lines on Virgil in The Motto :

8 For the reader’s convenience, to make the point even more clear, I give

here Ernest Myers’ prose translation of the passages cited above:

“They after long toils bravely borne took by a river’s side a sacred

dwelling place, and became the eye of Sicily, and a life of good luck clave

to them, bringing them wealth and Honour to crown their inborn worth.

O son of Kronos and of Rhea . . . guard ever graciously their native

fields for their sons that shall come after them.

Now of deeds done whether they be right or wrong not even Time the

father of all can make undone the accomplishment, yet with happy fortune

forgetfulness may come. For by high delights an alien pain is quelled and

dieth, when the decree of God sendeth happiness to grow aloft and widely. . . .

Ay but to mortals the day of death is certain never . . .”

134 Wisconsin Academy of Sciences, Arts, and Letters .

Virgil the Wise

Whose verse walks highest , but not flies.

Who brought green Poesie to her perfect Age;

And made that Art which was a Rage.

The definitions in the poems Of Wit and Of Reason have the

same quality, as also do epigrammatic elements in Davideis,

or, to take a final example, the last stanza in the Pindaric to

Dr. Scarborough, ending with the line, “When all's done, Life

is an Incurable Disease.” The spirit of the sweep of the Re¬

naissance is in him, but in the chop of the two tides, his tem¬

per answers more readily to the new mode of analytical defi¬

nition and reflection. Cowley’s own phrase of syllogisms and

enthymemes might be taken as a figure of it, when he says

of Pindar and Isaiah, “They wrote in enthymemes, we write in

syllogisms.” The age was no longer ready to drop a premiss

and flash to its conclusion, but was beginning to think and to

speak in the step by step movement of the syllogism, and Cow¬

ley led the way.

In what I have said above and in what I shall say, I am

aware that it was Pope who asked, “Who now reads Cowley?”

Pope, however, was exaggerating for a special point; more¬

over, to continue the quotation, “the language of his heart”

which Pope praises means much more than the essays suggest¬

ed by Courthope; and it is to be noted, too, that “his moral

pleases” Pope. Further, Pope’s dismissal, such as it is, must

not be taken at its face value. This becomes even clearer if

we give brief consideration to the detail of Cowley’s style.

Pope and Dryden consciously studied Denham and Waller as

the models of their technique — Dryden especially as he ma¬

tured. But as their work shows, and as they indicate in vari¬

ous passages, they read and absorbed Cowley in the years when

their imaginations were taking shape.6 That silent formative

influence must have been immense. As Thompson has shown,

it was from Cowley that Dryden and Pope learned the breadth

and spirit — the substance we may say — of the couplet which

they finished on the models of Waller and Denham.7 His in¬

fluence on the rhetoric of their couplet was no less great. Not

6 In Pope’s case the influence of Crashaw too was strong, and in part in the

same direction.

7 A Hamilton Thompson, Abraham Cowley. Cambridge History of English

Literature, Vol. VII, Chapter III, pp. 70-79.

Wallerstem ■ — Cowley as a Man of Letters .

135

only is the type of rhetoric indicated in the paragraphs above

present in Cowley, but it finds in him its characteristic suc¬

cinct, general, definitive language, even in descriptive or nar¬

rative passages. It permeates Davideis, as one example from

the wooing of David and Michel will show:

She weighed all this as well as we may conceive,

When those strong thoughts attacked her doubtful breast;

His beauty no less active than the rest.

But the shaping quality of Cowley's reasoning temper is

equally manifest in the more obvious pervasive element of his

style, in its wit and so called “metaphysical” quality. “Meta¬

physical” style is a very different thing in Donne and Cowley.

The stuff, the manner, the imaginative effect of the imagery

is very different, although they have in common the intellectual

form of statement and the logical extension of image. Donne's

images most often represent the truth of the world of inner

experience, their outer curiousness being a simple truth in that

world, their logic the logic of philosophical imagination seek¬

ing to pierce through objects in their casual temporal juxtapo¬

sition, to spiritual and emotional reality ; and in the satires, his

figures make appeal from worldly values to human. Cowley’s

imagery, if we define it by this comparison, represents not in¬

ner life, but sentiment, wit, fancy. Whereas Donne has ima¬

gination in the Coleridgean sense, handling and shifting objects

to get at what lies behind them, Cowley's is fancy musing upon

objects and playing with them, indeed, to see all their facets,

but still focusing upon them as fixed objects.

To analyse several of Donne’s characteristic images and ap¬

proach Cowley's figures through them will explain my mean¬

ing. In Donne's song “Sweetest love, I do not go”, the whole

theme, as in many others, is the significance, to a lover, of per¬

sonal experience as against the outer mechanical fact of life.

The images recreate this personal reality, following boldly and

directly the psychological flow of the emotion. “When thou

sigh'st, thou sigh’st not wind” (He will return as surely as

the sun, he has said. Now the mind articulates a fact of na¬

ture in another sphere, the fact of her physical life, and the

emotion, catching hold of the fact by its symbol, the word, and

pivoting on that, pulls the fact back into its own reality of

the spirit.) “When thou sigh'st, thou sigh’st not wind, But

136 Wisconsin Academy of Sciences , Arts, and Letters .

sigh’st my soul away”. The great figure in one of the sonnets

on death,

At the round earth’s imagined corners blow

Your trumpets, Angels, —

is not literally reasonable on the face of it. But reason may

deal with it profoundly if its full thought is looked at, if the

enthymeme be expanded. It does not matter, says Donne, if

the traditional pictures with which we figure the universe be

scientifically false; they are but symbols, and the reality is not

circumscribed by them, either in itself or for us. This truth he

gives us in one flash. The compass figure in the Valediction,

again, takes us step by step deeper into the perception of the

ideal thought on which the emotion rests, and the emotion in¬

tensifies and expands as the concept clarifies. What does this

teach us of Cowley?

Cowley's imagery develops in a quite different spirit. Like

his thought, it is many things in many types of poem. The

Anacreontics have little essentially in common with the Pinda¬

rics. But certainly the trick of image elaborated in detail be¬

comes a habit on the surface of his whole style. And he is set

going by the extension of figures in earlier poets, particularly

Donne's figures. It is this “witty” or “metaphysical” imagery

that we now want especially to look at. Some of the poems of

The Mistress are little allegories of love play, written in the

manner of the Elizabethans or Herrick's songs for music, but

with the figure of wit and fact caught from Donne taking the

place of a myth. Written in Juice of Lemon illustrates this.

Again, The Change begins with courtly Petrarchean play:

Love in her Sunny Eyes does basking play;

Love walks the pleasant Mazes of her Hair;

Love does on both her Lips forever stray; —

But this shifts to analysis:

With me alas, quite contrary it fares;

Darkness and Death lies in my weeping eyes,

Despair and Paleness in my face appears,

And Grief, and Fear, Love’s greatest Enemies;

But like the Persian-Tyrant , Love within

Keeps his proud Court , and ne’re is seen.

The analysis itself, however, becomes not the analysis of emo¬

tion or ideal situation, but rather the analysis of the figure

Waller stein— Cowley as a Man of Letters.

137

itself and hence draws out to its logical conclusion, the epi¬

gram:

Oh take my Heart, and by that means you’ll prove

Within, too stor’d enough of Love:

Give me but Yours, I’ll by that change so thrive,

That Love in all my parts shall live.

So powerful is this change, it render can,

My outside Woman, and your inside Man.

Poems which suggest specific comparison with Donne define

Cowley very clearly. The lyric My Heart Discovered , in its

search of heart and soul through body, evokes a close compari¬

son with Donne, and in particular the whole conception of the

opening lines,

Her body is so gently bright,

Clear and transparent to the sight,

That through her flesh, methinks, is seen

The brighter soul that dwells within;

carries us back to Donne's description of Elizabeth Drury,

Her pure and innocent blood

Spoke in her cheeks and so distinctly wrought,

That one might almost say her body thought.

In this poem, one of Cowley’s most delightful, the turn of wit

answers deeply to the theme of the poem. Skipping a few

lines :

I through her Breast her Heart espy,

As Souls in hearts do Souls descry,

I see’t with gentle Motions beat;

I see Light in’t, but find no Heat .

But, oh, what other Heart is there,

Which sighs and crouds to hers so neer, . . . .

The wounds are many in’t and deep;

Still does it bleed, and still does weep.

Whose ever wretched heart it be,

I cannot chuse but grieve to see;

What pity in my Breast does raign,

Methinks I feel too all its pain.

So torn, and so defac’d it lies,

That it could ne’re be known by th’ eyes;

But, oh, at last I heard it grone,

And knew by th’ voice that ’twas mine own.

138 Wisconsin Academy of Sciences , Arts and Letters .

So poor Alcione _ [Here he tells briefly how she unknow¬

ing weeps the body of her husband.]

What should the wretched Widow do?

Grief chang’d her straight; away she flew,

Turn’d to a Bird : and so at last shall I,

Both from my Murther’d Heart, and MurtWrer fly.

The theme is not as in Donne the yearning of personal feeling,

or imagination trying to transcend fact. It is objective reflec¬

tion, half pathetic, half comic, and the temper in which it views

its subject might well be called that of social comedy. And

again it works up to the epigram. Another lyric, Silence,

which suggests comparison with The Triple Fool, is, in theme,

the Donnian passionate analysis of passion, but in expression,

the logic moves, like the Hervey ode, with the slow definition of

prose, and ends not in the deeper focus of emotion into which

Donne takes us, but in the sententious phrase. The given

Heart, which is frankly and wholly the extended epigram, is on

the other hand, rapid, gay, integrated. We see, then, that this

prose quality of Cowley's mind, this literal development, char¬

acteristically steps in. Where the figure has started to be emo¬

tional, it often becomes merely ludicrous. Where the figure is

a figure of idea, it can, like Dry den's figures in Absalom, be

made to carry forward an argument very prettily. In The

Welcome this style of figure gives an enchanting gaiety ; in To

Wit it heightens and makes rapid the definition. These two

are akin in objectivity and lucid definition to Ben Jonson. In

My Dyet the subject matter once more comes from Donnian

logic, but the objectivity of the new manner gives it its char¬

acter and charm. The spirit of the conceit in Cowley has be¬

come intellectual and reasoning, and grows sometimes to reflec¬

tion and sometimes to social comedy.

The conceit pushed too far. And Pope and Dryden wisely

reprobated its excess of elaboration. In even great contexts,

the image pushed to the limit not only loses its emotion but for¬

gets its idea, and carries on in mere literal play of word. Start¬

ing in the passion to be complete in every detail of feeling and

ratiocination, it is abstracted from its theme; it becomes an

empty cleverness; it just trembles into periphrasis. The ap¬

proach to periphrasis is widespread in his work, though not

sharply focused. My Muse and To Mr . Hobs, in the lesser de¬

tail, well illustrate this:

Wallerstein — Cowley as a Man of Letters.

139

The Baltique, Euxin, and the Caspian,

And slender limb’d Mediterranean,

Seem narrow Creeks to Thee, and only fit

For the poor wretched Fisher-boats of Wit.

Thy nobler Vessel the vast Ocean tries,

And nothing sees but Seas and Skies,

Till unknown Regions it descries,

Thou great Columbus of the Golden Lands of new Philosophies.

Cowley approaches here a distinct “poetic diction”. His ab¬

stract intellection and the fact that ratiocination usurped the

ground of his thought were tendencies pushing him, perhaps

inevitably, towards such a diction. Because of these qualities

in his creation he tends to generalize his adjectives; and his

images, because one element of life is abstracted from the rest,

lose their sensousness and become abstract symbols of ideas of

objects.8 Finally, he tends as a result of these two develop¬

ments, and perhaps seeking an elegance congruous with this

rhetoric, to a theory and practice of poetic words. The notes

to Davideis, Book II, for example, explain the use of wife in¬

stead of spouse on the ground of poetic suitability of word.9

Here too Pope absorbed before he criticised. The abstract

play of reason caught him. There were other great influences

at work in forming the poetic diction of Pope.10 But from

what I have said it is, I hope, clear that the Popean periphrasis

received a significant impulse from Cowley.11 It will be re¬

membered that Pope himself called his early work “fancy's

maze” in contradistinction to the “truth” and “moralized song”

of his later work.12

8 This point may perhaps be more clear if we look at imagery in another

field than Cowley’s. It is interesting to note that in our own day we have a

group of lesser lyric poets who go through the same process. They concentrate

on the experiencing of the emotion, and in so doing they isolate the emotion from

the experience which caused it, and its relation to their thought and action,

hoping to intensify it by the very analysis of it. They abound, of course, in

series of images very specific in detail; but in effect, though their object is

feeling and not reason, they strike us just as does the “poetic diction” of the

Eighteenth Century. Elinor Wylie illustrates this tendency, and it is very

strongly marked in some of the very minor poets.

9 Davideis, Book II. Poems, Ed. A. R. Waller, Cambridge 1905, p. 321.

10 See, for example, the discussion of the influence of Milton in Sir Walter

Raleigh’s Milton, London, 1900, Chapter V, pp. 218-255 ; and the discussion of

the style of Dryden’s Virgil in Mr. Mark Van Doren’s Dryden.

11 The influence in this direction would not be limited to Cowley ; the extended

image in Crashaw must be borne in mind.

12 Epistle to Dr. Arhuthnot. Poems, Ed. H. W. Boynton, Boston, 1903, p. 181.

140 Wisconsin Academy of Sciences , Arts , and Letters .

Thus if Cowley did not find himself as a poet, he did find

himself as a man of letters. To bring to the fore this intellec¬

tual and logical temper, even though it was to prune away so

much of his own time that was dear to him, was one of his

tasks as a man of letters, sensitive to the current of ideas, just

as to keep alive the breadth and range of perceptions and the

objective imagery of the Pindarics , ready for greater hands to

kindle to flames, was another. Cowley is an important factor

in understanding how English letters passed so rapidly from

the multifarious splendor of the Renaissance to the more limited

but the assured accomplishment of Dryden.

WISCONSIN MYXOMYCETES

H. C. Greene

Foreword

The Myxomycetes of Wisconsin have already been the sub¬

ject of a comprehensive paper. In 1914 Alletta F. Dean pub¬

lished an article entitled The Myxomycetes of Wisconsin',

(Trans. Wis. Acad. Sci. 17 : 1221-1299. 1914), in which a

thorough treatment of the Wisconsin Myxomycetes then known,

largely her collections, was undertaken. Reprints of this paper

are no longer available. Moreover, it was, unfortunately, not

illustrated, and since that date some forms not at that time

recorded for Wisconsin have been collected. This combination

of facts seems to the writer to justify the publication of a brief,

illustrated paper which will bring the subject up to date, and

will, it is hoped, by means of the illustrations, stimulate curi¬

osity regarding this very interesting, but somewhat neglected

group.

The classification and the generic key presented follow Mac-

bride (North American Slime-Moulds , New and Revised Edi¬

tion 1922, 299 pp., Macmillan Co.), and certain of the illus¬

trations are redrawn from Lister ( Mycetozoa , Third Edition

1925, 296 pp., 222 pis. British Museum, N. H., publication.)

It will be noted that the generic key has reference only to

the Sub-class Myxogastres, and that therefore Ceratiomyxa,

the first genus treated in the paper, is not included in the key.

It is sufficiently distinctive, however, to be easily recognized on

its own account. After the genus name in the key appears the

number of the plate in which the illustration of the genus in

question is to be found. Descriptions of the figures in the

plates will be found in the latter part of this paper.

The valued and congenial aid of Dr. E. M. Gilbert of the

Department of Botany of the University of Wisconsin has made

this paper possible. To him, and to others who have been of

assistance, the writer desires to convey his sincere thanks.

Introduction

The Myxomycetes are a compact group of animal-like fungi

(or fungus-like animals, if you prefer) low in the scale of life

142 Wisconsin Academy of Sciences , Arts , and Letters.

and usually tiny and inconspicuous, which, by their great beau¬

ty, their diversity, and their availability in any moist, wooded

country, form a fascinating and easily accessible subject for

study by naturalists.

Myxomycetes (also called slime-moulds and Mycetozoa), in

the course of their relatively simple life cycle, display char¬

acters which are diversely ascribed to both plant and animal

kingdoms, so that students of classification have been, and still

are, somewhat at a loss as to the proper position of these curi¬

ous forms. During their vegetative existence the Myxomycetes

form so-called plasmodia which superficially resemble giant

amoebae, while during their reproductive period they resemble

the Gasteromycetes, a group of higher fungi, but are distin¬

guished from the Gasteromycetes by the total lack of a myceli¬

um. Dr. T. H. Macbride, the preeminent American student of

the Myxomycetes, makes the following remarks in the intro¬

duction to his North American Slime-Moulds: “But why call

them either animals or plants? Was nature then so poor that

forsooth only two lines of differentiation were at the beginning

open for her effort? May we not rather believe that life’s

tree may have risen at first in hundreds of tentative trunks of

which two have become in the progress of the ages so far

dominant as to entirely obscure less progressive types? The

Myxomycetes are independent: all that we may attempt to

assert is their near kinship with one or other of life’s great

branches.”

As above stated, Myxomycetes are found in considerable

abundance in any moist, wooded area at the proper season of

the year. In the Middle West these diminutive forms are to

be had from the middle of May until late fall. The type of

substratum chosen for vegetative growth is diverse, but must

be moist and in not too bright light. Old rotten logs, sticks, hu¬

mus, various large fungi, and all sorts of vegetable debris sup¬

port the plasmodia of Myxomycetes. When the time for fruit¬

ing arrives the amoeboid plasmodia creep out of their dark

feeding grounds, and come to fruit, as a rule, in fairly open

situations such as the tops of logs, sticks, leaves, and so forth.

In general, the mature fruits, representing the plant-like

phase, are much more conspicuous, than are the vegetative

plasmodia, which represent the animal-like phase. Therefore,

Greene— -Wisconsin Myxomycetes.

143

the fruiting phase will be first discussed in the following ac¬

count of the life cycle characteristic of Myxomycetes.

The fruiting bodies of Myxomycetes are called sporangia,

that is, (with the exception of one small genus, Ceratiomyxa)

they contain within them the numerous spores which, under

favorable conditions, germinate to produce the vegetative phase.

The diverse morphological characters of the sporangia and the

spores form the basis of our classification of these forms.

There may be many thousands of sporangia in a single fruc¬

tification, there may be hundreds, or there may be only a few,

or even one in the case of compound fructifications. Certain

species customarily have much larger fructifications than oth¬

ers. The sporangia may be very closely clustered, or merely

gregarious, or quite widely scattered, a matter again depending

in large degree upon the species concerned. Many rudimentary

sporangia may be closely crowded together in a single layer, or

superimposed over each other in several layers, and the aggre¬

gate is then known as an aethalium. In numerous cases indi¬

vidual sporangia are not differentiated at all and the flatly

rounded, irregular, elongated, structures resulting are termed

plasmodio carps.

Distinct, individual sporangia are, in most genera, small and

delicate, not over 1 millimeter in their dimension. In some

genera, however, as in Stemonitis and Comatricha, the length

of sporangia may be as great as 25 mm. the width not exceed¬

ing 1 or 2 mm. In the case of Arcyria, and in lesser degree in

that of Hemitrichia, the capillitium may expand beyond the ori¬

ginal limits of the sporangium until the effect is one of droop¬

ing plumes, with a length of some 12 or 15 mm. or more in a few

instances.

Aethalia, composed of many rudimentary sporangia, are

sometimes very large. Some of the flat aethalia of Lindbladia

effusa have been reported as having a diameter of 50 centi¬

meters, — about 20 inches ! Most aethalia are much smaller than

this, but are, nevertheless, often quite conspicuous.

As mentioned above, instead of definitely delimited sporangia,

there are often found elongate, sessile, flat sporangia known as

plasmodiocarps. Physarum sinuosum furnishes a good example

of the plasmodiocarp habit. Plasmodiocarps may be short and

straight, with their long diameter not more than twice that of

144 Wisconsin Academy of Sciences , Arts , and Letters .

the short. They may be short and curved, long and straight,

long and curved, ring-shaped, scalariform, or netted, or any

combination of these. The exact shape, as is also true in the

case of aethalia, is likely to be dependent on the contours of the

substratum to a greater or less extent. Plasmodiocarps appear

in some cases in company with perfectly formed individual

sporangia, while in the case of such a form as Hemitrichia

serpula the plasmodiocarp habit is most pronounced and dis¬

tinctive, resulting in an elaborate looped, reticulate fructifica¬

tion. Most plasmodiocarps are slender, not over a few milli¬

meters in width.

The typical fruiting body of a Myxomycete is composed of

(1) The sporangium proper which is a more or less rounded

structure enclosed by a membrane called the peridium. In many

species the peridium is coated with lime incrustations of varied

hue. Sometimes, as in the genus Diderma , the coating of lime

is more or less remote from the membranous peridium, giving

the appearance of a double wall. In species where lime is lack¬

ing the peridium may be shining or iridescent. The tint of cer¬

tain peridia is definitely due to the color of the spore mass

enclosed. In the genera, Cribraria and Dictydium, the peridium

is pierced by many small openings. In all cases, sooner or later,

it is the rupture of the peridium which permits the dispersal

of the spores. In most genera a system of membranous threads

or tubules is to be found within the sporangium. These threads

collectively form the capillitium. In different forms the capil-

litium varies greatly and forms one of the principal differential

diagnostic features. It may form an intricate branching system,

the threads may radiate from a central point of attachment,

or they may be free within the sporangium, or still otherwise

arranged. The capillitial threads may be entirely charged with

granules of lime as in Badhamia, or only partly so as in

Physarum. In most genera lime is not noticeably present in the

capillitium. The capillitium may be colorless or colored, depend¬

ing upon the species concerned, and the threads in some forms

are profusely sculptured, while other capillitial filaments are

practically smooth. The spores are one-celled, usually spherical,

small (not often more than 12 micra in diameter), and of vari¬

ous tints of brown or yellow. The color of spores in mass is

usually much deeper than that of the individual spore as seen

under the microscope. Spores may be practically smooth as in

Greene — Wisconsin Myxomycetes.

145

Cribraria, they may be covered with warts of varying size and

abundance, or they may be covered by a system of netlike bands,

spoken of as reticulations. Spore size and markings are impor¬

tant diagnostic characters.

(2) The stalk or stipe which bears the sporangium is mem¬

branous in nature, may be charged with lime or waste matter,

or both, and is most variable in shape, length, and color in

different species and genera. The stipe in many forms is con¬

tinued upward into the sporangium proper, and this upward

continuation is called the columella. The columella is variable

in size, shape, branching, and composition, and may or may

not, according to the form concerned, attain to the apex of the

sporangium. Collumellae are also found in some forms which

are without stipes in which case the columella may be regarded

as the thickened base of the sporangium. In many species the

stipes arise from a more or less effused base spoken of as the

hypothallus. There may be a distinct hypothallus associated

with each sporangium, as for example, in the case of Didymium

nigripes , or all sporangia of a given fructification may arise

from a common hypothallus, as in Stemonitis . The hypothallus

may be dull or shining, is also present in the case of certain

forms without stipes, and is likewise found in connection with

some aethalia and plasmodiocarps.

As may have already been noted from the context, the typi¬

cal fruiting body thus characterized is by no means the type

universally found. The stipe may be lacking and the sporangia

are then sessile, the columella is often lacking, and there is no

hypothallus in many species. Some species lack a capillitium,

and in still other species the peridium is so early evanescent

as to seem to the casual observer to be entirely absent. The

reader is referred to the illustrations accompanying the article

in order to gain an idea of the great variation found.

When the sporangia reach maturity, the spore mass is blown

away or falls to the ground, and with the germination of the

spores under the proper conditions, the vegetative animal-like

phase is initiated, and will now be described.

The disseminated spores, under suitable conditions of mois¬

ture and temperature germinate, that is to say, the spore cell-

wall is ruptured and the content escapes. The spore-content at

this stage resembles an amoeba, has a contractile vacuole, and a

146 Wisconsin Academy of Sciences , Arts , and Letters .

nucleus, and creeps slowly about. Usually in a short time the

amoeboid body produces a whip-like flagellum, with which it

lashes about through its moist life-medium, showing consider¬

able activity. This swarm-cell as it is now called, rotates

actively for a time, and then slows down once more, again

developing a creeping movement. At this point the swarm¬

cell becomes capable of ingesting food particles such as

fungous spores and bacteria. The swarm-cells grow and multi¬

ply by division, for an indefinite period, then withdraw the

flagellum, become amoeboid once more, and are henceforth

known as myxamoebae. The myxamoebae grow and divide as

do the swarm-cells, and eventually fuse in pairs. The pairs then

fuse with many other pairs and a slimy, multinucleate plasmo-

dium results. The plasmodium possesses the power of locomo¬

tion and creeps about, over or through the substratum in search

of food, as a slimy vein-like mass of greater or lesser extent,

its size often to some degree depending upon the species con¬

cerned. If conditions become unfavorable for continued activity,

the plasmodia possess the ability to enter a resting stage, form¬

ing dry, horny masses which may assume again the plasmodial

characteristics at the resumption of proper conditions.

The plasmodia are variously colored, and the colors have been

used to some extent as a diagnostic feature, but ordinarily the

student is unable to follow plasmodial development, and thus

these colors, which are said also to vary from time to time, are

of comparatively little practical aid.

This plasmodial condition may persist for months, usually

in the cracks and fissures of moist logs, and in darkness more

or less absolute. The plasmodia then, when conditions are

favorable, creep out into the brighter light, and, after a longer

or shorter period, pass rather suddenly to fruit, usually in an

advantageous position for spore dissemination. This transition

from the slimy amoeboid mass to the beautiful and intricate

fruiting bodies is a most interesting phenomenon. That such

conglomerates so similar to one another can suddenly assume

forms so radically different, to some extent sets the group apart,

and has made it an object of vitally engrossing study to biolo¬

gists for many years.

Greene — Wisconsin Myxomycetes.

147

Key to the Principal Genera of the

Sub-Class Myxogastres

(after Macbride)

1. Spores in mass violet-black, or sometimes reddish-brown.

2. Capillitium of simple or branching threads; sporangia more or less

completely charged with lime. Order Physarales.

3. Capillitium of hyaline, branching tubules, more or less completely

charged with lime; peridium and stipe, one or the other or both,

usually containing lime. Family Physaraceae.

4. Fructification of cushion-like aethalia. Fuligo (Plate I).

4. Fructification plasmodiocarpous, or of distinct sporangia.

5. Peridium bearing lime externally.

6. Capillitial threads charged with lime throughout.

Badhamia (Plate I).

6. Capillitial threads not completely charged with lime, but

bearing lime nodules.

7. Sporangia other than goblet-shaped; dehiscence irreg¬

ular. Physarum (Plate I).

7. Sporangia goblet-shaped.

8. Dehiscence by a distinct lid, or if not by a lid quite

regularly circumscissle. Craterium (Plate II).

8. Dehiscence irregular, with peridial segments appear¬

ing as “petals”. Physarella .

5. Peridium without lime externally.

9. Fructification plasmodiocarpous. Cienkowskia.

9. Sporangia distinct. Leocarpus (Plate II).

3. Capillitium of simple or rather sparingly branched threads with¬

out lime; lime, when present, usually on peridium only.

Family Didymiaceae.

10. Fructification an elongate, thick aethalium.

Mueilago (Plate II).

10. Fructification plasmodiocarpous, or of distinct sporangia.

11. Lime deposits present on peridium.

12. Lime desposits of distinct, stellate crystals.

Didymium (Plate II).

12. Lime deposits not in form of distinct, stellate crystals.

13. Lime deposits of small, rounded granules only;

peridium usually distinctly double.

Diderma (Plate II),

13. Lime desposits in the form of large, scattered

scales, imbedded in, or resting loosely on the carti¬

laginous peridium. Lepidoderma.

148 Wisconsin Academy of Sciences, Arts, and Letters,

11. Lime deposits not present on peridium; peridium double,

the outer layer gelatinous. Colloderma.

2. Capillitium thread-like, much branched, arising from a columella;

sporangia without lime (except in Diachaea). Order Stemonitales.

14. Fructification of blackish aethalia.

14. Fructification of blackish aethalia. Family Amaurochaetaceae.

15. Capillitium of fibrous threads, without vesicles.

Amaurochaete.

14. Fructification of distinct sporangia, except in Brefeldia; eapil-

litum springing from all points on the columella.

Family Stemonitaceae.

16. Fructification aethalioid; capillitium charged with vesicles.

Brefeldia (Plate III).

16. Fructification of distinct sporangia.

17. Stipe and columella black; no lime present.

18. Capillitium forming a surface net over the coarser

branches ; sporangia usually closely clustered and

long-cylindrical. Stemonitis (Plate III).

; 18. Capillitium without surface net; sporangia usually

scattered and globose, or shortly cylindrical.

Comatricha (Plate III).

17. Stipe and columella white; heavily charged with lime.

Diachaea (Plate III).

14. Sporangia distinct, but capillitium developed from summit of

columella. Family Lamprodermaceae.

19. Columella extending entirely through the sporangium, and

capillitium springing from a disk at the apex.

Enerthenema (Plate III).

19. Columella shorter, not reaching beyond center of the spo¬

rangium; capillitium intricate; peridium shining, iridescent.

Lamproderma (Plate IV).

19. Columella very short or rudimentary; capillitium of stiff,

little-forked branches which at maturity bear plate-like rem¬

nants of the sporangial wall at their tips.

Clastoderma (Plate IV).

1. Spores in mass usually some shade of brown or yellow, rarely purplish

or rosy, never violaceous black.

20. Capillitium absent; pseudo-capillitium present in Retieularia and

Enteridium; spores in mass yellowish through brown, rarely

purplish. Order Cribrariales.

21. Fruiting bodies plasmodiocarpous, scattered. Family Liceaceae.

22. Plasmodiocarps variously shaped; peridium simple. Licea.

21. Fructification aethalioid; sporangia well defined; closely ap-

pressed, tubular, rupturing at top, side walls entire.

Family Tubiferaceae.

Greene-— Wisconsin Myxomycetes.

149

23. Spores in mass olivaceous-yellow; sporangia in one or sev¬

eral layers. Lindbladia (Plate IV).

23. Spores in mass rusty-red ; sporangia in one layer. Tubifera.

21. Fructification aethalioid; sporangia not well indicated, and with

side walls not entire. Family Reticulariaceae.

24. Spores in mass brownish or reddish.

25. Aethalia covered by a firm, mottled silvery cortex;

pseudo-capillitium of membranous expansions, much

frayed into fine threads; hypothallus white.

Reticularia (Plate IV).

25. Cortex evanescent, the aethalia when mature ap¬

pearing as a soft, red-brown, cushion-like mass;

pseudo-capillitium of broad, membranous, inter¬

weaving bands, not frayed out.

Enteridium (Plate V).

24. Spores in mass ochraceous; sporangial tops well-defined.

Dictydiaethalium (Plate V).

21. Sporangia separate and distinct, with the membranous sporan¬

gial well perforated so as to form a net, especially above.

Family Cribrariaceae.

26. Peridium more or less dissipated so as to form an apical net,

the intersections of which may or may not be expanded into

definite nodes. Cribr aria (Plate V).

26. Peridium more or less dissipated into a series of parallel

ribs, radiating from below, and connected with each other

by delicate transverse threads. Dictydium. (Plate V).

20. Fructification aethalioid, the individual aethalia resembling minia¬

ture puff-balls; peridium membranous; capillitium of tubules from

peridium; spore mass rosy or ashen, becoming yellowish.

Order Lycogalales.

27. As under 20. Family Lycogalaceae.

28. As under 20. Ly cog ala (Plate V).

20. Capillitium of sculptured threads, attached or free, simple or

branched; spores usually yellow; sporangia without lime, stalked

or sessile, distinct, rarely plasmodiocarpous. Order Trichiales.

29. Capillitium plain, roughened or spinulose, simple; the threads

sometimes attached to the sporangial walls.

Family Perichaenaceae.

30. Sporangia plasmodiocarpous; dehiscence irregular.

Ophiotheca.

30. Sporangia polygonal; dehiscence circumscissle.

Perichaena (Plate VI).

29. Capillitium a distinct net, attached below; threads roughened,

but not sculptured with smoothly continuous spiral bands.

Family Arcyriaceae.

150 Wisconsin Academy of Sciences , Arts , and Letters .

31. Peridium fragmentary, but persistent; capillitium non¬

elastic. Lachnobolus.

31. Peridium evanescent above, persistent below; capillitium

elastic. Arcyria (Plate VI).

29. Capillitial threads usually free, but in some forms composing

a loose, branched net; sculptured with definite spirals; or some¬

times with rings. Family Trichiaceae.

32. Capillitial threads forming a loose, attached net, the threads

of which are spirally sculptured. Hemitrichia (Plate VI).

32. Capillitial threads free.

33. Threads distinctly spirally sculptured.

Trichia (Plate VI).

33. Sculpture irregular or wanting. Oligonema.

Characterization of Wisconsin Slime Moulds

Class Myxomycetes (Link) De Bary

Fungus-like organisms, with an amoeba-like multinucleate

vegetative phase, and a fungus-like reproductive phase; the

spores, borne in connection with the reproductive phase, usually

enclosed in sporangia, but rarely free, germinating to produce

swarm-cells which fuse to form the vegetative phase.

Sub-Class Phytomyxinae Schroeter

This sub-class contains parasitic forms with which this paper

is not concerned, and it is therefore omitted from consideration.

Sub-Class Exosporeae Rost.

Spores developed externally on columnar sporophores ; sporo-

phores white, somewhat translucent, branching; spores white,

borne on stalks which project from the sporophore.

Genus Ceratiomyxa Schroeter. This peculiar genus is

unique in that the spores are borne externally on clustered,

columnar, white papillae. The genus is easily recognized in

the field by the fragile, translucent spore columns, character¬

istic of no other Myxomycetes.

Ceratiomyxa fruticolosa (Muell.) Macbr. (Plate I, fig. 1).

Sporophores clustered in little tufts, white, bearing the spores

externally, about 1 mm. high; spores white, ellipsoidal, each

borne on a separate stalk. Found in moist situations, especially

on wet logs, in spring and early summer. Common.

Greene — Wisconsin Myxomycetes.

151

Sub-Class Myxogastres (Fries) Macbr.

This sub-class comprises practically the entire group under

consideration. It is characterized by small, unicellular, usually

spherical spores of various tints, enclosed in sporangia which,

in most cases, bear a capillitium.

Order Physarales

Spores in mass violaceous-black; capillitium composed of

hyaline tubules, either completely or partly charged with lime ;

peridium sometimes without lime, but in most cases containing

lime in greater or less amount.

Family Physaraceae

Capillitium netted and branched, intricate, containing lime

to a greater or less extent ; peridium and stipe, one or the other

or both, usually containing lime deposits.

Genus Fuligo (Haller) Pers. Aethalioid fructifications,

usually large and conspicuous, and giving no indication exter¬

nally of the presence of individual sporangia; the outer layer

of the aethalium a thin, calcareous crust usually disappearing

after maturity; capillitium charged with lime to a greater or

lesser extent.

Fuligo septica (Linn.) Gmel. This species can be subdivided

into several forms, of which two are here characterized :

F. septica f. ovata (Schaeff.) Pers. (Plate I, fig. 2). Aetha¬

lium yellow to light brown, one to many centimeters in dia¬

meter, 1-2 or more cm. thick, variable in shape, lime crust

prominent, with the “surface when mature extremely friable,

like dry foam”; capillitium with branching yellow lime-knots;

spores violet-black in mass. On dead wood, and other debris.

Common.

F. septica f. violacea Pers. “Aethalium thin, two or three

inches wide, covered by a cortex at first dull red and very soft,

at length almost wholly vanishing, so that the entire mass takes

on a purple violet tint, upper surface varied with white ; capil¬

litium rather open, the more or less inflated, large, irregular

nodes joined by long, slender, delicate, transparent filaments;

spores dark violet . . .” (Macbr.) Reported by Dean, but not

seen by the writer.

152 Wisconsin Academy of Sciences , Arts, and Letters .

Fuligo cinerea (Schw.) Morg. Aethalioid or plasmodiocarp-

ous, white, irregular in shape and size, flat upon the substrate ;

capillitium charged with irregular lime-knots; spores violet-

black in mass, ellipsoidal. Fruiting bodies appearing in mass on

living, young tobacco plants near Madison, Wis., June 1931.

Given to the writer for identification by Dr. J. J. Davis, Curator

of the Herbarium, University of Wisconsin.

Genus Badhamia (Berk.) Rost. Sporangia usually well-

defined, sessile or stipitate; peridium of one layer; capillitium

intricate, of branching tubules charged with lime throughout

their extent. In external characters this genus much resembles

Fhysarum, next following.

Badhamia panicea (Fries) Rost. Sporangia gregarious, or

closely clustered, bluish-gray, with white lime-scales over sur¬

face, globose, about 1 mm. diam., sessile; lime tubules of

capillitium white, abundant, and showing plainly when spores

are blown away ; spores violet-black in mass. Collected by H. J.

Gorcica in the vicinity of Westboro, Wis., on dead poplar wood,

September 1931.

Badhamia papaverace a Berk, and Rav. Sporangia gregarious,

iridescent-gray with scanty lime-deposits on sporangium wall,

globose, about 1 mm. diam.; stipes short, blackish brown;

capillitium white, persistent after spores have blown away;

spores violet-black in mass. On dead wood.

Badhamia ruhiginosa (Chev.) Rost. Sporangia gregarious,

brownish, the sporangium wall without lime above, but calcare¬

ous below, egg-shaped, .5 mm. diam. ; stalk erect, stiff, reddish-

brown, prolonged within the sporangium as a columella and

arising from a small hypothallus ; capillitium dense, white, per¬

sistent, spores violet-black in mass. On dead wood.

Badhamia utricularis (Bull.) Berk. (Plate I, fig. 3). Spor¬

angia clustered, iridescent-gray, with lime-granules scanty;

sporangia single, or lobed and confluent, about 1 mm. diam.;

stalks long, membranous, straw-colored, prostrate; capillitium

white, not especially prominent; spores violet-black in mass.

On dead wood.

Genus Physarum (Pers.) Rost. Sporangia distinct, sessile

or stipitate, or tending toward the aethalioid or plasmodiocarp-

ous; peridium simple or double, with more or less lime;

Greene — Wisconsin Myxomycetes.

153

capillitium a branching net of hyaline tubules, more or less

densely charged with lime at the points of branching.

Physarum auriscalpium Cooke. Reported for Wisconsin by

Dean. Specimen not available for examination by the writer.

Physarum cinereum (Batsch) Pers. Sporangia gregarious,

or closely clustered, gray, sessile, subglobose, through various

forms to definite plasmodiocarps, usually about .5 mm. or less

in diameter; capillitium of branching threads with rather pro¬

nounced white lime-knots ; spores violet-black in mass. Frequent

on turf and lawns, where large numbers of sporangia may

appear.

Physarum confertum Macbr. Sporangia heaped in masses,

dark gray with limeless peridium, globose, .2-.4 mm. diam. ;

capillitium scanty, consisting of colorless threads with very

small, angular lime-knots; spores violet-black in mass. On

sticks, grass, etc.

Physarum contextum Pers. Sporangia closely crowded, often

so much so as to be interwoven, yellow, sessile, elongate, about

.5 mm. diam. ; peridium in two layers, the outer thick, crustose,

yellow, the inner thin, yellowish; capillitium of branching

threads, with large, irregular lime-knots ; spores violet-black in

mass. On sticks, bark, etc.

Physarum globuliferum (Bull.) Pers. Sporangia gregarious,

white, globose, .5 mm. diam. ; stipe pale above, becoming brown¬

ish below, rather slender, of variable length, continued into the

sporangium as a small conical columella; capillitium a dense

network of colorless threads with rounded, whitish lime-knots ;

spores violet-black in mass. On dead wood and leaves.

Physarum leucopus Link. (Plate I, fig. 5) . Sporangia gregari¬

ous, snow-white, with a dense covering of lime granules,

globose, .5 mm. diam., mounted on stout, conical, fluted white

stipes which arise from a white hypothallus; capillitium of

colorless threads, with large, angular, white lime-knots ; spores

violet-black in mass. On dead leaves.

Physarum melleum (Berk, and Br.) Mass. Sporangia scat¬

tered, yellow, globose, .5 mm. diam., mounted on short, stout,

furrowed white stipes, which terminate in a small, conical col¬

umella; hypothallus lacking; capillitium of colorless threads,

154 Wisconsin Academy of Sciences , Arts , and Letters.

with large, angular, white lime-knots; spores violet-black in

mass. On dead wood.

Physarum notabile Macbr. Sporangia gregarious, gray,

globose, .5 mm. diam., through stipitate, sessile, to plasmodio-

carpous forms, frequently in the same fructification; stipe,

when present, short, dark, tapering upward, furrowed ; capilli-

tium of colorless threads, with white lime-knots of variable

shape; spores violet-black in mass. On dead wood.

Physarum nucleatum Rex. Sporangia gregarious, white,

spherical, .5 mm. diam., stipe yellowish white, rough awl-

shaped ; capillitium dense, compact, and remarkably persistent,

with usually a shining ball of lime suspended in the central

portion of the capillitial net; lime-knots small, white, rounded;

spores violet-black in mass. On dead wood.

Physarum nutans Pers. Sporangia gregarious, grayish white,

subglobose, umbilicate below, nodding, .5 mm. diam. ; peridium

thin; stipes long, delicate, tapering decidedly upward, dark

below, white above; capillitium of delicate, colorless threads

with a few small, white lime-knots ; spores violet-black in mass.

On dead wood and bark.

Physarum polycephalum Schw. Sporangia confluent in clus¬

ters of 5-10, or thereabouts, the clusters in turn gregarious,

yellowish-gray, lobed, total height up to 2 mm. ; stipes reddish-

yellow, slender, weak, combined in clusters which accord with

the sporangial number; capillitium a branching network of

colorless threads with irregular yellowish lime-knots; spores

violet-black in mass. On leaves and dead wood.

Physarum psittacinum Ditm. Sporangia gregarious, irides¬

cent-blue, mottled with red-orange spots, globose, .5 mm. or

more in diam.; stipe orange-red, columnar, arising from a

hypothallus of the same color; capillitium a dense network of

yellowish threads, with many angular, orange lime-knots ;

spores violet-black in mass. On dead wood. Collected by E. M.

Gilbert, Hayward, Wis., August 1980.

Physarum sinuosum (Bull.) Weinm. (Plate I, fig. 4). Spo¬

rangia usually forming long, vein-like plasmodiocarps ; plasmo-

diocarps snow-white on upper portion, due to aggregated lime-

granules, gray below, sessile, breaking open by a longitudinal

Greene — Wisconsin Myxomycetes.

155

fissure; capillitium with numerous, large, white lime-knots;

spores violet-black in mass. On dead leaves and twigs.

Physarum variabile Rex. Sporangia scattered, dull yellow,

globose to egg-shaped or cylindric, .5 mm. or more in diam. ;

shortly stalked or sessile; peridium roughened; capillitium a

compact network of delicate, colorless threads, with irregular

white or yellowish-white lime-knots; spores violet-black in

mass. On dead leaves and wood.

Physarum virescens Ditmar. Sporangia heaped in groups of

a dozen or more, sessile, yeliow or greenish yellow, spherical to

elongate, less than .5 mm. diam. ; capillitium a delicate network

with small, irregular, yellow lime-knots; spores violet-black in

mass. On dead wood, moss or leaves. Collected by H. C. Greene,

Devil's Lake, Wis., July 1929.

Physarum viride (Bull.) Pers. Sporangia gregarious, various

shades of yellow or orange, subglobose, flattened below, .5 mm.

diam., nodding, borne on slender, tapering yellowish stipes,

which become red-brown toward the base; capillitium a loose

network of colorless threads with elongated yellow lime-knots ;

spores violet-black in mass. On dead wood.

Genus Craterium Trentepohl. This genus is closely allied

to Physarum , and is differentiated from it largely by the mode

of sporangial dehiscence. When the sporangia mature and

break open, a cup-like, persistent basal porton is left. The

sporangia generally exhibit a distinct lid, and the peridium

is generally of two layers.

Craterium leucocephalum (Pers.) Ditm. (Plate II, fig. 1).

Sporangia gregarious, white above, brown below, ovate or top¬

shaped, or definitely vasiform, .5 mm. diam., mounted on short,

stout stipes of the same color as the lower sporangium; spo¬

rangia often dehiscing rather irregularly; capillitium white,

with large lime-knots; spores violet-black in mass. On dead

leaves.

Craterium minutum (Leers) Fries. Sporangia gregarious,

brown below, with a distinct, usually convex lid lighter in col¬

or, goblet-shaped, .5 mm. diam., mounted on brown stipes which

arise from a circular hypothallus ; capillitium white, with large

lime-knots which are usually aggregated in a mass at the cen-

156 Wisconsin Academy of Sciences, Arts, and Letters .

ter of the cup; spores violet-black in mass. On dead leaves.

Collected by E. M. Gilbert, Hayward, Wis. August 1980.

Genus Leocarpus (Link) Rost. This genus includes a sin¬

gle species, L. fragilis (Dicks.) Rost.

Leocarpus fragilis (Dicks.) Rost. (Plate II, fig. 2). Spo¬

rangia gregarious, or closely clustered, shining red-brown to

various shades of yellow, egg-shaped, 2 mm. or more in length,

with weak, usually prostrate, white or yellowish stipes; capil-

litium of threads bearing closely clustered grayish-white lime-

knots; spores violet-black in mass. On dead wood, leaves, and

twigs.

Family Didymiaceae

Lime-deposits, when present, affecting the peridium only, or

sometimes the stipe also; capillitium of simple threads, usually

radiating from a central columella.

Genus Mucilago (Mich.) Adams. This genus includes a

single species, M . spongiosa (Leyss.) Morg.

Mucilago spongiosa (Leyss.) Morg. (Plate II, fig. 3).

Aethalium white, elongated, the shape depending upon the sub¬

stratum to a considerable degree, large, up to several cm.;

aethalium with a crusty, porous cortex; capillitium of darkish,

anastomosing threads; spores violet-black in mass. On living

herbaceous stems.

Genus Didymium (Schrad.) Fries. Sporangia distinct, not

forming aethalia; peridium covered with lime granules which

are distinctly crystalline, a feature serving to set the genus

apart from all others except Mucilago which differs in its aetha-

lioid condition. Capillitium of simple threads, radiating out¬

ward from the columella.

Didymium clavus (Alb. and Schw.) Rabenh. Sporangia gre¬

garious, grayish-white, discoidal, 1 mm. diam., the dark peri¬

dium frosted with lime-crystals above, naked and black below ;

sporangia mounted on slender, cylindrical, black stipes; col¬

umella lacking; capillitium of scantily branched pale brown

threads; spores violet-black in mass. On dead leaves.

Didymium melanospermum (Pers.) Macbr. Sporangia gre¬

garious, white, hemispherical, umbilicate below, up to 1 mm.

diam., mounted on short, stout, black stipes; columella large,

Greene — Wisconsin Myxomycetes.

157

hemispherical, dark-colored; capillitium of brown, sparingly

branched threads, often with thickenings; spores violet-black

in mass. On dead wood, and other debris.

Didymium nigripes (Link) Fries. (Plate II, fig. 4). Spo¬

rangia remotely gregarious, white, hemispherical, umbilicate

below, .5 mm. diam., mounted on slender black stipes, each

arising from a rounded black hypothallus ; columella globose,

dark-colored; capillitium of pale brown, scantily branched

threads ; spores violet-black in mass. On dead leaves and twigs.

Didymium squamulosum (Alb. and Schw.) Fries. Sporan¬

gia gregarious, white, globose, umbilicate below, up to 1 mm.

diam., mounted on stout, cylindrical, furrowed white stipes

each arising from a small, discoidal, white hypothallus; col¬

umella hemispherical, white; capillitium of pale, violet-brown

branching threads ; spores violet-black in mass. On dead leaves.

Genus Diderma Persoon. Sporangia ranging from plasmodio-

carps to distinct stipitate forms; peridium, as a rule, plainly

double ; outer wall calcareous or cartilaginous ; inner wall mem¬

branous, often widely separated from the outer ; columella usu¬

ally in evidence, but sometimes only rudimentary or lacking;

capillitial threads much as in Didymium; lime amorphous.

Diderma crustaceum Peck. (Plate II, fig. 5). Sporangia

closely crowded, and imbedded in the massed white hypothal¬

lus, white, globose, about 1 mm. diam., sessile ; outer sporangial

wall of lime, smooth, pure white, fragile, and distinct from the

bluish, membranous inner wall ; capillitium of purplish threads ;

spores violet-black in mass. On leaves, twigs, and bark.

Diderma effusum (Schw.) Morg. var. reticulatum Rost. Spo¬

rangia gregarious, white, rounded, sessile, somewhat flattened,

sometimes tending toward plasmodiocarpous, up to 1 mm.

diam.; inner peridium bluish white, distinct from the fragile,

lime-incrusted outer peridium; columella not well differentiat¬

ed; capillitium of short, colorless, scantily branching threads;

spores violet-black in mass. On dead leaves and twigs.

Diderma globosum Pers. Reported by Dean. Scarcely to be

differentiated from D. crustaceum Peck.

Diderma hemisphericum (Bull.) Hornem. Sporangia gre¬

garious, white, pronouncedly discoidal, about 1 mm. diam.,

158 Wisconsin Academy of Sciences , Arts, and Letters .

borne on stout white stipes, about 1 mm. high, each from a

small hypothallus; capillitium scanty, with colorless threads;

spores violet-black in mass. On dead leaves.

Diderma simplex (Schroet.) Lister. Sporangia gregarious,

or closely clustered, reddish-brown, globose, sessile, .5 mm.

diam., inner peridium not to be differentiated from the outer

wall of lime ; brownish hypothallus present ; columella not well

differentiated; capillitium scanty, of pale, sparsely branching

threads. Collected by H. C. Greene, Univ. of Wis. Greenhouse,

May 1930. On sphagnum and leaves.

Diderma spumaroides Fries. Sporangia closely clustered,

white, appearing grayish when lime is scanty, globose, sessile,

.5 mm. or slightly more in diam.; inner peridium closely ap-

pressed to the outer; white hypothallus usually present; col¬

umella white, small, globose; capillitium of branching brown

threads ; spores violet-black in mass. On dead leaves and bark.

Order Stemonitales

Spores in mass, blackish, violet-brown, or reddish-brown.

This order is characterized by the lack of lime in all genera ex¬

cept Diachaea, easily known by its stipes which are heavily

charged with lime. The capillitium in this order is thread-like

and branched, arising usually from a columella.

Family Stemonitaceae

Sporangia stalked (aethalioid in the case of Brefeldia) with

a definite columella from which the abundant and branching

threads of the capillitium arise; sporangium wall delicate and

usually disappearing at maturity; sporangia usually arising

from a common hypothallus.

Genus Brefeldia. This genus includes a single species, B .

maxima (Fries) Rost.

Brefeldia maxima (Fries) Rost. (Plate III, fig. 1). Spo¬

rangia combined in an aethalium, deep blackish brown ; aethali-

um large 3-30 cm. or more across, 5-10 mm. thick; capillitium

composed of dark brown threads expanded at regular intervals

into many-chambered vesicles; spores blackish in mass. On

dead wood, leaves, bark, etc.

Green — Wisconsin Myxomycetes.

159

Genus Stemonitis (Gleditsch) Rost. Sporangia cylindric

and stipitate with the abundant and flexuose capillitium spring¬

ing from all parts of the long columella; spores in mass black,

brown, or reddish brown, plainly in evidence in mature speci¬

mens, owing to the evanescent character of the sporangium

wall; capillitium covered by a delicate surface net.

Stemonitis ferruginea Ehr. Sporangia in dense clusters,

reddish-brown, cylindric, variable in height, from 5-15 mm.;

stipes black, one-half to one-third total height, arising from a

membranous hypothallus, and continuing through the greater

portion of the sporangium as a columella ; capillitium composed

of branching, brownish threads springing from the columella,

and connected with an outer surface network of rounded mesh¬

es ; spores red-brown in mass. On dead wood.

Stemonitis fused (Roth) Rost. (Plate III, fig. 2). Sporangia

clustered in tufts, blackish brown, cylindric, height variable,

from 5-20 mm.; stipes black, shining, about one quarter the

total height; arising from a common hypothallus, and contin¬

ued through the greater portion of the sporangium as a col¬

umella; capillitium of dark threads, springing from the col¬

umella, and ultimately giving rise to a close-meshed surface

net; spores blackish-brown in mass. On dead wood.

Stemonitis pallida Wingate. Sporangia gregarious, violet-

brown, cylindrical, blunt at apex, height about 5 or 6 mm.;

stipes short, black, polished, rising from a thin hypothallus,

and continued into the sporangium as a columella; capillitium

terminating in a close-meshed surface net ; spores violet brown

in mass. On dead wood. Collected by H. C. Greene, Devil’s

Lake, Wis., July 1929.

Stemonitis splendens Rost. Sporangia clustered, purple-

brown, cylindrical, height from 10-20 mm. ; stipes short, black,

polished, arising from a common, shining, silvery hypothallus ;

columella continuous through the sporangium; capillitium of

dark threads which ultimately give rise to a surface net with

rounded meshes ; spores purple-brown in mass. On dead wood.

Stemonitis webberi Rex. Reported for Wisconsin by Dean.

Original material not seen. Lister regards this as a variety of

S. splendens .

Genus Comatricha (Preuss) Rost. Sporangia usually

stalked, usually cylindrical, without capillitial surface net, and

160 Wisconsin Academy of Sciences , Arts , and Letters .

with peridium disappearing at maturity; spores in mass, vari¬

ous shades of brown to black. This genus merges easily into

Stemonitis, on the one hand, and Lamproderma on the other.

From Stemonitis it differs in the absence of a capillitial sur¬

face net, from Lamproderma in the evanescent peridium. The

genus is somewhat artificial and perplexing forms are repeated¬

ly met.

Comatricha flaccida Morg. Reported by Dean. Original ma¬

terial not seen by writer.

Comatricha longa Peck. Sporangia in tangled, prostrate

tufts, black, very long, up to several cm. ; stipes shining, black,

short, arising from a blackish hypothallus; columella black,

slender, disappearing below apex of sporangium; capillitium

of slender free threads extended stiffly outward from the col¬

umella, and often forking; spores black in mass. On dead

leaves, wood, etc.

Comatricha nigra (Pers.) Schroeter. (Plate III, fig. 3). Spo¬

rangia scattered, deep brown, globose to cylindrical, mounted

on long, slender, black, tapering stipes; height over all from

2-4 mm. ; columella continued upward into the sporangium and

becoming dissipated in the capillitial branching; capillitium of

repeatedly branching purple-brown threads with very few free

ends ; spores dark brown in mass. On dead wood.

Comatricha pulchella (Bab.) Rost. Sporangia scattered, pale

brown, ovate, mounted on short black stipes, very small, total

height about 1 mm. ; columella attaining almost to the apex of

the sporangium; capillitium a dense network of slender brown

threads ; spores brown in mass. On dead leaves.

Comatricha typhoides (Bull.) Rost. Sporangia gregarious

or scattered, silvery, becoming brown, cylindric, mounted on

black stipes which are “clothed with the silvery, membranous

continuation of the sporangium wall” ; columella reaching al¬

most to summit of the sporangium ; spores deep brown in mass.

On dead wood.

Comatricha typhoides (Bull.) Rost. var. similis Lister. Spo¬

rangia as in C. typhoides , except that stipes have no silvery

sheath, and spores have slightly different markings. Collected

by E. M. Gilbert, Hayward, Wis., July 1930.

Greene — Wisconsin Myxomycetes.

161

Genus Diachaea Fries. Sporangia distinct, stipitate, globose

or cylindric, peridium fragile, iridescent ; stipes heavily charged

with lime; capillitium much as in Comatricha.

Diachaea leucopoda (Bull.) Rost. (Plate III, fig. 4). Spo¬

rangia gregarious, bluish-purple, with a metallic sheen, cylin¬

dric, mounted on white, columnar stalks, which arise from a

continuous vein-like hypothallus and .are continued as columel-

lae through the height of the sporangia; capillitium springing

from the columella as a network of slender fiexuous brown

threads; spores violet-brown in mass. On living and dead

leaves and twigs.

Family Lamprodermaceae

This family is characterized by the fact that the capillitium

is borne chiefly from the tip of the columella.

Genus Enerthenema Bowman. Sporangia distinct, stipi¬

tate, gregarious, small, globose, the apex of the trans-sporan-

gial columella expanded into a disk from which the capillitial

threads spring.

Enerthenema papillatum (Pers) Rost. (Plate III, fig. 5).

Sporangia gregarious or scattered, blackish brown, spherical,

approaching 1 mm. diam., mounted on short, black stipes, which

pass entirely through the sporangia as columellae which emerge

at the top as flattened, shining disks; sporangium wall evane¬

scent ; capillitium of long, slender, rather stiff, scantily

branched threads, springing chiefly from the expanded disk of

the columella; spores blackish brown in mass. On dead wood.

Genus Clastoderma Blytt. This genus contains a single

species, C. debaryanum Blytt.

Clastoderma debaryanum Blytt. (Plate IV, fig. 1). Spo¬

rangia more or less gregarious, brown, globose, borne on slen¬

der brown stipes, very minute, total height not more than 1

mm., sporangia proper .1-.2 mm.; peridium not persistent ex¬

cept for plates which adhere to the ultimate branches of the

capillitium; capillitium composed of stiff, forking threads;

spores brown in mass. Collected by E. M. Gilbert, Hayward,

Wis., August 1930.

Genus La m proper m a Rost. Sporangia distinct, stipitate,

characterized by the. fairly persistent, shining, metallic peridi-

162 Wisconsin Academy of Sciences, Arts, and Letters .

um which, when shed, leaves a ring around the base of the col¬

umella ; columella short, stout, with the capillitium usually aris¬

ing plainly from the apical region.

Lamproderma violaceum (Fries) Rost. (Plate IV, fig. 2).

Sporangia gregarious, metallic purple, or bronze, globose, .5

mm. or slightly more in diam. ; stipe dark, moderately stout,

usually about equal to sporangium ; capillitium of dark, branch¬

ing threads, anastomosing to form an open network; spores

violet-black in mass. On dead wood and leaves.

Order Cribrariales

Spores in mass usually pale brown or yellowish, rarely pur¬

plish or violaceous; sporangia ranging from aethalioid forms

to distinct, stipitate forms, peridium membranous, without

lime; capillitium lacking.

Family Tubiferaceae

Fructifications of aethalia or distinct sporangia; sporangia

tubular, irregularly dehiscent, rupturing at the apex.

Genus Lindbladia Fries. This genus includes a single spe¬

cies, L. effusa (Ehr.) Rost.

Lindbladia effusa (Ehr.) Rost. (Plate IV, fig. 4). Spo¬

rangia closely crowded in a single layer, or superimposed to

form an aethalium, dull brown, sessile, tubular, about .5 mm.

short diam.; fruiting masses often very extensive, up to 25

cm. or more; capillitium lacking; spores deep yellow-brown in

mass. On dead wood. (esp. coniferous.)

Genus Tubifera Gmelin. Sporangia red-brown and angled

by mutual pressure, tubular; hypothallus prominent; capilliti¬

um lacking; spore mass red-brown.

Tubifera ferruginosa (Batsch) Macbr. (Plate IV, fig. 3).

Sporangia densely crowded, red-brown, cylindrical, angled by

mutual pressure, up to 3 mm. high ; capillitium lacking ; spores

red-brown in mass. On dead wood.

Tubifera stipitata Macbr. Sporangia similar to those of T.

ferruginosa, except that they are clustered on a dark brown

hypothallus which has the form of a stout stipe, 2-3 mm. high.

On dead wood.

Greene — Wisconsin Myxomycetes.

163

Family Reticulariaceae

Sporangia indistinctly defined and aggregated in aethalia on

a common hypothallus and covered by a common cortex; pseu-

do-capillitium composed of interwoven or fragmentary sporang-

ial walls ; spore mass brown or yellowish.

Genus Reticularia Bull. This genus is represented by a

single species, R . lycoperdon Bull.

Reticularia lycoperdon Bull. (Plate IV, fig. 5). Aethalium

subglobose, silvery- white, 2-5 cm. diam. ; solitary, arising from

a well-developed white hypothallus; true capillitium lacking;

pseudo-capillitium formed from remnants of sporangial wall,

appearing in blown specimens as a mass of strands arising

from the hypothallus; spores red-brown in mass. On dead

wood.

Genus Enteridium Ehr. Sporangia fused to form aetha¬

lia ; and covered by a thin, evanescent membrane ; pseudo-capil¬

litium composed of the sporangial walls interwoven to form a

network of broad, membranous bands.

Enteridium splendens Morg. (Plate V, fig. 1). Aethalium

cushion-like, lobed, covered by a thin, smooth, shining brown

cortex, from 1-6 cm. diam. ; hypothallus white ; capillitium none,

the sporangial wall perforate and interwoven; spores reddish-

brown in mass. On dead wood.

Genus Dictydiaethalium Rost. This genus is represented

by a single species, D. plumbeum (Schum.) Rost.

Dictydiaethalium plumbeum (Schum.) Rost. (Plate V, fig.

2). Aethalium thin, flat, smooth, olivaceous, up to several cm.

broad, in section showing columnar, hexagonal, perforated spo¬

rangia, with six threads extending from base to apex, where

the sporangium is entire in the form of a cap; hypothallus

white, prominent; spores in mass dull yellow. Collected by E.

M. Gilbert, Hayward, Wis. 1921, and by Charles Drechsler,

Park Falls (1917?).

Family Cribrariaceae

Sporangia distinct and stipitate; peridium usually well-de¬

fined, but sometimes lacking as a continuous layer, and, at any

rate, opening above to form a meshed network with nodes of

164 Wisconsin Academy of Sciences , Arts , and Letters.

varied shape; spore mass various shades of yellow to brown,

rarely purplish.

Genus Cribraria (Pers.) Schrader. Sporangia distinct,

stipitate, the membranous peridium forming a basal cup, more

or less developed ; perforated above to form a meshed network ;

spores in mass, ochraceous or violaceous.

Cribraria aurantiaca Schrader. (Plate V, fig. 3). Sporangia

gregarious, yellowish-brown, globose, approaching 1 mm. diam. ;

stipe brown, in length about twice the diam. of the sporangium ;

cup large, indented; net with irregular, expanded nodes, con-

col or ous with the yellow-brown-cup; spores in mass yellowish-

brown. On dead wood, especially coniferous wood.

Genus Dictydium (Schrad.) Rost. This genus includes a

single species, D . cancellatum (Batsch) Macbr.

Dictydium cancellatum (Batsch) Macbr. (Plate V, fig. 4).

Sporangia gregarious, reddish- or purplish-brown, of peculiar

bell-like contour, or subglobose, nodding, about .5 mm. diam.,

mounted on dark red-brown stipes ; capillitium none ; sporangi¬

um wall ribbed, with ribs connected by slender cross-threads,

spores reddish- or purplish-brown in mass. On dead wood.

Order Lycogalales

This order includes a single family, Lycogalaceae , and the

single genus, Lycogala Micheli.

Genus Lycogala Micheli. Aethalia with membranous peri-

dia, resembling miniature puff-balls; capillitium of irregular

tubules extending inward from the peridium; spore mass pale

pinkish or ashen, becoming sordid or yellowish.

Lycogala epidendrum (Buxb.) Fries. (Plate V, fig. 5).

Aethalia single or clustered, olivaceous, spherical, or angled by

mutual pressure when clustered; inconspicuously warted, 3-10

mm. diam.; peridium tough and persistent; capillitium of

branching, wrinkled tubules which extend inward from the

peridium ; spores ashen or yellowish in mass. Common on dead

wood.

Lycogala exiguum Morg. Perhaps only a form of L. epiden¬

drum differing in its small (2-5 mm. diam.), blackish, gregari-

Greene — Wisconsin Myxomycetes.

165

ous aethalia, which are never closely clustered as in typical

L. epidendrum.

Ly cogala flavo-fuscum (Ehr.) Rost. Aethalia single, or clus¬

tered in small groups, purplish-gray, tending toward spherical,

usually very large, up to 4 cm. diam. ; capillitium consisting of

branching, irregular tubules, arising from the peridium and

continued inward, enclosing within its meshes, rounded, yel¬

lowish, protoplasmic vesicles ; spores grayish in mass. On dead

wood.

Order Trichiales

Spore mass usually pale, brownish or yellowish, or with a

reddish tinge ; sporangia membranous, without lime, plasmodio-

carpous, or distinct, sessile or stipitate, scattered or crowded;

capillitial threads attached to wall or free, sculptured, single

or combined into a net.

Family Perichaenaceae

Capillitial threads smooth or variously roughened, not form¬

ing a network, attached at one end to sporangium wall.

Genus Ophiotheca Currey. Fructification characteristical¬

ly of flexuose plasmodiocarps, the thin peridium breaking open

irregularly ; capillitium of roughened threads ; spores yellowish

in mass.

Ophiotheca wrightii Berk, and Curt. Plasmodiocarps brown¬

ish to black, usually bent or ring-shaped ; .2-1 mm. diam. ; capil¬

litium of long threads studded with prominent spines; spores

yellow in mass. On dead wood and bark, especially the inner

bark of fallen trees.

Genus Perichaena. Sporangia flattened, round or angled

when closely clustered; peridium thick, breaking open with a

more or less definite lid; capillitium of warted, yellowish

threads ; spores yellow in mass.

Perichaena corticalis (Batsch) Rost. Sporangia gregarious,

chestnut brown, sessile, depressed hemispherical; .5-1 mm.

diam., opening by a lid, the outlines of which are clearly marked

in mature specimens; capillitium of slender threads, attached

to the lid ; spores bright yellow in mass. On bark of dead trees.

166 Wisconsin Academy of Sciences , Arts, and Letters .

Perichaena depressa Libert. (Plate VI, fig. 1). Sporangia

crowded, angled by mutual pressure, chestnut brown, sessile,

flattened, opening by a definite lid ; capillitium of slender yellow

threads ; spores bright yellow in mass. Closely allied to P . cor-

ticalis . On the inside bark of logs.

Family Arcyriaceae

Capillitium forming a more or less definite net, usually at¬

tached below; capillitial threads variously roughened or sculp¬

tured, but lacking a smooth spiral sculpturing; spores pale.

Genus Lachnobolus. This genus is set apart from Arcyria

on the basis of the non-elastic capillitium, and by the fact that

the upper peridium is somewhat persistent, not completely

evanescent as in Arcyria .

Lachnobolus occidental is Macbr. Sporangia scattered or

crowded, metallic rose in color, later becoming brownish, usu¬

ally sessile, globose to ellipsoidal, about .5 mm. diam. ; spo¬

rangium wall breaking away above, leaving a membranous,

lobed cup below; capillitium a network of warted threads, not

expanding as in Arcyria ; spores brownish flesh-colored in mass.

On dead wood.

Genus Arcyria (Hill) Pers. Sporangia ovoid or cylindric,

stipitate, the upper portion of the peridium completely evane¬

scent; capillitium forming an elastic net; color of spores in

mass variable, pale, yellow, greenish or reddish.

Arcyria cinerea (Bull.) Pers. Sporangia scattered, or fairly

closely clustered, ashen gray, short-cylindrical, 2-4 mm. high,

cup small; stipe 1-2 mm. high, dark gray to black; capillitium

dense ; spores ashen gray in mass. On dead wood.

Arcyria denudata (Linn.) Sheldon. Sporangia crowded or

gregarious, bright red, becoming brownish with age, ovoid, ta¬

pering upwards, 1-2 mm. high; cup well-defined; stipe about

half total height of sporangium, and of same color; capillitium

fairly lax, but firmly attached to cup; spores reddish in mass.

On dead wood.

Arcyria incarnata Persoon. Similar to A. denudata, except

that the sporangia are of a more delicate rosy hue, the stipe is

usually shorter, and the mature capillitium is so loosely at-

Greene — Wisconsin Myxomycetes.

167

tached to the cup that a slight current of air suffices to blow it

away.

Arcyria magna Rex. Reported for Wisconsin by Dean, but

probably rare and not likely to be met with. Lister regards

this as a mere variation of A . oerstedtii, a form not reported

for Wisconsin.

Arcyria nutans (Bull.) Grev. (Plate VI, fig. 2). Sporangia

closely clustered, pale yellow, cylindrical, 2 mm. high when un¬

expanded; cup very small; stipe short or lacking; capillitium

becoming tremendously expanded (up to 3 cm. or more), giving

rise to the drooping yellow plumes so characteristic of the

species.

Family Trichiaceae

Capillitium spirally sculptured, the threads free or in a loose

net; spore mass of various shades through yellow to reddish.

Genus Hemitrichia Rost. Capillitium a loose, branching

net of spirally sculptured threads centrally attached ; sporangia

plasmodiocarpous or distinct, sessile or stipitate.

Hemitrichia clavata (Pers.) Rost. Sporangia gregarious,

scattered or crowded, yellowish, the upper portion of the peri-

dium breaking away to leave a cup-shaped structure below,

mounted on slender, brownish stipes; sporangia usually large,

up to 3 mm. high, but sometimes much smaller; capillitium of

slender threads, branched to form a net, yellow, the threads

sculptured with spiral bands ; spores yellow in mass. On dead

wood.

Hemitrichia serpula (Scop.) Rost. Plasmodiocarps often of

rather wide extent, branching freely, reticulate, tubular, yellow,

about .5 mm. wide ; spores yellow in mass. On dead wood.

Hemitrichia stipata (Schw.) Macbr. Sporangia crowded,

superimposed, of a shining metallic copper color, cylindrical,

mounted on short, red-brown stipes ; upper portion of sporangi¬

um wall falling away, leaving a cup as in Arcyria ; capillitium

of copper-colored threads branched to form a net; spores red¬

dish-brown in mass. On dead wood.

Hemitrichia vesparium (Batsch) Macbr. (Plate VI, fig. 3).

Sporangia clustered, usually with the dark stipes welded to-

168 Wisconsin Academy of Sciences, Arts, and Letters.

gether, dark red, clavate, 1 mm. or more high; peridium metal¬

lic gray; capillitium of reddish, sparingly branched, spirally

sculptured threads ; spores red in mass. On dead wood.

Genus Trichi a (Haller) Rost. Sporangia distinct, sessile

or stipitate; capillitium of free, spirally sculptured threads,

called elaters ; yellowish ; peridium membranous ; spores in mass

various shades of yellow, except in T. lateritia (not reported

for Wisconsin) where they are brick red.

Trichia decipiens (Pers.) Macbr. (Plate VI, fig. 4). Spo¬

rangia gregarious, shining olive, depressed egg-shaped, often

breaking open above in regular fashion, about .7-. 8 mm. diam.,

mounted on olive stipes of variable length ; capillitium consist¬

ing of olive-colored, much tapered elaters, sculptured with 3-5

spiral bands ; spores olivaceous or yellowish in mass. On dead

wood.

Trichia favoginea (Batsch) Pers. Sporangia crowded, olive-

yellow, cylindric, .5 mm. or more in diam. ; capillitium of cylin¬

drical elaters with 3-5 spiral bands, usually escaping from spo¬

rangia at maturity, and hanging above them as a woolly yellow

mass ; spores bright yellow in mass. On dead wood.

Trichia persimilis Karst. Sporangia somewhat crowded, as

a rule, bright yellow or brownish, sessile, seated on a thin hypo-

thallus, .5-.8 mm. diam., capillitium yellow, consisting of spiral¬

ly sculptured elaters ; spores bright yellow in mass. On dead

wood.

Trichia scabra Rost. Sporangia very closely crowded on a

common hypothallus, orange-brown, globose, .7-.8 mm. diam.;

capillitium an orange-yellow mass of elaters, sculptured with

3-4 closely wound spiral bands ; spores orange-yellow in mass.

On dead wood.

Trichia varia (Pers.) Rost. Sporangia gregarious or crowd¬

ed, yellowish, globose to egg-shaped, up to 1 mm. diam., sessile

or with a short, stout black stipe ; capillitium of long, irregular

threads, marked with only 2 spiral bands ; spores yellow in

mass. On dead wood.

170 Wisconsin Academy of Sciences, Arts, and Letters.

Plate I

Fig. 1. (a) Sporophores of Ceratiomyxa fruticolosa, X 35.

(b) Portion of sporophore wall, showing attachment of spore

(after Lister), X 250.

Fig. 2. (a) Aethalium of Fuligo septica f. ovata, X 1.

(b) Capillitium, X 300.

(c) Spore, X 625.

Fig. 3. (a) Sporangia of Badhamia utricularis, X 5.

(b) Capillitium, X 150.

(c) Spore, X 500.

Fig. 4. (a) Plasmodiocarp of Physarum sinuosum, X 3.

(b) Capillitium, X 250.

(c) Spore, X 400.

Fig. 5. (a) Sporangia of Physarum leucopus, X 25.

(b) Capillitium and spores, X 350.

TRANS. WIS. ACAD., VOL. 27

PLATE I

172 Wisconsin Academy of Sciences , Arts , and Letters.

Plate II

Fig. 1. (a) Sporangia of Craterium leucocephalum, X 20.

(b) Capillitium, X 250.

(c) Spores, X 500.

Fig. 2. (a) Sporangia of Leocarpus fragilis, X 5.

(b) Capillitium, X 250.

(c) Spore, X 500.

Fig. 3. (a) Aethalium of Mucilago spongiosa , X 1.

(b) Capillitium, X 150.

(c) Spore, X 500.

(d) Lime crystals and portion of sporangial wall, X 500.

Fig. 4. (a) Sporangia of Didymium nigripes, X 15.

(b) Blown sporangium, showing globose columella and capillitium,

X 30.

(c) (d) Spores and lime crystals, X 500.

Fig. 5. (a) Sporangia of Diderma crustaceum, X 15.

(b) Capillitium, X 150.

(c) Spore, X 400.

TRANS. WIS. ACAD., VOL. 27

PLATE II

174 Wisconsin Academy of Sciences , Arts, and Letters .

Plate III

Fig. 1. (a) Aethalium of Brefeldia maxima , X Vz

(b) Capillitium, X 100.

(c) Spore, X 400.

Fig. 2. (a) Sporangia of Stemonitis fusca, X 2.

(b) Capillitium, X 150.

(c) Spore, X 500.

Fig. 3. (a) Sporangia of Comatricha nigra, X 12.

(b) Capillitium, X 450.

(c) Spore, X 750.

Fig. 4. (a) Sporangia of Diachaea leucopoda, X 20.

(b) Capillitium, X 150.

(c) Spore, X 750.

Fig. 5. (a) Sporangia of Enerthenema papillatum, X 20.

(b) Capillitium and spores, X 35.

(c) Spore, X 500.

PLATE III

TRANS. WIS. ACAD., YOD. 27

176 Wisconsin Academy of Sciences , Arts, and Letters .

Plate IV

Fig. 1. (a) Sporangium of Clastoderma debaryanum, X 45.

(b) Section through sporangia! head showing capillitium, X 75.

(c) Spore, X 600.

Fig. 2. (a) Sporangia of Lamproderma violaceum, X 25.

(b) Capillitium, X 500.

(c) Spore, X 600.

Fig. 3. (a) Sporangia of Tubifera ferruginosa, X 5.

(b) Spores, X 800.

Fig. 4. (a) Aethalium of Lindbladia effusa , X 1.

(b) Individual sporangia, X 4.

(c) Spore, X 1000.

Fig. 5. (a) Aethalium of Reticularia lycoperdon, X 1.

(b) Pseudo-capillitium, X 50.

(c) Spore, X 600.

Q r

TRANS. WIS. ACAD., VOL. 27

PLATE IV

178 Wisconsin Academy of Sciences , Arts, and Letters .

Fig. 1. (a)

(b)

(c)

Fig. 2. (a)

(b)

(c)

(d)

Fig. 3. (a)

(b)

(c)

Fig. 4. (a)

(b)

(c)

Fig. 5. (a)

(b)

(c)

Plate V

Aethalium of Enteridium splendens, X 1.

Pseudo-capillitium, X 50.

Spore, X 600.

Aethalium of Dictydiaethalium plumb eum, X 1.

Individual sporangia (after Lister), X 20.

Single sporangium, showing perforated walls, X 50.

Spores, X 500.

Sporangia of Cribraria aurantiaca, X 8.

Sporangial head, showing net and nodes, X 50.

Spore, X 1000.

Sporangia of Dictydium cancellatum, X 15.

Sporangial ribs, X 500.

Spores, X 800.

Aethalia of Ly cog ala epidendrum , X 1.

Capillitial tubules, X 400.

Spore, X 1000.

TRANS. WIS. ACAD., VOL. 27

PLATE V

2

180 Wisconsin Academy of Sciences , Arts , and Letters .

Plate VI

Fig. 1. (a) Sporangia of Perichaena depressa, X 15.

(b) Capillitium and portion of sporangial wall, X 500.

(c) Spore, X 600.

Fig. 2. (a) Sporangia of Arcyria nutans , X 5.

(b) Section of capillitium, showing twisted threads, X 100.

(c) Capillitium, X 350.

(d) Spore, X 600.

Fig. 3. (a) Sporangial cluster of Hemitrichia vesparium, X 6.

(b) Capillitial threads, X 500.

(c) Spore, X 500.

Fig. 4 (a) Sporangia of Trichia decipiens , X 12.

(b) Single elater, X 150.

(c) Tip of elater, X 400.

(d) Spore, X 500.

Greene — Wisconsin Myxomycetes .

181

TRANS. WIS. ACAD., VOL. 27

1

PLATE VI

NOTES ON PARASITIC FUNGI IN WISCONSIN. XIX.

J. J. Davis

The summer of 1930 was hot and dry, which had a marked

deterrent effect on the development of parasitic fungi.

Synchytrium pulvereum Davis, on Laportea canadensis ,

having been found to bear summer sori similar to those of S.

cellulare on Boehmeria cylindrica , is now referred to that spe¬

cies and the binomial reduced to synonymy.

Record has been made of the presence of Peronospora mel-

ampyri (Bucholtz) in Wisconsin at Friendship and Radisson (as

Plasmopara melampyri Bucholtz, Notes XVI, pp. 287-8). In

July 1931 it was found at Washington Island with oospores,

mostly immature, in the leaves. Those measured were globose,

27-33/a in diameter; oogonia 40-50/a. The conidia are strongly

fuscous.

In “Notes” XI, pp. 294-295, record was made of the occur¬

rence of Doassansia sagittariae (West.) Fisch on Lophotocar-

pus calycinus at Blue River. The collection was a small one,

there being but few host plants. In 1930 it was found in abun¬

dance at a station on the bottom lands of the Mississippi river

near Glen Haven. As at the first locality no Doassansia was

found on Sagittaria and as in the first collection the sori are

often irregular. Field evidence points to close host adaptation

as might be expected of parasites of this character.

In 1931 plants of Amphicarpa monoica were exposed to in¬

fection in the greenhouse from overwintered Puccinia on An-

dropogon from two localities without result. One of the locali¬

ties was visited later and Aecidium on Comandra umbellata was

found to be abundant there.

Pestalozziella subsessilis Sacc. & Ell. was recorded in the

first supplementary list (1894) on the authority of Dr. Tre-

lease who had found it at Madison on Geranium maculatum.

It was not seen by the writer until May, 1930, when it was

found at Viroqua on the same host. Examination of this col¬

lection shows that the sporules are formed in definite pycnidia

in which there is apparently no ostiole, the spore discharge be-

184 Wisconsin Academy of Sciences, Arts, and Letters .

ing through rupture. The genus therefore should find place

in the Sphaerioidaceae. Notes on spore measurements read

17-80 X 5-10/a.

A specimen on an Aster of the paniculatus group collected

May 18, 1929 bears sporules some of which exceed 80/a in length

but has been referred to Septoria astericola Ell. & Ev. be¬

cause they are but about 1/a in diameter. This species develops

in spring, S. atropurpurea Pk. in midsummer.

A specimen on Solidago latifolia from Kenosha Co. (July 4,

1892) was recorded in the Supplementary list as Septoria at¬

ropurpurea Pk. It has the following characters: spots circu¬

lar to angular, 1-2 mm. or more elongate up to 4 mm. in length,

dark purple above, pale below, often confluent and sometimes,

by death of intervening tissue, forming pale brown areas;

pycnidia epiphyllous, few, scattered, black, 60-80/a in diameter ;

sporules straight or somewhat curved, 37-66 X An¬

other specimen (Kenosha Co., June 10, 1894) is more mature

and bears larger spots, 4 mm. in diameter, which become paler

in the center.

Stigmatea robertiani Fr. on Geranium robertianum. Fish

Creek. The material is not mature but appears to be of this

species.

The Cercospora that occurs on Smilax hispida in Wisconsin

was refered to C. mississippiensis Tracy & Earle in the

fourth supplementary list and in the provisional list. In

“Notes” X this was referred to C. SMILACIS Thuem. following

Peck. Solheim refers it to C. petersii (B. & C.) Atk. which

appears to be the proper designation.

Additional Hosts

A Synchytrium occurring in small quantity on Fragaria vir-

giniana at Superior has been referred to S. AUREUM Schroet.

It is much like the form on Geum.

Bremia lactucae Regel. On Lactuca villosa. Madison.

Plasmopara halstedii (Farl.) Berl. & De Toni. On Eupa-

torium purpureum . Viroqua.

Peronospora grisea Unger. In a collection of Veronica ser-

pyllifolia from Reedsburg one of the plants bears a little of the

mildew on the upper part of the stem.

Davis — Parasitic Fungi in Wisconsin . XIX,

185

Peronospora polygoni Thuem. On Polygonum Convolvulus ,

Lancaster. V. H. Young & J. J. Davis.

PUCCINIA HIERACII (Schwm.) Mont. On Agoseris cuspidata.

Pine Bluff. (N. C. Fassett).

Stagonospora atriplicis (West.) Lind. On Spinacia olera-

cea (Cult.). Madison. This is S. spinaciae Ell. & Ev., which

does not seem to me to be distinct. The sporules in this collec¬

tion are 14-24 X 6-8//,, 1-3 septate.

Septoria hellianthi Ell. & Kell. On Helianthus scaberri-

mus . Spring Green.

Colletotrichum malvarum (A. Br. & Casp.?) Southworth.

On Althaea (Cult.). Baraboo. (L. R. Jones, 1911).

Phyllachora graminis (Pers.) Fckl. On Elymus striatus.

Madison.

Septoria rudbeckiae Ell. & Hals. On Rudbeckia subtomen-

tosa on the Wisconsin river bottom lands in Iowa county. In

this collection the white arid portion constitutes most of the

spot which has a dark purple border. Similar spots, except that

the border is brown, are usual on Rudbeckia laciniata but not

on R. hirta. The development of the pycnidia on the arid spots

is often poor.

Of a collection on twigs of Viburnum Lentago made at Arena

June 4, 1929 the following notes were made: Acervuli various

in extent on the young growth of the season often extending

up the petioles, sometimes on the principal veins very excep¬

tionally on the lamina, subcuticular, discoid ; conidia soon erum-

pent in white masses, hyaline, cylindrical to fusoid, usually

more or less acute at the proximal end, 12-23 X 31^-6/x. This

has been provisionally referred to Gloeosporium cingulatum

Atk. Usually death of some of the young leaves results.

An Ovularia on leaves on Phalaris arundinacea referred to

0. pulchella (Ces.) Sacc. has been collected at Spring Green.

Glomerularia corni Pk. var. lonicerae Pk. On Lonicera

tatarica. Madison. I have seen no record of the occurrence

of this parasite on a host of foreign origin. The plants were

growing, without cultivation, on the railroad right of way.

Dearness & House consider the form on Lonicera to be spe-

186 Wisconsin Academy of Sciences , Arts , and Letters .

cifically distinct. ( New York State Museum Bulletin . Report

of the Botanist for 1921 , p. 85.)

Cercospora fingens Davis. A scanty collection on Thalic-

trum dioicum from Mazomanie. Amphigenous.

Ramularia virgaureae Thuem. On Solidago patula. Conidia

catenulate. Septation and catenulation often seem to be degrees

of the same process.

Cercospora davisii Ell & Ev. On Melilotus officinalis.

Spring Green. While this parasite is common on Melilotus alba

in Wisconsin it is much less frequent on M. officinalis. In this

collection the leaflets bear also small arid barren spots re¬

sembling those caused by Phyllosticta decidua Ell. & Kell,

suggesting a prior infection by another parasite.

In Mycologia 21: 804 et seq. (1929) Horsfall reports results

of examination of Cercospora zebrina Pass, on Trifolium, C.

davisii E. & E. on Melilotus alba and C. medicaginis Ell. & Ev.

on Medicago in which he failed to find definite morphological

characters by which to separate them and proposed that they

be united as a single species. In view of the variability of

conidiophoral and conidial characters in this genus perhaps it

is better to await the results of cross inoculation work and com¬

parison of ascigerous stages, if such exist, before decision.

Cercospora antipus Ell. & Hoi. On Lonicera dioica. Wash¬

ington Island.

Cercospora galii Ell. & Hoi. On Galium triflorum. Solon

Springs.

Alternaria herculea (Ell. & Mart.) Elliott. On Brassica

nigra. Gratiot.

Entyloma compositarum Farl. On Bidens vulgata. Big

Bend.

Aecidium plantaginis Burrill. The aecial stage of Uro-

myces seditiosus Kern was found on Plantago aristata at

Avoca May 28, 1929.

Puccinia bartholomaei Diet. The aecial stage, Aecidium

JAMESIANUM Pk. on Asclepias tuberosa. Ferry bluff, Sauk Co.

Puccinia canaliculata (Schw.) Lagh. II III on Cyperus

esculentus . Gratiot.

Davis — Parasitic Fungi in Wisconsin . XIX.

187

Since the connection of Aecidium on Erigeron with a Puc¬

ci nia on Carex was shown by Arthur all aecia on Erigeron in

Wisconsin have been referred to Puccinxa caricis-erigerontis

Arth. ( Journ . Mycol. 8: 58-4) now considered to be a race of

P. extensicola Plowr. which develops aecia on Aster and Soli-

dago as well. Of the two rusts on Cyperus in Wisconsin Puc¬

cinxa canaliculata (Schw.) Lagh. and P. cyperi Arth. the

former had been found to develop aecia on Xanthium (Arthur,

Journ. Mycol. 12: 23) but the aecial host of the latter has been

unknown. In conversation with Dr. H. S. Jackson he expressed

the opinion that P. cyperi Arth. develops its aecial stage on

Erigeron. Recalling that abundant development of aecia on

Erigeron canadensis had been observed in localities where Cy¬

perus was abundant rusted material of Cyperus Schweinitzii

bearing P. cyperi was overwintered in the open and the fol¬

lowing spring plants of Erigeron canadensis were brought into

the greenhouse and infected from the overwintered material

resulting in abundant aecia while the controls remained nor¬

mal. This Aecidium is quite similar to the one on Erigeron

connected with Puccinia extensicola Plowr., the only differ¬

ence that has been observed being a tendency to thickening of

the spore wall at or near the apex as was pointed out by Dr.

Arthur. The character is most readily seen when the spores

are concatenate and hence are seen in side view. A collection

on Erigeron ramosus from Spring Green is also referred to this

species.

Telial material from the same station has since been used to

infect Erigeron annuus in the greenhouse. This indicates that

there is no physiological difference between the rust occurring

on Euerigeron and that on what is considered by some to be

the distinct genus Leptilon.

Although Puccinia karelica Tranz. has been collected in

Wisconsin on Carex the aecial stage Aecidium trientalis

Tranz. was not found until 1930 when it was collected on Trien¬

talis americana in a large swamp near Cedarburg by A. M.

Fuller and the writer.

Puccinia violae (Schum.) DC. Aecia on Viola sagittata.

Arena.

A collection on Asplenium acrostichoides from Solon Springs

is presumably Uredinopsis copelandi Syd. The fronds are

188 Wisconsin Academy of Sciences , Arts , and Letters.

well infected but there are scarcely any uredospores and the

teliospores do not furnish distinctive characters. Another col¬

lection on the same species of host from Haugen however bears

both kinds of spores.

Puccinia menthae Pers. On Monarda didyma (Cult.).

Madison. (Sam Chechik.)

Puccinia xanthii Schw. On Ambrosia psilostachya. Spring

Green.

The aecial stage of Coleosporium solidaginis (Schw.)

Thuem. occurs on needles of young planted trees of Finns resi -

nosa in Peninsula State Park.

The pre-sclerotial stage of Sclerotium deciduum Davis oc¬

curred at Viroqua on Ranunculus septentrionalis.

Additional Species

Synchytrium fulgens Schroet. The summer spore stage

on Oenothera biennis was found under a railroad bridge at

Browntown.

Sclerotinia vaccinii Wor. On Vaccinium macrocarpon.

Cranmoor. (E. E. Honey).

Phyllosticta podophylli (Curt.) Wint. On Podophyllum

peltatum. Big Bend.

A dead fallen leaf of Pinus Strobus from “the elephant's

back" near “the dells" in Adams county bears what is probably

Vermicularia libertiana Roum. The conidia are somewhat

long, 10-13/a, and usually narrower and the bristles range up to

130/a in length. That this is parasitic is doubtful.

A small collection on a twig of Pinus Banksiana from Goth¬

am, June 18, 1930, shows on dead needles depressed globose

pycnidia 175-200 X 135/a in which develop deep brown (black

in mass) fusoid-oblong sporules which became uniformly tri-

septate, 16-20 X 4-7/a; basidia indistinct. The uninfected

needles on the twig were living. This probably bears relation

to Hendersonia foliicola (Berk.) Fckl. with which it has

been filed.

Septoria fumosa Pk. A collection on leaves of Solidago

serotina from Readstown, May 23, 1930, is referred to this spe-

Davis — Parasitic Fungi in Wisconsin . XIX.

189

cies. The spots are greyish brown, paler below, more or less

angular and limited by the veinlets, sometimes confluent, 1-5

mm. in diameter; pycnidia epiphyllous, scattered, subepider-

mal, sometimes imperfect distally, about 100/* in diameter;

sporules hyaline, curved, 50-75 X 2/*. Septoria davisii Sacc. is

probably not distinct from this.

Septoria cynoglossi n. sp. Spots definite, orbicular to irregu¬

lar, brown, paler below, 2-5 mm. in diameter ; pycnidia epiphyl¬

lous, scattered, rather thin-walled but usually with a more or

less prominent black thickening around the pore, 50-80/* in

diameter; sporules straight, 20-30 X %/*. On Cynoglossum

boreale , Winneboujou, Wisconsin, August 9, 1930. The ma¬

terial is not mature and the sporules are probably larger when

fully developed.

Septoria hieracicola Dearn. & House. On Hieracium lon-

gipilum. Spring Green. The spots are conspicuous but often

sterile, the small pycnidia inconspicuous.

Colletotrichum SOLITARIUM Ell. & Barth, on Solidago lati-

folia . Washington Island.

Cercospora briareus Ell. & Ev. On Acerates viridi flora.

Spring Green. In this collection the conidiophores are shorter

(mostly 20-35/*) and some of the conidia longer (100/* or more)

than in the type as described.

Puccxnia arenariae (Schum.) Wint. On Arenaria stricta .

Belmont. (N. C. Fassett.)

Hyalopsora cheilanthis (Pk.) Arth. On Cryptogramma

Stelleri. Viroqua. (N. C. Fassett.)

University of Wisconsin Herbarium,

April, 193 1.

Index to “Notes” XVIII and XIX.

(Names of parasites in italics.)

Acerates viridiflora 189

Aecidium ceanothi 258

Aecidium falcatae 258

Aecidium jamesianum Pk. 186

Aecidium lupini Pk. 258

Aecidium mariae-wilsoni Pk. 258

Aecidium onobrychidis Burr. 258

Aecidium plantaginis Burr. 186

Aecidium polygalinum Pk. 259

Aecidium pustulatum Curtis 258

Aecidium trientalis Tranz. 187

Aecidium xanthoxyli Pk. 258

Agoseris cuspidata 185

Alternaria herculea (E. & M.) Elli¬

ott 186

Althaea 185

Ambrosia psilostachya 188

Amorpha fruticosa 256

Amphicarpa monoica 183

Andropogon furcatus 253, 258

Andropogon Hallii 258

Anemone virginiana 259

Aquilegia canadensis 253

Arenaria stricta 189

Asclepias tuberosa 186

Ascochyta aquilegiae (Rabh.)

Hoehn. 253

Ascochyta imperfecta Pk. 260

Ascyrum stans 254

Asplenium acrostichoides 187

Aster (?) paniculatus 184, 254

Aster umbellatus 254

Baptisia tinctoria 258

Bidens vulgata 186

Boehmeria cylindrica 183

Brassica nigra 186

Bremia lactucae Regel 184

Caeoma (Aecidium) pentastemonis

Schw. 257

Calyptospora goeppertiana Kuehn

260

Carex Bebbii 259

Ceanothus amerieanus 258

Ceanothus ovatus 258

Cephalanthus occidentalis 256

Cercospora antipus E. & Hoi. 186

Cercospora briareus E. & E. 189

Cercospora cephalanthi E. & K. 257

Cercospora davisii E. & E. 186

Cercospora dulcamarae (Pk.) E. &

E. 261

Cercospora fingens Davis 186

Cercospora galii E. & Hoi. 186

Cercospora gentianicola E. & E. 256

Cercospora hyperici Tehon & Dan¬

iels 256

Cercospora junci n. sp. 259

Cercospora medicaginis E. & E. 186

Cercospora mississippiensis Tracy

& Earle 184

Cercospora molluginicola Lieneman

256

Cercospora molluginis Hals. 256

Cercospora molluginis Davis 256

Cercospora pas saloroides Wint. 256

Cercospora peter sii (B. & C.) Atk.

184

Cercospora smilacis Thuem. 184

Cercospora viticola (Ces.) Sace. 256

Cercospora zebrina 186

Cladosporium gloeosporioides Atk.

254, 255

Coleosporium solidaginis (Schw.)

Thuem. 188

Collet otrichum 255

Colie to trichum cladosporioides (E.

& E.) Atk. 255

Colie to trichum gloeosporioides Pen-

zig 255

Collet otrichum malvarum (A. Br. &

Casp.?) South worth 185

Colletotrichum solitarium Ell. &

Barth. 189

Gomandra umbellata 183

Cryptogramma Stelleri 189

Cylindrosporium passaloroides

(Wint.) Gilman & Archer 256

Cylindrosporium tradescantiae Ell.

& Kell. 254

Cynoglossum boreale 189

Cyperus esculentus 186

Cyperus Schweinitzii 187

Doassansia sagittariae (West.)

Fisch 183

Elymus striatus 185

Entyloma compositarum Farl. 186

Epilobium densum 260

Erigeron annuus 187

Erigeron canadensis 187

Erigeron ramosus 187

Eupatorium purpureum 184

Fragaria virginiana 260, 184

Galium concinnum 259

Galium triflorum 186, 261

Geranium maculatum 183

Geranium robertianum 184

Gloeosporium 254

Gloeosporium balsameum Davis 253

Gloeosporium cingulatum Atk. 185

Gloeosporium cladosporioides Ell. &

Hals. 254, 255

Davis— Index to Notes on Fungi

191

Glomerularia comi lonicerae Pk.

185

Gnaphalium decurrens 261

Halenia deflexa 256

Helianthus seaberrimus 185

Hendersonia foliicola (Berk.) Fckl.

188

Hieracium longipilum 189

Hyalopsora cheilanthis (Pk.) Arth.

189

Hypericum adpressum 256

Hypericum canadense 257

Hypericum majus 257

Hypericum mutilum 254

Hypericum prolificum 259

Hypericum virginicum 255

Iris lacustris 260

Juncus brachycephalus 259

Lactuca sativa (Cult.) 259

Lactuca villosa 184

Laportea canadensis 183

Larix laricina 257

Leptothyrium tumidulum Sacc. 253

Linaria canadensis 260

Lonicera dioica 186

Lonicera tatarica 185

Lophotocarpus calycinus 183

Lupinus perennis 258

Maianthemum canadense 259

Medicago 186

Medicago sativa 260

Melampsora bigelowii Thuem. 257

Melampsora medusae Thuem. 257

Melilotus alba 186, 259

Melilotus officinalis 186

Microsphaera alni (Wallr.) Wint.

259

Monarda didyma 188

Mycosphaerella personata Higgins

256

Oenothera biennis 188

Ovularia pulchella (Ces.) Sacc. 185

Pentstemon gracilis 261

Peridermium balsameum Pk. 257

Peridermium ingenuum Arth. 257

Peronospora ealotkeca DBy. 259

Peronospora grisea Unger 184

Peronospora linariae Fckl. 260

Peronospora melampyri (Bucholtz)

Davis 183

Peronospora polygoni Thuem. 185

Pestalozziella subsessilis Sacc. &

Ell. 183

Phalaris arundinacea 185

Phyllachora graminis (Pers.) Fckl.

185, 253

Phyllosticta aquilegiae Tehon &

Daniels 253

Phyllosticta aster icola E. & E. 254

Phyllosticta cruenta pallidior Pk.

260

Phyllosticta decidua Ell. & Kell

186, 260

Phyllosticta podophylli (Curt.)

Wint. 188

Phyllosticta punctata Ell. & Dearn.

260

Phyllosticta similispora Ell. & Da¬

vis 253

Phyllosticta sphaeropsispora E. &.

E. 254

Picea canadensis 257

Pinus Banksiana 188

Pinus resinosa 188

Pinus Strobus 188

Plantago aristata 186

Plasmopara halstedii (Farl.) Berl.

& De Toni 184

Plasmopara melampyri Bucholtz

183

Plasmopara pygmaea (Ung.)

Schroet. 259

Podophyllum peltatum 188

Polygala Senega 258

Polygonum Convolvulus 185

Populus nigra italica 260

Potentilla arguta 259

Pseudopeziza medicaginis (Lib.)

Sacc. 259

Psoralea Onobrychis 258

Pteris aquilina 260

Puccinia andropogonis Schwein.

257, 258

Puccinia arenariae (Schum.) Wint.

189

Puccinia bartholomaei Diet. 186

Puccinia bolleyana Sacc. 259

Puccinia canaliculata (Schw.)

Lagh. 186

Puccinia ceanothi Arth. 258

Puccinia cyperi Arth. 187

Puccinia ellisiana Thuem. 258

Puccinia hieracii (Schum.) Mont.

185

Puccinia investita Schw. 261

Puccinia Kaernbachii (P. Henn.)

Arth. 258

Puccinia karelica Tranz. 187

Puccinia menthae Pers. 188, 260

Puccinia patruelis Arth. 259

Puccinia pustulata Curt. 258

Puccinia violae (Schum.) DC. 187

Puccinia windsoriae 257

Puccinia xanthii Schw. 188

Pucciniastrum americanum (Farl.)

Arth. 257

Pucciniastrum arcticum (Lagh.)

Tranz. 257

Pucciniastrum galii (Lk.) Ed.

Fisch. 261

Pucciniastrum pustulatum (Pers.)

Diet. 260

192

Davis — Index to Notes on Fungi

Ramularia cephalanthi (E. & K.)

Heald 257

Ramularia virgaureae Thuem. 186

Ranunculus septentrionalis 188

Rhabdogloeopsis 253

Rubus allegheniensis 260

Rudbeckia hirta 185

Rudbeckia laciniata 185

Rudbeckia subtomentosa 185

Satureja vulgaris 260

Sclerotinia vaccinii Wor. 188

Sclerotium deciduum Davis 188, 260

Septoria astericola E. & E. 184, 254

Septoria atropurpurea Pk. 184, 254

Septoria cymbalariae Sacc. & Speg.

261

Septoria cynoglossi n. sp. 189

Septoria davisii Sacc. 189

Septoria fumosa Pk. 188

Septoria helianthi E. & K. 185

Septoria hieracicola Dearn. &

House 189

Septoria nolitangeris Gerard 254

Septoria pentstemonicola E. & E.

261

Septoria rudbeckiae E. & Hals. 185

Septoria tradescantiae (E. & K.)

n. comb. 254

Shepherdia canadensis 259

Smilax hispida 184

Solanum Dulcamara 261

Solidago confinis 254

Solidago latifolia 184, 189

Solidago patula 186

Solidago rigida 253

Solidago serotina 188

Sphaerotheca humuli (DC.) Burr.

259

Spinacia oleracea 185

Stagonospora atriplicis (West.)

Lind 185

Stagonospora spinaciae E. & E. 185

Stigmatea robertiani Fr. 184

Synchytrium aureum Schroet. 184

Synchytrium cellulare Davis 183

Synchytrium fulgens Schroet. 188

Synchytrium pulvereum Davis 183

Taphrina aurea (Pers.) Fr. 260

Thalictrum dioicum 186

Trientalis americana 187, 260

Trifolium 186

Uredinopsis 257

Uredinopsis copelandi Syd. 187

U romyces acuminatus magnatus

Arth. 259

Uromyces andropogonis Tracy 258

Uromyces hyperici-f rondo si

(Schw.) Arth. 257, 259

Uromyces pedatatus (Schw.) J. L.

Sheldon 258

Uromyces seditiosus Kern 186

Vaccinium canadense 260

Vaccinium macrocarpon 188

Vaccinium pennsylvanicum 260

Vermicularia libertiana Roum. 188

Veronica serpyllifolia 184

Viburnum Lentago 185

Viburnum Opulus 259, 260

Viola 258

Viola sagittata 187

Zanthoxylum americanum 258 •

IMPERMEABILITY IN MATURE AND IMMATURE SWEET

CLOVER SEEDS AS AFFECTED BY CONDITIONS

OF STORAGE.

Earl A. Helgeson

Introduction

Many seeds, in common with other plant structures, undergo

a dormant or so-called rest period. Methods of securing dor¬

mancy in seeds seem to be generally associated with: (1) em¬

bryo characteristics which prevent the immediate germination

of a seed even though conditions favorable for germination

obtain, and (2) seed coat characteristics which prevent either

the ready passage of liquids or gases, or the expansion of the

embryo. Crocker (1) has made a classification of the mechan¬

isms of dormancy. In so far as immature embryos are a con¬

sideration, he finds that at the time of seed ripening, these may

vary from an undifferentiated group of cells to a mature em¬

bryo. Individuals exhibiting this type of dormancy are present

in practically all the large groups of seed plants.

Impermeable or so-called “hard” seeds are defined by Har¬

rington (4) as “seeds whose coats are impermeable to water at

temperatures favorable for germination.” This type of dor¬

mancy seems to be of common occurrence in many species of

the Leguminosae with the result that their seeds may lie in the

soil for a number of years germinating a few at a time. A con¬

siderable amount of work has been done on fully mature imper¬

meable seeds but comparatively few studies are available on the

physiological processes which take place during the maturation

of such seeds. This study was undertaken to demonstrate at

what stage of maturation the impermeable condition was as¬

sumed and what the nature of the processes leading to the im¬

permeable condition might be.

Sweet clover, Melilotus, was chosen as experimental material

because of the constant high percentage of impermeable seeds

produced and because of the extreme resistance to the entrance

of water shown by these seeds.

194 Wisconsin Academy of Sciences , Arts, and Letters.

Experimental

Desiccation and the production of impermeability in sweet

clover seeds. Seeds used in these studies were collected when

they were slightly immature. Two degrees of immaturity were

recognized: (1) “Brown pod,” seeds with brown pods from

racemes having green peduncles, (2) “Yellow pod,” seeds with

slightly green or yellow pods and green peduncles. Brown pod

seeds had the bright yellow color of a fully matured seed, but

they had not dried down completely. Yellow pod seeds had at¬

tained their full size but had not started to dry down.

Racemes with seeds in the states of maturity mentioned

above were collected, taken to the labratory, and the pods care¬

fully stripped from the peduncles. Lots of 100 pods were count¬

ed out, an effort being made to select only healthy one-seeded

pods, and these were placed in Gooch crucibles. The crucibles

were then stored in a desiccator over calcium chloride or in a

vacuum desiccator with the same drying agent, according to

the nature of the drying desired. In the preliminary tests, the

lots to be treated in vacuo over calcium chloride were placed

in a desiccator with a side arm and the desiccator was evacuat¬

ed from time to time by means of a hand pump, until the pres¬

sure was equal to 61 centimeters of mercury. It was found,

however, that there was a slow leak in the desiccator, so this

method was abandoned and an automatically controlled Freas

vacuum oven at room temperature was used for further tests.

To obtain additional dryness in the oven, a pie pan containing

calcium chloride was placed under the tray supporting the

seeds.

Unless otherwise stated, germination tests were always made

in a Minnesota type germinator set at 20°C, moist blotters or

filter paper being used as a substrate. After counts of germi¬

nated seeds had been made, the blotters were allowed to dry,

the remaining seeds were then hulled by hand and the imper¬

meable seeds counted. The seeds which softened but did not

germinate were calculated by subtracting the sum of germinat¬

ed and impermeable seeds from the original number of pods

put to germinate. In the case of hulled seeds, all three frac¬

tions could be counted at the end of a test. Throughout this

paper the following abbreviations have been used to denote the

three fractions : Gm, germinated seeds ; HS, impermeable

Helgeson — Impermeability in Sweet Clover Seeds . 195

Table I. Influence of desiccation over extended 'periods upon the production oj

impermeability in yellow pod white sweet clover seeds.

*Total after 7 and 10 days respectively.

fPut to germinate without hulling, all other lots were hulled before being put

to germinate.

seeds ; and SS, soft seeds or seeds which took up water but did

not germinate.

The behavior of immature seeds in a vacuum desiccator and

in an ordinary desiccator is shown in Table I. The control and

196 Wisconsin Academy of Sciences, Arts , and Letters .

the first 2 day lot were unhulled. All the other lots were hand

hulled and 100 seeds counted out for the germination test.

From these data it is clear that even a 2 day period of desic¬

cation causes a considerable increase in impermeability and

that a treatment in a vacuum over calcium chloride causes a

somewhat greater increase in impermeability. The maximum

“hardening” effect seems to have been reached after 3 days in

storage in a vacuum over calcium chloride and after 4 days in

storage over calcium chloride without vacuum.

To test the effects of a higher degree of desiccation on the,

permeability of immature seeds, the Freas vacuum oven kept

at constant partial vacuum of 37.5 ±0.2 cm. Hg. and contain¬

ing calcium chloride as stated above was used. The results

obtained are given in Table II.

Table II. Influence of desiccation in constant partial vacuum over calcium chlo¬

ride at room temperature upon the production of impermeability in immature white

sweet clover seeds. The seeds used in this series of tests were unhulled.

*Put to germinate without hulling. Lots not marked were hulled before being

put to germinate.

The data in the above table are in line with those in Table

I. In general, there seems to be a more rapid increase in im¬

permeability in brown pod lots than there is in yellow pod lots.

The decrease in seeds which germinate and in soft seeds is

also more rapid. From the third day on, the lots used in Table

II were hand-hulled before being put to germinate, this served

Helgeson — Impermeability in Sweet Clover Seeds . 197

as a rather good check on the unhulled lots used in the preced¬

ing tests and seemed to show that results secured with unhulled

lots are fairly reliable.

The effect of storage over extended periods upon mature and

slightly immature sweet clover seeds . A preliminary germina¬

tion test, on a lot of unhulled brown pod white sweet clover

seeds, brought out the fact that seeds harvested at the right

stage could be held in storage under moist, cool conditions for

some time without hardening (Table III).

Accordingly, a considerable number of collections from indi¬

vidual plants in the brown pod and yellow pod stages were

made. These lots were placed in manila envelopes and stored

in three places as follows: (a) Laboratory, (b) cold room, and

Table III. Germination tests of brown pod white sweet clover seeds , held after

harvesting under three different conditions of storage.

*That is, the percentage germination at the beginning of the experiment.

fOut-of-doors 515 days; then in the cold room 75 days.

(c) out-of-doors. The cold room had a fairly constant tem¬

perature of 7° C. and had a relative humidity of approximately

85 per cent; the seeds stored out of doors were placed in a

special louvered shelter. They were thus protected from rain

but subjected to seasonal changes in temperature and humidity.

In practically all germination tests 100 unhulled pods were

used and the percentages of germinated seeds, impermeable

seeds, and soft seeds determined as for the desiccation experi¬

ments. The counts were taken after seven days in the germi-

nator. As often as possible the first or original germination

test was made as soon as the lots were collected. Whenever

that could not be done, the lots were held in a moist chamber

at room temperature until the test could be made, which was

usually the next day. Immediately after the test sample had

been taken the lots were placed in storage.

198 Wisconsin Academy of Sciences , Arts , and Letters .

A few lots of ripe seeds were also collected, from some of the

same individual plants from which immature lots were taken,

and placed under similar storage conditions. These would

serve as controls on any softening influence due to the various

storage conditions. Some mixed lots of immature seeds were

made up by using seeds from 10 or more plants of the same

variety. Ripe, mixed lots were collected from the same plants

from which immature lots were taken. As no significant dif¬

ference between seeds from wild plants as against cultivated

plants of the same variety were noted no mention is made of

the seed source. Most of these samples were collected during

August 1928 at Madison, Wisconsin. The data for the first

tests, after a period of 7 to 8 months, are presented in Table IV.

Table IV. Germination of brown pod yellow and white sweet clover seeds , after

storage for 7 to 8 months in the laboratory and cold room respectively. In each case

the seeds of any particular lot were from a single plant.

*50 seeds used for each test.

Germination test at beginning of experiment.

As only 50 seeds were used in the tests recorded in Table IV,

the findings can be regarded only as relative. It is seen that

even after 7 to 8 months in storage the cold room lots gave

rather high percentages of germination. The less mature lots,

i. e., those giving around 80 per cent original germination,

showed a greater reduction in seedling production than did

those lots germinating around 40 per cent. This reduction was

largely due to an increase in soft seeds, brought about prob¬

ably by the action of micro-organisms during the germination

test. The lots stored in the laboratory had practically all be¬

come impermeable.

After a period of from 15 to 17 months in storage, the ger¬

mination of lots kept in the cold room was still relatively high,

in fact, practically the same as it was after 8 months. The

Helgeson — Impermeability in Sweet Clover Seeds. 199

data in Table V show further that the percentages of soft seeds

in the cold room lots are very high and seem to have increased

at the expense of the hard seeds. The lots stored in the labora-

Table V. Germination of brown pod white and yellow sweet clover seeds after

storage from 15 to 17 months under the several conditions indicated below. Y = yellow ;

W = white) I —from an individual plant ; M — mixed.

*Based on 105 seeds.

fHand-hulled before testing.

jlO days on germinator before counting.

Table VI. Germination of mature and brown pod white and yellow sweet clover

seeds after storage under the different conditions indicated below. Y — yellow ; R —ripe;

W —white; M —mixed; I =from an individual plant.

*Ripe seeds from same plants as immature lots of corresponding number.

tWhite mixed yellow pod seed.

JBased on 101 seeds.

§Based on 115 seeds.

1 1 Based on 104 seeds.

If Based on 102 seeds.

200 Wisconsin Academy of Sciences , Arts , and Letters .

tory still show a high percentage of impermeable seeds and

relatively low percentages of permeable and soft seeds. There

seems to be no apparent softening of seeds owing to storage

in the dry condition in the laboratory.

Although the laboratory lots served as a good check on the

effects of drying in storage, they did not give any information

as to the possible softening effect of humidity and low tempera¬

ture on the lots held in the cold room. To test what effect these

latter agencies might have, a number of the mature lots from

the same plants from which immature seeds were collected

were put to test. These data, together with tests of immature

lots from the same plant, are recorded in Table VI.

That there may have been some slight effect on the mature

seeds stored in the cold is shown in the preceding table, but the

differences are hardly large enough to account for all the per¬

meable seeds occurring in the brown pod lots. There was also

a small increase in soft seeds in ripe lots stored in the cold

room as compared with the corresponding lot stored in the la¬

boratory.

The data set down in Table VII show the effect of storing

mature and immature seeds out-of-doors where they are sub¬

ject to seasonal variations in temperature and humidity and

then giving them constant storage in the cold room for a short

period. Checking these lots against lots stored continually in

the cold room, it is clear that storage out of doors reduces the

Table VII. Germination of mature and brown pod white sweet clover seeds stored

out-of-doors and then in the cold room.

R = ripe ; M = mixed.

*Ripe seed from same plants as immature lots of corresponding number.

fBased on 104 seeds.

JBased on 102 seeds.

§Mixed yellow pod.

Helgeson — Impermeability in Sweet Clover Seeds. 201

number of impermeable seeds in the mature lots to a consid¬

erable extent. The percentages of soft seeds in both types of

seed are reduced to a minimum.

It is to be regretted that more material of this type was not

available as there seems to be an indication that storage, im¬

mediately after harvest, in a place where the seeds are pro¬

tected from direct exposure to weather but subject to natural

temperature and humidity changes, reduces the ratio of im¬

permeable seeds without effecting the total viability of the

seeds.

The preceding storage tests have shown that cold, moist con¬

ditions prevent slightly immature seeds from becoming im¬

permeable. The question next arose as to whether this inhibi¬

tion was permanent or lasted only so long as seeds were kept

in the cold room. To test this a number of 100-pod lots were

counted out and placed in a desiccator over calcium chloride

and left for 7 days. The desiccator was exhausted by means

of a vacuum pump until the gauge registered 60 cm. of mer¬

cury and it was held at approximately this level for the first 24

hours. Then the pump was disconnected and the desiccator

was allowed to come to atmospheric pressure and was so main¬

tained for the remainder of the 7 days. At the end of the dry¬

ing period the lots were taken out and put to germinate. The

data for this experiment are presented in Table VIII.

Unfortunately there was not enough material left after se¬

lecting the lots for drying to run a germination test at the time

the drying test was started. The counts on germination before

drying were therefore made from the last previous test. The

results brought forward in Table VIII indicate that the imma¬

ture lots stored in the cold room are only temporarily kept from

hardening. A short drying period suffices to reduce the num¬

ber of permeable seeds to an extent almost equal to the reduc¬

tion effected in the original lots by storage in the laboratory.

Another interesting fact brought out by these data is the great

reduction in the percentage of soft seeds. Here again the

dried lots are comparable with the original lots which were

stored in the laboratory.

The high percentage of impermeable seeds present in the

dried lots suggested that perhaps many of the soft seeds had

also become impermeable. To arrive at the true value of the

202 Wisconsin Academy of Sciences , Arts, and Letters .

■52

CO

§2

e

S- ' ~

^ 00

o <SS

•sf

% e

la

CD r>J

«o cs

If

“1

*c*a

§ II

co HH

'Sfe

a s

g S

<=> I3

>*2,

w

*Based on 105 seeds.

Helges on— Impermeability in Sweet Clover Seeds . 203

impermeable seeds remaining after the drying test, these were

rubbed between sandpaper and germinated. That a consider¬

able number of soft seeds did become impermeable during the

drying process is apparent from results obtained in this germi¬

nation test. The percentages of germination in some of the

scarified dried lots are, however, somewhat higher than the

percentages obtained in the cold-room lots before drying. This

seems to confirm the statement made before, that the cold room

lots showed rather high percentages of soft seeds because of

the attack of micro-organisms during the germination test, and

that drying reduced the susceptibility of the seeds to such at¬

tacks (Table VIII).

To test the effects of various conditions of storage on mature

lots of commercial white sweet clover seeds the following ex¬

periment was set up : Three cloth bags containing respectively,

(1) scarified seeds, (2) seeds which had been hulled but not

scarified, and (3) unhulled Grundy County seeds, were placed

out of doors in a small house built in such manner that rain

was kept out but no protection against other weather changes

was afforded. Similar lots of the same seeds were kept in the

cold room and in the laboratory. Small pans containing the

three kinds of seeds mentioned above were also stored in desic¬

cators in the cold room, and in the laboratory. The commer¬

cial seeds used were obtained from the Agronomy department

and were of the 1927 crop. The unhulled Grundy County seeds

were purchased from the L. L. Olds Seed Co. and were grown

and harvested in North Dakota in 1927. The out-door lots

were placed in storage October 29, 1927, and the other lots

were stored February 23, 1928. On this latter date all lots

were tested for germination and this test is given as the con¬

trol. The final germination test of all lots was made on March

14, 1930, or after a period of 24 months and 19 days.

Two hundred seeds were used in each of the tests in this

experiment and the Grundy County seeds were hand-hulled be¬

fore testing. The data obtained from these tests (Table IX)

show that dry storage is the best means of conserving the

vitality of commercial seeds. Storage in the cold room with

the prevailing high humidity was especially detrimental to the

scarified and commercial-hulled seeds. In the desiccators the

seeds stood up about as well as did those stored in the dry

204 Wisconsin Academy of Sciences, Arts, and Letters .

laboratory. The lots stored out of doors were intermediate be¬

tween the cold room lots and laboratory lots. Here again, as

in tests on immature lots, the effects of seasonal changes in

reducing the number of impermeable seeds is apparent. About

Table IX. Effect of storage under various conditions on mature sweet clover seeds :

All counts were made after 5 days in the germinator. D —in desiccator.

the same percentage of impermeable seeds in the hulled com¬

mercial and the unhulled Grundy County samples softened.

General Discussion

The data presented have brought out a number of facts which

seem to throw some light on the nature of the processes leading

to the impermeable state in sweet clover seeds.

The desiccation studies, as well as the studies on immature

and mature seeds held under different conditions of storage, in¬

dicate that the change from the permeable to the impermeable

state takes place as a final step in the maturation of the seed.

The experiments have shown that the assumption of the im¬

permeable state can be prevented by storage in a cool, moist

place, or brought about by storage in a dry place. Esdorm

(1), working with yellow lupine, found the same to hold true

for these seeds. That immature seeds of Gymnocladus and of

vetch have very few impermeable seeds and that the imperme¬

able seed percentage is increased by dry storage has been

shown by Raleigh (7) and by Jones (5). The former worker

found rather high percentages of pectic materials to be pres¬

ent in the seeds of Gymnocladus dioica and concluded that per¬

haps drying caused these to change to some resistant anhydride

Helgeson— Impermeability in Sweet Clover Seeds . 205

form. Jones states that with vetch, Vicia villosa, the imperme¬

able condition is brought about by some dehydration process

which operates independently of the plant. He finds that, if

the drying is continued, these seeds finally become permeable

again and that the presence of moisture increases the rate

at which such seeds become permeable.

In the case of sweet clover, some such dehydration process

probably takes place, but it does not seem likely that pectic

materials alone are concerned although they seem to be pres¬

ent to some extent. With fully mature impermeable seeds

neither continued drying nor high humidity seem to have any

appreciable effect on permeability. When, however, immature

seeds are rendered impermeable by drying they seem to be

rather sensitive to changes in temperature and humidity. That

immature impermeable seeds of certain of the Leguminosae be¬

come permeable in storage more rapidly than do mature im¬

permeable seeds was shown by Harrington (U). That the im¬

mature seeds stored in the cold room were only temporarily

kept from hardening and that such seeds become impermeable

after a short period of desiccation is a matter of interest.

These facts seem to indicate that dehydration, possibly asso¬

ciated with an oxidative process, brings about an irreversible

change in some material which is colloidal in nature.

The effects of various conditions of storage on mature com¬

mercial seeds are of considerable interest. It appears that stor¬

age in a dry situation is the best means of conserving the vitali¬

ty of all such seeds whether scarified or not. The rapid loss

of viability in scarified sweet clover seeds stored in a dry place

is in line with the results obtained by Graber (3) for alfalfa,

and by others (6) for sweet clover. That high humidity is

especially detrimental to the viability of all the sweet clover

seeds is also brought out. It is interesting to note that very

few impermeable seeds become permeable in the cold room;

while the natural variations in humidity and temperature to

which all lots stored out-of-doors were subjected to, caused a

considerable reduction of impermeable seeds.

It is with sincere gratitude that the writer acknowledges the

valuable suggestions and kindly criticisms given from time to

time by Dr. B. M. Duggar and Dr. L. F. Graber.

206 Wisconsin Academy of Sciences , Arts, and Letters .

Summary and Conclusions

(1) Impermeability in sweet clover seeds is probably brought

about by dehydration in the late stages of maturation.

(2) Slightly immature sweet clover seeds are practically all

permeable and produce high percentages of germination.

(3) The impermeable state in such immature seeds can be

induced by storage in a dry place. On the other hand, hard¬

ness may be prevented for at least 19 months by storage in a

moist, cold room.

(4) Immature permeable seeds taken from the cold room

and placed over calcium chloride for 7 days become imperme¬

able.

(5) Storage for 16 months in a moist, cold room causes a

notable reduction in the viability of permeable seeds. Storage

under dry conditions for the same period of time had little

effect on either the impermeability of hand-picked seeds or on

their viability.

(6) The viability of scarified seeds was greatly reduced un¬

der all conditions of storage. Storage out-of-doors reduced the

percentage of impermeable seeds in all mature and immature

lots.

Literature Cited

1. Crocker, Wm. 1916. Mechanics of dormancy in seeds. Amer. Jour.

Bot. 3 : 99-120.

2. Esdorm, J. 1928. Beitrage zur Keimungsphysiologie hartsch&liger

Samen. Angewandte Bot. 10 : 469.

3. Graber, L. F. 1922. Scarification as it effects longevity of alfalfa

seeds. Jour. Amer. Soc. Agron. 14 : 298-302.

4. Harrington, G. T. 1916. Agricultural value of impermeable seeds.

Jour. Agr. Res. 6 : 761-796.

5. Jones, J. P. 1928. A physiological study of dormancy in vetch seed.

N. Y. (Cornell) Agr. Exp. Sta. Mem. 120 : 1-50.

6. Bull. Univ. Minn. Rpt. N. W. Exp. Sta. ( Crookston) 20 : 28-31. 1926.

7. Raleigh, G. J. 1930. Chemical conditions in maturation, dormancy,

and germination of seeds of Gymnocladus dioica. Bot. Gaz. 89 :

273-294.

PRELIMINARY REPORTS ON THE FLORA OF

WISCONSIN. XV.

POLYGONACEAE

Kenneth L. Mahony

The station records used in this study were taken from the

following herbaria: Northland College, the Milwaukee Public

Museum, S. C. Wadmond, and the University of Wisconsin.

Rumex

A. None of the leaves halberd-shaped or arrow-shaped; flowers perfect

B. Valves entire or denticulate, 3-27 mm. broad

C. Grains of fruiting calyx none, or single and minute, not one-

third as long as the valves: valves 5-7 mm. broad

D. One valve bearing a small grain or its midrib merely thick¬

ened at the base . R. Patientia

D. Valve with its grain conspicuous, 2-3 mm. long .

. R. Patientia var. kurdicus

C. Grains 1-3, well developed, mostly one-half to three-fourths as

long as the valves; valves 1.5-6 mm. broad

E. Pedicels filiform, curved or flexuous

F. Leaves crisped on margin . R. crispus

F. Leaves flat

G. Pedicels with tumid joints, rarely exceeding the cori¬

aceous, greenish, straw-colored, or dull brown calyx

H. Valves 1.5 -3. 5 mm. broad, 2. 5-5.5 mm. long; pedi¬

cels semi-nodding; grains on midrib usually three;

stem usually much branched and only semi-

ascending . R. mexicanus

H. Valves 3-6 mm. broad, 3.5-8 mm. long; pedicels

definitely nodding; grains on midrib usually one;

stem not much branched and strictly ascending. . . .

. . . . . . R. altissimus

G. Pedicels obscurely jointed, mostly exceeding the mem¬

branous finally purplish calyx

I. Grains three . R. Britannica

I. Grains solitary . [ R . occidentalism 1

1 Species in brackets are not represented in our collections from Wisconsin,

but should be looked for, as they probably occur in the state.

208 Wisconsin Academy of Sciences, Arts, and Letters.

E. Pedicels clavate, deflexed, straight ish and slightly rigid, 2-3

times as long as the subacuminate valves . R. verticillatus

B. Valves with long sharp salient teeth at least near the base

J. Perennial; valves ovate-halberd-shaped; the whorls of flowers

loose and distant on the leafless spike . R. obtusifolius

J. Annual ; valves rhombic-oblong, lance-pointed ; the whorls of

flowers excessively crowded in leafy and compact spikes .

. . . . R. maritimus var. fueginus

A. Some or all of the leaves halberd-shaped ; flowers dioecious

K. Valves much exceeding the fruit . . . . [i?. hastatulus]

K. Fruit exserted from the minute scarcely changed calyx . . . .

. . . R. Acetosella

R. Patientia L. (Fig. 1) seems to be almost confined to the

western half of the state, but is found as far north as Bay-

field County and as far south as Iowa, Dane, and Racine Coun¬

ties. This plant is one that has been introduced into this coun¬

try from Europe and should have a wide distribution in the

state.

R. Patientia L. var. kurdicus Boiss. (Fig. 2) is also an in¬

troduced plant, but is represented in this study by only two

specimens, one collected in each Trempealeau and Pepin Coun¬

ties.

R. CRISPUS L. (including R. elongatus Guss.) (Fig. 8) is

found over the entire state. Such a distribution is to be expect¬

ed from an introduced plant.

R. mexicanus Meisn. (Fig. 4) is general over the state from

north to south, but is found only in the western half of the

state except for three stations, one in each, Milwaukee, Brown,

and Ozaukee Counties.

The specimen from Ozaukee County was collected by the

writer and was found growing along the harbor where Lake

Michigan boats unload at Port Washington, and might possibly

have been brought there in ballast. It is very evident from the

distribution that this plant tends to avoid the Archean Rock

region in Wisconsin.

R. altissimus Wood (Fig. 5). This species is general over

the western and southern parts of the state.

R. Britannica L. (Fig. 6) seems to be quite general over the

glaciated portion of the state. It is often called the “great

water dock”, and as the name indicates it likes water or marshy

Mahony — Reports on Flora of Wisconsin. XV.

209

Rumex Patientia

Rumex crispus

Rumex Patientia

var. kurdicus

210 Wisconsin Academy of Sciences, Arts, and Letters.

places. It is usually found in swamps and along margins of

lakes and streams which are numerous in glaciated regions.

R. verticillatus L. (Fig. 7) is found north as far as Oneida

County, and is especially abundant along the Mississippi river

as far north as Pierce County.

R. obtusifolius L. (Fig. 8) is found at scattered stations

ranging from the Apostle Islands south to Racine County, but

seems to be absent in the northeast portion of the state. It

has been naturalized from Europe and such a scattered distri¬

bution should be expected.

R. maritimus L. var. fueginus (Phil.) Dusen.; Rhodora vol.

17 : 73-83, 1915. R. persicarioides of Gray’s Man. 7th ed. 357,

1908. Britton and Brown, Ill. FI. 2nd ed. : 659, 1913; and re¬

cent American authors, not L. (Fig. 9). This is a plant that

was found to be in the southeastern and northwestern portions

of the state as far north as Superior.

Fernald writes of certain species of plants centering on the

gulf of St. Lawrence, and then reappearing in the western

United States.2 The general range of R. maritimus var. fuegi¬

nus in North America would suggest that it is such a species.

It appears to be a preglacial plant that only persisted in the

unglaciated areas and is now working back into its old range

from the west and is coming into Wisconsin from the south¬

west, or it may have persisted in these scattered areas in Wis¬

consin and is now gradually working its way westward.

At practically every station in Wisconsin this plant has been

collected in a region of newly formed soil, for example : along

the Mississippi river the plant is found growing between wing

dams on the river; in Dodge County the station is at Horicon

Marsh, which is a drained lake. One station in central Dane

County is an abandoned beach of Lake Mendota near Pheasant

Branch ; the other, in the southeastern part of the county is at

Rice Lake, an intermittent lake ; also the station in Polk County

is on an intermittent lake.

R. Acetosella L. (Fig. 10) is very general and probably can

be found in every county of the state. This is another Rumex

that has been introduced from Europe and such a distribution

is expected.

2 Mem. Am. Acad, of Arts and Sci., vol. 15, no. 3. 1925.

Mahony —Reports on Flora of Wisconsin . XV.

211

+ Polygonum aviculare

var. crass if olium

212 Wisconsin Academy of Sciences, Arts, and Letters,

Polygonum

A. Leaf -blades ovate to lanceolate, tapered at base to the petiole

B. Flowers in axillary fascicles, or spicate with foliaceous bracts .....

. . . . . . . . . Subgenus Avicularia

C. Achenes conspicuously exserted . . P. exsertum

C. Achenes nearly or quite included in fruiting calyx

D. Branches terete or nearly so

E. Pedicels included; sepals usually white or bluish-green,

1.3-1.8 mm. long

F. Stems prostrate or nearly so

G. Leaves thick and broadly spatulate .

. . P. aviculare var. crassifolium

G. Leaves neither thick nor broadly spatulate

H. Leaves lanceolate, 6-20 mm. long. .P. aviculare

H. Leaves thin and linear, 5-9 times as long as

broad . P. aviculare var. angustissimum

F. Stems erect or nearly so ... P. aviculare var. vegetum

E. Pedicels exserted ; sepals yellowish-green, bluish-green,

greenish, yellow, pink or roseate, often 2-3 mm. long

I. Leaves elliptical

J. Leaves yellowish-green, obtuse . . . P. erectum

J. Leaves bluish-green, very rounded at the tips ......

. . . . . P. achoreum

I. Leaves lanceolate

K. Flowers greenish, yellow, or pink . . .

. . P. ramosissimum

K, Flowers roseate _ P. ramosissimum f. atlanticum

D. Branches rather sharply angled

L. Leaves strongly pilcate ; flowers erect _ ...... P. tenue

L. Leaves flat, with revolute margins ; flowers nodding .....

. . . . . . . . [P. DouglasU]

B. Flowers not in axillary fascicles, but spicate without foliaceous

bracts

M. Flowers in dense spikes, with small scarious bracts ; styles not

deflexed in fruit and not hooked . . Subgenus Persicaria

N. Sheaths nearly or quite free from ciliation

O. Annual ; achene compressed ; plants with a branched

fibrous root system

P. Peduncle not glandular ; achenes 1-2 mm. broad

Q. Under surface of leaves glabrous . . .

. . . . . P. lapathifolium

Q. Under surface of leaves tomentose, at least the

lower ones ..... P. lapathifolium var. salicifolium

Mahony — Reports on Flora of Wisconsin. XV.

213

P. Peduncle glandular; achenes 2-3 mm. broad

R. Under surface of leaves tomentose, at least the

lower leaves . P. scab rum

R. Under surface of leaves not tomentose

S. Glands of hairs on peduncle with red pigment . .

. P. pennsylvanicum var. laevigatum

S. Glands of hairs on peduncle without red pigment

P. 'pennsylvanicum var. laevigatum f. pallescens

0. Perennial; achene compressed or turgid; plants with

floating or creeping, subligneous rhizomes

T. Peduncle glabrous; panicle ovoid, 1-5 cm. long

U. Aquatic form; stems floating or somewhat emersed;

leaves glabrous; sheaths without herbaceous mar¬

gin . P. natans f. genuinum

U. Terrestrial form; stems upright and leafy; leaves

more or less hairy; sheaths with herbaceous mar¬

gin . . . P. natans f. Hartwrightii

T. Peduncles pubescent; panicle dense, cylindric, 3-10 cm.

long

V. Aquatic form; stem floating or somewhat emersed;

leaves glabrous . [P. coccineum f. natans ]

V. Terrestrial form; stems upright and leafy; leaves

more or less hairy . P. coccineum f. terrestre

N. Sheaths bristly-ciliate

W. Stems and peduncles glandular-hispid . P. Careyi

W. Stems and peduncles not glandular-hispid

X. Sepals dotted with dark glands

Y. Achenes dull and pitted

Z. Pedicels not strongly exserted from the ocreolae ;

achenes mostly 3-3.5 mm. long. .P. Hydropiper

Z. Pedicels strongly exserted from the ocreolae;

achenes mostly 2-2.5 mm. long .

. P. Hydropiper var. projectum

Y. Achene shining and smooth

a. Stems 0.6-1. 6 m. high; leaves lanceolate, at¬

tenuate, 7-12 cm. long, taper-pointed .

. P. punctatum

a. Stems 3-6 dm. high; leaves lanceolate, smaller,

thinner, and lighter green than in the type ....

. P. punctatum var. leptostachyum

X. Sepals not dark-dotted

b. Stems hairy . . .

b. Stems not hairy

P. orientate

214 Wisconsin Academy of Sciences , Arts , and Letters .

c. Flowers in erect, or short-cylindrical, densely

flowered spikes 0.5-2.75 cm. long . . P. Persicaria

c. Flowers in erect slender loosely flowered often

interrupted spikes 3-6 cm. long . . .

. P. hydropiperoides

M. Flowers in loose naked long and slender spikes with no small

scarious bracts; styles deflexed in fruit and hooked .

. Subgenus Tovar a . P. virginianum

A. Leaf-blades sagittate to heart shaped, or round-ovate, truncate or cor¬

date at the base

d. Stems armed with reflexed prickles on angles .

. Subgenus Echinocaulon

e. Leaves halberd-shaped . P. arifolium

e. Leaves arrow-shaped . P. sagittatum

d. Stems not armed with reflexed prickles

f. Stems twining and slender . Subgenus Tiniaria

g. Achenes minutely roughened, dull, black . P. Convolvulus

g. Achenes smooth, shining, black

h. Sheaths fringed at base with reflexed bristles. . . .P. cilinode

h. Sheaths not fringed at the base with reflexed bristles .

. . . P. scandens

f. Stems erect and stout. . .Subgenus Pleuropterus . . .P. cuspidatum

The subgenus Avicularia Meisn. is represented in Wisconsin

by six species. This subgenus contains plants that are classed

as European, Asiatic, road side, and prairie plants. They are

a very aggressive group of plants and present many taxonomic

problems.

P. exsertum Small is represented in this study by only one

collection. The plant was collected by J. H. Schuette, in 1881,

and the station is recorded as the Fox River valley.

P. aviculare L. (Fig. 11). This plant seems to have a scat¬

tered distribution around the margins of the state. Very few

collections have been made from the central part of the state.

P. AVICULARE L. var. angustissimum Meisn. (Fig. 12) also

seems to be confined to the margins of the state.

P. AVICULARE L. var. crassifolium Lange ; House, N. Y. State

Mus. Bull. 254 : 291, 1924. P. aviculare var. littorale of

Gray’s Manual, 7th ed. (Fig. 12). Represented in Wisconsin

by only two collections.

P. AVICULARE L. var. VEGETUM Ledeb. (Fig. 13). This plant

is scattered widely over the state.

215

Mahdny— Reports on Flora of Wisconsin. XV.

Polygonum avieulare

var. vegetum

Polygonum erectum

Polygonum tenue

Polygonum pennsylvanicum

var. laevlgatum

216 Wisconsin Academy of Sciences, Arts, and Letters.

P. ACHOREUM Blake, Rhodora 19 : 232, 1917, (Fig. 14) is

of a widespread distribution in Wisconsin.

P. erectum L. (Fig. 15). This plant has been collected in the

north, central, eastern and southern parts of the state, but no

collections have been made from the western part.

P. ramosissimum Michx. (Fig. 16). This plant is also of a

general widespread distribution over the state. It has been

collected in Minnesota opposite Alma, Wisconsin ; and should be

found in this state on the sands of the Mississippi river.

P. RAMOSISSIMUM Michx. f. ATLANTICUM Robinson, Rhodora

4 : 72, 1902 (Fig. 16). We have only one collection of this in

the state, from the south end of Lake Winnebago.

P. tenue Michx. (Fig. 17) is restricted to the southern half

of the state. It has been collected as far north as Polk County

but the major part of the collections have been made along the

Wisconsin river south of Marathon County.

Subgenus Persicaria (Tourn.) L. has by far a larger repre¬

sentation in Wisconsin than any of the other subgenera. There

are eleven species of this subgenus found in Wisconsin and

numerous varieties and forms.

This subgenus like the Avicularia group contains European,

Asiatic, road side and prairie plants. They are weedy aggres¬

sive plants and present many taxonomic problems.

P. PENNSYLVANICUM L. var. laevigatum Fernald, Rhodora

19 : 73, 1917. (Fig. 18). Of general distribution over the

state, being found largely on the rivers and lakes. It appears

to be less common in the Mississippi river bottoms than the

form.

P. PENNSYLVANICUM L. var. laevigatum Fernald f. PALLE-

SCENS Stanford, Rhodora 27 : 180, 1925. (Fig. 19). This

plant is confined to the western half of the state except for one

collection made near Green Bay in Brown County.

P. SCABRUM Moench. ( P . tomentosum Schrank) (Fig. 20).

For a complete synomymy see Rhodora 23 : 259, 1921. We

have only a few collections of this plant from the state, and

with one exception all have been collected from the northwest¬

ern part of the state.

Mahony— Reports on Flora of Wisconsin . XV .

217

Polygonum pennsyl van! e um

var. laeYigatum

f • pallescens

Polygonum scabrum

s:

30

Polygonum lapathifolium

yar# salicifolium

Polygonum nataas Polygonum natans

f. Hartwrightii f* genuinum

218 Wisconsin Academy of Sciences, Arts, and Letters .

P. lapathifolium L. (Fig. 21). This plant is of a general

distribution over the state.

P. LAPATHIFOLIUM L. var. SALICIFOLIUM Sibth. ; Wiegand and

Eames, Cornell Univ. Agr. Exp. Stat. Memoir 92 : 190, 1926

(Fig. 22). This plant is largely confined to the northwestern

corner of the state. However, we have one specimen that was

collected near Verona in Dane County and another that was col¬

lected by the writer on a dam built along the Rubicon river

at Hartford, in Washington County.

P. NATANS A. Eaton f. Hartwrightii (Gray) Stanford, Rho-

dora 27 : 160, 1925. (P. amphibium Small var. Hartwrightii

(Gray) Bissell of Gray’s Manual, 7th ed.) (Fig. 28). Most of

the collections of this plant have been made in the southeastern

part of the state, however, we do have two stations in the north¬

eastern part of the state.

P. NATANS A. Eaton f. genuinum Stanford; for complete

synonymy see Rhodora 27 : 159, 1925. (The floating form

of P. amphibium Small of Gray’s Manual, 7th ed.) (Fig. 24).

This plant has been found in the northwestern and southeast¬

ern part of the state.

P. COCCINEUM Muhl., f. TERRESTRE (Willd.) Stanford; for

complete synonymy see Rhodora 27 : 163, 1925. (P. Muhlen-

bergii (Meisn.) Wats, of Gray’s Manual, 7th ed.) (Fig. 25).

This plant is found over the northwestern and southeastern

part of the state.

During the late spring of 1930 the writer gathered from be¬

side a railroad track at Madison, Wisconsin a few living speci¬

mens of Polygonum natans f. Hartwrightii and transferred

them to the University green house. They were planted in a

tank and the water level kept at a mark which allowed the plants

to be completely submerged. At first the leaves came off and

roots developed at the nodes of the upright stems, but soon

the whole aerial part of the plants turned brown and appar¬

ently died. About the first of May 1931 new green branches

developed from these old branches of the year before. This

time the leaves borne had all of the characteristics of the Poly¬

gonum natans f. genuinum . These results are similar to those

obtained by other men working with the same plants. How¬

ever, during the summer months the plants grew so much that

Mahony ■ — Reports on Flora of Wisconsin. XV. 219

Polygonum coccineum

var. terrestre

Polygonum Hydropiper

var. projectum

Polygonum punctatum

var, l.eptostacfcyum

220 Wisconsin Academy of Sciences , Arts , and Letters .

it was impossible to keep the plants entirely submerged. As a

result they grew several inches above the surface of the water.

The leaves produced above the water were not like those of

the submerged form, but were typical of the P. natans f. Hart-

wrightii. By September the submerged and floating leaves had

all died and the plant gave the appearance of being the land

form.

P. natans f. genuinum, P. natans f. Hartwrightii and P. coc-

cineum f. terrestre seem to avoid the region of the Archean

Rock. The rocks here are largely granite, and are covered

over to various depths by glacial deposits. However, this gran¬

ite rock formation tends to make most of the lakes of an acid

nature. These two species seem to frequent the lake regions

but apparently avoid these acid lakes.

We have two collections of P. natans f. genuinum from north¬

ern Wisconsin, one from each, Vilas and Oneida Counties.

Their presence here may be due to the fact that these two lakes

through some factor have had their acid nature neutralized and

have become favorable for the growth of this species.3

There are other species of the Polygonaceae, for instance

Rumex mexicanus, Polygonum punctatum var. leptostachyum,

P. Hydropiper var. projectum and P. Persicaria that tend to

stay out of this Archean Rock region. One can hardly say that

it is this granite rock below these glacial deposits that is the

factor in the distribution of these plants, although it may play

quite an influential part.

P. Careyi Olney (Fig. 26). We have only three collections

of this in the state.

P. Hydropiper L. (Fig. 27). We have only one collection of

this taken from Racine County. According to Stanford this is

a plant that has been introduced from Europe.

P. Hydropiper L. var. projectum Stanford, Rhodora 29 :

86, 1927. (Fig. 28). This plant is of a scattered distribution

over the state, and is according to Stanford native.

P. punctatum Ell. (P. acre HBK. in Gray's Manual 7th

ed.) For a complete synonymy see Rhodora 29 : 77, 1927 (Fig.

29) . Of a general distribution, but most of the collections seem

to have been made in the northern half of the state.

8 Fassett, N. C. Trans. Wis. Acad. 25 : 167, 1930.

221

Mahony — Reports on Flora of Wisconsin . XV.

222 Wisconsin Academy of Sciences, Arts, and Letters.

P. PUNCTATUM Ell. var. LEPTOSTACHYUM (Meisn.) Small, ( P .

acre HBK. var. leptostachyum Meisn. in Gray’s Manual 7th

ed.) (Fig. 30) is most common along the Mississippi and St.

Croix river valleys and in southern Wisconsin.

P . punctatum and P . punctatum var. leptostachyum are some¬

times very hard to distinguish and an accurate separation when

using herbarium sheets is almost impossible; although when

found growing together the two plants are sometimes quite dis¬

tinct.

P. orientals L. (Fig. 31) is a plant that has escaped from

gardens and has been collected in the southeastern part of the

state.

P. Persicaria L. (Fig. 32). This plant seems to follow the

Mississippi and St. Croix river valleys on the west, Lake Mich¬

igan shore line on the east, and is very abundant in the south¬

ern part of the state. This plant is an introduced species and

we should expect a scattered distribution.

P. H ydropi peroides Michx. (Fig. 33). Only four collections

have been made in the state.

Subgenus Tovar a (Adans.) Gray consists of only one species

which has a rather restricted distribution. It is found in the

eastern part of North America and the eastern part of Asia.

P. virginianum L. (Fig. 34). This plant has not been col¬

lected many times in the state, but those collections with a very

few exceptions have been made along the river valleys and

Lake Michigan shore line.

Sub genus Echinocaulon Meisn. has two species and both are

represented in Wisconsin. The plants of this subgenus, like

that of Tovar a, have a distribution in eastern North America

and eastern Asia.

P. arifolium L. (Fig. 35). This species seems to be found

largely in the southwestern half of the state.

P. sagittatum L. (Fig. 36) is a very common plant in Wis¬

consin. It seems probable that it might be collected from every

county in the state.

A green flowered form of this species has been described, f.

chloranthum Fernald. N. C. Fassett says: “Although this is

usually an estuarine form, green-flowered plants may occur

223

Mahony —Reports on Flora of Wisconsin . XV.

Fagopyrum esculentum Polygonella articulata

224 Wisconsin Academy of Sciences , Arts , and Letters .

elsewhere; I have found them in damp places at Ocean Point,

Maine, and in some parts of northern Wisconsin. These speci¬

mens lack the habit which is ordinarily characteristic of this

form, and which is probably a direct result of frequent submer¬

gence, for the estuarine plant has not only the green flowers in¬

dicated by the name, but slender stems and weak prickles.”4

Of the abundant material collected in Wisconsin that I have

worked with, I have been unable to find any specimens that

could be called f . chloranthum , although the flower coloring may

be white, pink or greenish.

Subgenus Tiniaria Meisn. consists of four species, three of

which are found in Wisconsin.

P. Convolvulus L. (Fig. 37) is of a general distribution in

Wisconsin. This plant has been introduced from Europe.

P. CILINODE Michx. (Fig. 38) is restricted to the northern

half of the state.

P. scandens L. (Fig. 39). This plant seems to have quite

a general distribution in the state except for the extreme north¬

east. I believe the plant to be more common in Wisconsin, how¬

ever, than the collections show.

Subgenus Pleuropterus (Turcz.) B. & H. consists of one spe¬

cies.

P. CUSPID ATUM Sieb. & Zucc. (Fig. 40). Only one speci¬

men from Wisconsin was found. This plant is often cultivated

in gardens and this one specimen might be from a plant that

had escaped from a nearby garden.

Fagopyrum

This genus is represented in Wisconsin by one cultivated spe¬

cies, the common buckwheat.

F. esculentum Moench. (Fig. 41). This plant being a cul¬

tivated plant and of some economic value in the state, has been

mapped by the State Department of Agriculture,5 and the map

in this study was taken from that publication and is as near

an exact copy as was possible to make. The original map

showed the farms over the state where buckwheat was grown.

4 Fassett, N. C. Proc. Bost. Soc. Nat. Hist. 39 : 107, 1928.

6 Wis, St. Dept, of Agr. Statistical Atlas of Wis. Agr., p. 49, 1926-1927.

Mahony — Reports on Flora of Wisconsin. XV.

225

As is evident from the map, the buckwheat region follows close¬

ly the St. Croixan Sandstone area which is outlined on the map.

POLYGONELLA

This genus is composed of two species, one of which is found

in Wisconsin.

P. articulata (L.) Meisn. (Fig. 42). This plant is quite

common in sandy areas over the state and seems to be particu¬

larly abundant in the northwestern part of the state.

PRELIMINARY REPORTS ON THE FLORA OF

WISCONSIN. XVI.

XYRIDALES

Norman C. Fassett

This report, and the four following it, are based on speci¬

mens in the herbaria of the Milwaukee Public Museum, North¬

land College, Mr. S. C. Wadmond, and the University of Wis¬

consin. The courtesies and cooperation of Mr. Smith and Mr.

Fuller, of Professor Bobb, and of Mr. Wadmond, are grate¬

fully acknowledged.

ERIOCAULACEAE— Pipewort Family

Eriocaulon septangulare With. E. articulatum Morong.

(Fig. 1). Usually found on the sandy or slightly peaty pond-

margins of northern Wisconsin. The circle in Trempeauleau

County represents an oral report from Mr. F. M. Uhler. A

sheet in the University herbarium bears the data: “Shebogen

Co,, Wisconsin. McM.” ; there is a pencilled comment by J. R.

Heddle : [ = Sheboygen Co. “McM.” = ? J. F. McMullen.] I

have seen no other material from the eastern counties.

XYRIDACEAE— Yellow-Eyed Grass Family

Xyris Montana Ries. (Fig. 2, cross) . Collected on July 23,

1896, on Lake Superior between Trout Stream and Long Island,

by L. S. Cheney. This is either in Iron County or in Ashland

County.

X. TORTA J. E. Smith. X. flexuosa Muhl. (Fig. 2, dots) . In

northwestern Wisconsin, definitely associated with small bodies

of water which are relicts of an almost extinct lake dating back

to early post-Wisconsin times1 ; in south-central Wisconsin prob¬

ably related to similar lakes. A more detailed discussion of this

and other like ranges is in preparation by Dr. W. T. McLaugh¬

lin.

1 See Aldrich & Fassett, Science 70 : 45-46. 1929.

228 Wisconsin Academy of Sciences , Arts, and Letters .

COMMELINACEAE — Spiderwort Family

Tradescantia reflexa Raf. Spiderwort (Fig. 8). Wad-

mond2 reports this (as T. Virginiana) as “common; along rail¬

road tracks, borders of woods, roadsides” in Racine and Ke¬

nosha Counties. Throughout much of southern Wisconsin it is

abundant on sand plains. Its range is quite typical of the prai¬

rie plants in the state; it is abundant across southern Wiscon¬

sin, but is not common in the interior of the Driftless Area ex¬

cept along the sand plains of the Mississippi and Wisconsin

Rivers and of the old Lake Wisconsin bed, and adventive in the

north after the forests have been cut from the sand barrens.

A specimen from Muscoda, with rose-colored petals, is prob¬

ably to be referred to f. Lesteri Standley, Rhodora 32 : 32.

1930. Forma albiflora Slavin & Nieuwland, Am. Midland

Nat. 11 : 600. 1929 has been collected at Boscobel, Spring Green,

Sparta, Fontana and Trempealeau.

T. OCCIDENTALIS (Britton) Smythe (Fig. 4). Sand plains

along our western borders on the Mississippi and St. Croix

Rivers ; adventive on the railroad tracks at Avoca, Iowa Coun¬

ty. Our specimens are very variable as to width of bracts and

denseness of pubescence ; some of them may be referable to T.

bracteata Small, if indeed T. occidentalis and T . bracteata are

distinct species. Our material has the capsules glabrous except

at summit, which would, according to the key in Rydberg’s

Rocky Mountain Flora, exclude them from both of these species.

Commelina communis L. Along roadsides in settlements,

escaping from cultivation ; Eau Claire, Pepin, Buffalo and Dane

Counties.

C. ERECTA L. A narrow-leaved plant, collected at Boscobel

in 1884, is apparently this species. It is without doubt a gar¬

den escape, and has probably not persisted.

PONTEDERIACEAE — Pickerel-weed Family

Pontederia cordata L. Pickerel-weed (Fig. 5, dots). Oc¬

casional throughout the state; least common in the Driftless

Area, where it is occasionally found along the Mississippi Riv¬

er. Forma latifolia (Raf.) House, N. Y. State Mus. Bui. 243-

2 Trans. Wis. Acad. 16 : 819. 1909.

Fassett — Reports on Flora of Wisconsin . XVI.

229

• X. torta

•Pontederla cor data

+ P cordata, f. latl folia

230 Wisconsin Academy of Sciences, Arts, and Letters .

244 : 62. 1923, is also common in the state ; it has the sides of

the leaf-blade rounded, instead of essentially straight as in

typical P. cor data. Forma angustifolia (Pursh) House, l . c .,

does not seem to have been collected in the state. The albino,

f. albiflora (Raf.) House, l. c., has been orally reported to

the writer by Mr. J. A. Moore of the Missouri Botanical Gar¬

den, as having been seen in Vilas County.

Heteranthera dubia (Jacq.) MacM. (Fig. 6). Across the

southern third of the state, and up the Mississippi River and

its tributaries. Although we have but four collections from

the Mississippi River, the plant is very common in that region ;

the true condition would probably be represented by a solid

line following that river in Fig. 6.

PRELIMINARY REPORTS ON THE FLORA OF

WISCONSIN. XVII.

MYRICACEAE; JUGLANDACEAE

Norman C. Fassett

MYRICACEAE— Sweet Gale Family

Myrica Gale L. Sweet Gale (Fig. 1, dots). Swales and

wet shores, northern Wisconsin, south to Manitowoc County.

M. Gale, var. subglabra (Chevalier) Fernald, Rhodora 16 :

167. 1914. (Fig. 2, crosses). This variety, distinguished by its

glabrous or glabrate lower leaf-surfaces, is said by Fernald to

replace the type in some areas. In Wisconsin it seems to be

much less common than typical M. Gale.

M. asplenifolia L. Sweet Fern (Fig. 2). Common in the

sandy and granitic areas of the state, and mostly absent from

the areas of limestone. But, like some of the Ericaceae1 2, it oc¬

curs about Green Bay, where the underlying rock is limy.

JUGLANDACEAE— Walnut Family

The four maps here presented are based on the unpublished

studies of Mr. L. S. Cheney, as were those of other groups of

trees previously published in this series. Localities represented

by herbarium specimens are indicated by large dots, being

shown only when they show the presence of the species where

it was not recorded by Cheney.

1. Juglans — Walnut

1. J. cinerea L. Butternut (Fig. 3). Cheney writes, “Jug¬

lans cinerea chooses a rich moist soil; it is therefore found

growing on low rocky hillsides, near the banks of streams, and

on alluvial lands, in the company of the black walnut, red oak,

white oak, bitternut and hackberry.”

2. J. nigra L. Black Walnut (Fig. 4; the crosses indicate

planted trees). “In Wisconsin, the tree is occasionally met

1 See Trans. Wis. Acad. 24 : 258. 1929.

232 Wisconsin Academy of Sciences, Arts, and Letters .

with in the wild state in all of the two southern tiers of coun¬

ties and in all counties fronting on the Mississippi River, north

to the St. Croix River; from Vernon County northward it is

confined to the immediate neighborhood of the Mississippi Riv¬

er or its largest tributaries. This tree chooses rich alluvial

bottom-lands or rich hillsides as its natural habitat.” We have

a number of cases of southern plants which follow up the Mis¬

sissippi River in this manner; see, for example, Juniper us vir-

giniana2, Anemone patens var. Wolfgangiana 3, and Anemonella

thalictriodes* .

2. Carya — Hickory

We have, apparently, but two native hickories common in the

state. These are C. ovata and C . cordiformis. They are listed

by S. C. Wadmond4 (as C. alba and C . amara) as being com¬

mon in Racine and Kenosha Counties. Russel5 lists both from

Milwaukee County, adding C. glabra as reported from Wauwa¬

tosa, where probably planted. Cheney and True6 list C. alba

and C. amara from the Madison area. In his notes, Cheney

lists besides these uHicoria glabra odorata”, of which he says

a single tree grows near the edge of Lake Monona in the sub¬

urb of Elmside, Madison. This, according to Professor R. H.

Denniston, was a large tree, perhaps old enough to antedate

the settling of the city, and probably of natural occurrence. It

did not satisfactarily fit descriptions of C. glabra, and was pos¬

sibly of hybrid origin. Professor Denniston and the writer

were unable to find the tree on October 9, 1931 ; it has appar¬

ently been cut. Dr. Denniston speaks of having seen the tree

as recently as within the last ten years. Cheney’s notes are

fully thirty years old.

Mr. S. C. Wadmond has collected C. glabra in Delavan, where

it is a shade tree in the city.

With but two common species, our hickories are easily distin¬

guished. C. ovata may be recognized by the tuft of white hairs

on each tooth on the leaflets, and C. cordiformis by its yellow

buds.

2 Trans. Wis. Acad. 25 : 180. 1930.

3 Ibid. 207.

*Ibid. 16 : 825. 1909.

e Bull. Wis. Nat. Hist. Soc. 5 : 186. 1907.

6 Trans. Wis. Acad. 9 : 98. 1892.

Fassett— Reports on Flora of Wisconsin . XVII. 238

My^ica aspleoif ©lla

234 Wisconsin Academy of Sciences , Arts, and Letters .

1. C. ovata (Mill.) K. Koch. Shag-bark Hickory (Fig. 5).

“This hickory is usually found growing on low hills or in the

vicinity of streams and swamps, in rather deep, rich, and only

moderately moist soils. With us its most constant companions

are the oaks, bitter nut and the hard maples.”

2 C. cordiformis (Wang.) K. Koch. Bitter Nut Hickory

(Fig. 6). “The bitter nut selects as its home low wet woods

near the borders of streams and swamps, or high rolling up¬

lands. It is commonly associated in our territory with the

hickory, the hackberry, the oaks, and in the northern part of

the state with the yellow birch, basswood and hard maples.”

PRELIMINARY REPORTS ON THE FLORA OF

WISCONSIN. XVIII.

SARRACENIALES.

Florence B. Livergood

SARRACENIACEAE— Pitcher-plant Family

Sarracenia purpurea L. Pitcher-plant (Fig. 1). Abun¬

dant in Sphagnum bogs, mostly in the glaciated areas of Wis¬

consin. (The Driftless Area, occupying the southwestern quar¬

ter of the state, is indicated on the map.)

DROSERACEAE — Sun-dew Family

Some confusion as to the identification of the species of Dro -

sera, especially between D. anglica and D. intermedia, has been

found in herbaria. To aid future collectors in the state, Dr. R.

I. Evans has prepared Fig. 6, showing the leaf-outlines of our

four species.

Drosera rotundifolia L. (Figs. 2 and 6A). Mostly in the

glaciated areas.

D. anglica Huds. D. longifolia L., in part; not of Gray's

Manual, ed. 7 (Figs. 3 and 6B). Lake Superior region; rare.

D. intermedia Hayne. D. longifolia L., in part ; Gray’s Man¬

ual, ed. 7 (Fig. 4, dots; Fig. 6C). Occasional in the glaciated

parts of the state. Subcaulescent forms collected in Bayfield,

Ashland, Sawyer and Marquette counties correspond to the

form described by J. R. Churchill in Rhodora 2 : 70-71. 1900,

and appear to be f. subcaulescens Mellvill, Mem. & Proc.

Manchester Lit. & Phil. Soc. 4 : ser. IV : 195. 1891; Diels,

Pflanzenreich 4 : pt. 112 : 84. 1906. (Fig. 4, crosses).

D. linearis Goldie (Figs. 5 and 6D). Rare. Interrogation

marks in Polk and Columbia Counties indicate old collections

without precise location.

236 Wisconsin Academy of Sciences, Arts, and Letters,

PRELIMINARY REPORTS ON THE FLORA OF

WISCONSIN. XIX.

SAXIFRAGACEAE

Norman C. Fassett

1. SULLIVANTIA

S. RENIFOLIA Rosendahl, Univ. Minn. Stud. Biol. Sci. 6 : 410,

pi. 43. 1927. S. Sullivantii, in part, of Gray’s Manual, ed. 7

(Fig. 1). Moist cliffs along streams in the Driftless Area. As

treated by Dr. Rosendahl, this is one of the six very localized

species of the genus; true S. Sullivantii occurs mostly south of

the glaciated area in Ohio, S. renifolia is in the Driftless Area

of Wisconsin and neighboring states, while the four remaining

species are localized in the Far West. The Driftless Area, oc¬

cupying the southwestern quarter of the state, is outlined on

the map.

2. Saxifraga— Saxifrage

S. pennsylvanica L. (Fig. 2). In swamps and on moist

cliffs throughout the state.

[ Tiarella cor difolia is reported in Gray’s Manual, ed. 7, as

being found westward to Minnesota; it should be sought in Wis¬

consin.]

3. Heuchera— Alum Root

H. hispida Pursh. (Fig. 3). On sandstone ledges and wood¬

ed sandy banks. Wadmond1 records this as being common on

prairies. Apparently more common southward, venturing

northward along the Wisconsin, Chippewa and St. Croix Riv¬

ers.

4. Mitella — Bishop’s Cap

1. M. diphylla L. (Fig. 4). Woodlands, apparently through¬

out the state. Forma opposxtifolia (Rydb.) Rosendahl, Eng-

ler, Bot. Jahrb. 50, suppl. : 380. 1914, characterized by having

1 Trans. Wis. Acad. 16 : 841. 1909.

238 Wisconsin Academy of Sciences, Arts, and Letters .

the cauline leaves short-petioled, is about as common (Fig. 5).

A collection from Ashland Junction, June 8, 1930, Otto West-

lund no. 150, consists of three specimens; two (in the Univer¬

sity herbarium) are, respectively, typical M. diphylla and f. op-

positifolia, while the third (in the herbarium of Northland Col¬

lege) is f. triphylla Rosendahl, 1. c ., with two petioled leaves

and a smaller sessile on higher up on the stem. A specimen

from Lynxville, September 3, 1915, /. /. Davis, lacks cauline

leaves, but appears otherwise normal (Fig. 5, circle).

2. M. nuda L. (Fig. 6). Northern Wisconsin, coming south

in the eastern counties. This is a common type of distribution :

see, for example, Pyrola chlorantha ,2 P. a sarifolia var. incarna-

ta,3 Corylus cornuta,4 and Betula lutea 4 Forma intermedia

(Bruhin) Rosendahl, l. c., p. 383, originally described from

Manitowoc County, is considered by Rosendahl to be a hybrid

with M. diphylla . Also Lake Owen, Bayfield Co., Gris com

12280.

5. Chrysosplenium— -Golden Saxifrage

C. americanum Schwein. (Fig. 7). In wet woods and

springy places northward, coming south to Manitowoc on the

Lake Michigan shore, and to cool woods in Vernon County and

cold canyons of the Dells of the Wisconsin River.

[C. tetrandrum Fries, which has been collected at Decorah,

Iowa, should be sought, probably on damp cliffs in the Drift¬

less Area.]

6. Parnassia — Grass of Parnassus

1. P. PARVIFLORA DC. (Fig. 8, dots). Known in the state

only from Door County. The range as mapped includes two

sheets in the Gray Herbarium, reported to me by Mr. C. A.

Weatherby.

2. P. multiseta Fernald, Rhodora 28 : 211. 1926. P. pa-

lustris of Gray’s Manual, ed. 7; not L. (Fig. 8, crosses),

known in the state only from Douglas County.

3. P. CAROLINIAN a Michx. (Fig. 9). In meadows, mostly

in the limy parts of the state (see page 233, fig. 2, for illustra¬

tion showing extent of limestone area).

2 Trans. Wis. Acad. 24 : 259, fig. 6. 1929.

*Ibid., 260, fig. 12.

4 Ibid., 25 : 191. 1930.

239

Fassett — Reports on Flora of Wisconsin . XIX.

Mitel la diphylla,

f, oppoaitifolla

Micella auda

240 Wisconsin Academy of Sciences, Arts, and Letters,

7. Philadelphia— Syringa

P. coronarius L. Escaped in Columbia, Dane and Rock

Counties.

8. Ribes— Currant ; Gooseberry

Through the cooperation of Mr. T. F. Kouba, the collections

of this genus made in Wisconsin by the Division of Blister Rust

Control, U. S. D. A., have been studied. This is in addition to

the herbaria listed on page 227.

The gooseberries, particularly, are a complex group and pre¬

sent many problems. While most of the specimens may be

identified readily, some may show exceptions to almost every

character listed in the key.

A. Flowers solitary or in bunches of 2-4, with spines at the base of each

bunch (The Gooseberries)

B. Calyx-lobes shorter than the tube; petioles with long hairs which

are usually unbranched, rarely slightly fringed, often gland-tipped ;

ovaries and berries usually prickly; peduncles 13-27 mm. long in

fruit

C. Ovaries and berries prickly . . R. Cynosbati

C. Ovaries and berries nearly or quite lacking prickles .

. . . . R. Cynosbati f. inerme

B. Calyx-lobes about equalling or longer than the tube; petioles with

long copiously fringed hairs; ovaries and berries never prickly ; pe¬

duncles and pedicels together 1-20 mm. long in fruit

D. Stamens about 1 cm. long, about twice as long as the calyx-

lobes, long-exserted, very conspicuous; bracts of the inflorescence

fringed with minute stalked red or yellowish glands ; peduncles

and pedicels together 8-20 mm. long in fruit; spines usually

stout, reddish, about 1 cm. long, rarely absent . . . R. missouriense

D. Stamens about equalling the calyx-lobes, or often slightly ex-

serted, not very conspicuous ; bracts of the inflorescence fringed

with minute hairs, but without glands in our common species ;

spines slender, about 5 mm. long, or often absent

E. Stamens about twice as long as the petals ; leaves and bracts

without glands ; leaf-blades wedge-shaped at base; peduncles

and pedicels together 3-10 mm. long in fruit

F. Leaves glabrous or somewhat pubescent beneath .

. . . . . R. hirtellum

F. Leaves velvety beneath ...... R. hirtellum var. calcicola

E. Stamens about equalling the petals ; leaves (particularly the

petioles and under sides of the blades) and bracts with mi¬

nute stalked glands intermixed with the fine hairs; leaf-

Fassett— -Reports on Flora of Wisconsin . XIX .

241

blades truncate or subcordate at base; pedicels 1-2 mm. long

in fruit, hardly exceeding the bud-scales . . R. oxyacanthoides

A. Flowers in racemes (The Currants)

G. Flowers about as broad as long

H. Ovaries without stalked glands

I. Calyx bell-shaped, with a well-developed tube; leaves usually

with resinous dots on the lower surface; bases of petioles

without gland-tipped hairs or stalked glands

J. Calyx-tube about equalling the lobes; racemes drooping;

flowers greenish-yellow; leaves often with resinous dots on

the upper surface

K. Bract at the base of each pedicel longer than the pedi¬

cel; calyx 8-10 mm. long; ovaries without resinous

dots; young twigs with a ridge running down from the

base of each petiole

L. Leaves with copious resinous dots beneath, and

sparsely dotted on the upper surface .

. R. americanum

L. Leaves without dots above, and nearly or quite lack¬

ing them on the lower surface . . .

. R. americanum f. pauciglandulosum

K. Bract shorter than the pedicel; calyx 5-6 mm. long;

ovaries with resinous dots; twigs without ridges run-

ing down from the petioles . R. nigrum

J. Calyx-tube much shorter than the lobes; racemes erect;

flowers white; leaves without resinous dots on the upper

surface; ovaries and fruits with resinous dots .

. R. hudsonianum

I. Calyx saucer-shaped, the united portion flat and scarcely

tube-like; leaves frequently with hairs, but never with res¬

inous dots, on the lower surface; bases of petioles with long

gland-tipped hairs, or with small stalked glands

M. Flowers yellowish or greenish; pedicels without stalked

glands; middle lobe of leaf ovate, a little longer than

broad . R. sativum

M. Flowers purple; pedicels with minute stalked glands; mid¬

dle lobe of leaf triangular, broader than long

N. Leaf-blades densely hairy beneath . R. triste

N. Leaf -blades nearly or quite without hairs .

. R. triste var. albinervium

H. Ovaries with stalked glands

O. Stems densely covered with prickles . . R. lacustre

0. Stems without prickles

242 Wisconsin Academy of Sciences , Arts , and Letters .

P. Calyx 2-2.5 mm. long . . . R. prostratum

P. Calyx 3-4 mm. long .... R. prostratum var. wiseonsinum

G. Flowers several times as long as broad . R, odoratum

1. R. Cynosbati L. Prickly Gooseberry. (Fig. 10, dots).

Abundant throughout the state. Forma inerme Rehder, Mitt.

Deutsch. Dendr. Ges. for 1910 : 250. 1910 (fig. 10, crosses),

which lacks most or all of the prickles from the berries, is oc¬

casional in the state, sometimes grading into the type.

2. R. mxssourxense Nutt. R. gracile of Gray's Manual, ed.

7; not Michx. Missouri Gooseberry. (Fig. 11). Common ex¬

cept northward.

3. R. hirtellum Michx. R. oxyacanthoides of Gray's Man¬

ual, ed. 7, in large part; not L. (Fig. 12, dots). Throughout

the state, but not common southward, where it is usually found

in bogs. Nearly all our material has the leaves more or less

pubescent beneath ; plants from Door and Racine Counties may

be referred to var. calcicola Fernald, which is also represent¬

ed in the Gray Herbarium by a sheet from Door County.

A specimen collected at Prentice, July 18, 1919, by J. J. Da¬

vis, lacks the plumose hairs on the petiole, and has in their

place unbranched hairs and glands suggestive of R. Cynosbati ,

but the short peduncles and sparsely pubescent leaf -blades with

cuneate bases seem to place it with R. hirtellum . A specimen

from Newbold, July 8, 1893, L. S. Cheney no. 1507, has the

leaves, but not the bracts, somewhat glandular.

4. R. oxyacanthoides L. Hawthorn-leaved Gooseberry.

The only Wisconsin specimen I have been able to refer to this

species is one from Ellison Bay, Door County, May 30, 1926,

E. J . Kraus et al This plant is unarmed, and the leaves are

extremely velvety, but it is otherwise characteristic.

5. R. americanum Mill. R. floridum L’Her. American

Black Currant (Fig. 13, dots). Common throughout the state,

except, apparently, in a large central area; perhaps future col¬

lections will show a uniform distribution throughout Wiscon¬

sin.

R. americanum f. pauciglandulosum, n. f., foliis supra sine

guttis et subtus saepe sine guttis.- — Algoma, June 1, 1905, Lou

Damas (type in the Herbarium of the University of Wiscon¬

sin) ; Delavan, June 2, 1907, S. C. Wadmond no. 1866; ap-

Fassett— Reports on Flora of Wisconsin . XIX. 243

ChrysoBplenium americanum

• Parnassia parvi flora

+ P. multioeta

f. inerrae

• Ribes hirtellum

+ R. hlrtellum,

var, calcicola

244 Wisconsin Academy of Sciences, Arts, and Letters .

proached by a specimen from Baraboo, May 8, 1927, N. C. Fas -

sett no. 5568. (Fig. 18, crosses).

6. R. NIGRUM L. Black Currant. Rarely escaping from cul¬

tivation ; collections have been made in Ashland, Door and Dane

Counties.

7. R. HUDSONIANUM Richards. Hudson Bay Currant. (Fig.

14) . Northward.

8. R. lacustre (Pers.) Poir. Swamp Black Currant. (Fig.

15) . Mostly northeastward; a single collection from Lake Su¬

perior shore.

9. R. prostratum L’Her. Skunk Currant. (Fig. 16, dots).

Across the northern half of the state.

R. prostratum var. wisconsinum n. var., calycibus 3-4 mm.

longis. — Bayfield Co. : wet sand, shore of Lake Superior, Port

Wing, June 14, 1928, Ludlow Griscom no. 12323 (type in the

Herbarium of Ludlow Griscom) ; poorly drained area, with

alder and aspen, near Cornucopia, June 5, 1931, T. F. Kouba;

Sand Island, June 20, 1897, L. S. Cheney no. 6155; mainland

east of Sand Bay, June 22, 1897, L. S. Cheney no. 6296. Bar¬

ron Co. : occasionally damp soil, Barron, May 26, 1917, Charles

Goessl no. 6814. Dunn Co.: moist woods, Wheeler, May 19,

1917, Charles Goessl no. 6720. (Fig. 16, crosses).

In its long calyx this resembles R. laxiflorum Pursh, of

northwestern America, to which it is perhaps as closely related

as to our eastern skunk currant. In associating it with the

latter species, the writer has taken into consideration the fol¬

lowing facts. The calyx-lobes of R . prostratum are character¬

ized both by Rydberg1 and by Coville and Britton2 as being

smooth, to distinguish them from the pubescent or glandular

calyces of R. laxiflorum. But many specimens of the former

have a sparse pubescence or even a few scattered glands on the

calyx, while R. laxiflorum var. japonicum is described by Berg¬

er3 as having glabrous sepals. A character which does not

seem to be noted in our manuals is found in the stipules ; these

are represented in R. prostratum by mere narrow subscarious

margins toward the bases of the petioles, while in R. laxiflorum

»F1. Rocky Mts., ed. 2 : 397. 1922.

2 N. Am. FI. 22, pt. 3 : 194. 1908.

3 N. Y. State Agric. Exper. Sta. Tech. Bull. no. 109 : 64. 1924,

Fassett— Reports on Flora of Wisconsin. XIX. 245

• Rites amerioanu®

+ R, americanum,

f. pauci$landulosum

Rites 1'acustre

+ R, prostratum,

var. wisconeinum

Rites triste

var. altinervium

246 Wisconsin Academy of Sciences, Arts, and Letters.

they are broader, and subtruncate at the summit in most cases.

The writer is not qualified to judge how generally this char¬

acter may hold, since he has at his disposal only five sheets of

the latter species, including var. coloradense. In placing the

plant of the Lake Superior region with R. prostratum rather

than with R. laxiflorum, he has placed reliance on the glabrous

or slightly glandular and pubescent calyces, on the narrow stip-

ular bases of the petioles, and on the narrow petals. The fruit

has not been collected.

Attention was first called to this plant by Professor Ludlow

Griscom, to whom the writer is indebted for the loan of the ma¬

terial cited as type. Professor Griscom was impressed by its

erect wand-like branches, utterly unlike the usual prostrate or

trailing stems of typical R. prostratum . Several of the collec¬

tions cited above appear to have had erect stems, but in ab¬

sence of collector's data it is impossible to say whether or not

that habit is always associated with the larger calyx. This

point must rest with future field observers.

A sheet from Kingston, Ontario, May 14, 1903, J. Fowler,

appears to belong with this variety.

10. R. sativum Syme. R. vulgare of Gray's Manual, ed. 7 ;

not Lam. Garden Currant. Occasionally escaping ; we have col¬

lections and reports from Waucapa, Dane, Rock, Racine4 and

Milwaukee5 Counties. This often appears as if native ; in Wau¬

paca County the writer found it growing in wet woods with R.

triste.

11. R. triste Pall. Swamp Currant. (Fig. 17). Northern

and eastern. Var. albinervium (Michx.) Fernald (fig. 18)

has essentially the same general range in Wisconsin, but there

seem to be few areas where both the typical form and the

variety occur, as may be seen by a comparison of figures 17

and 18. The locations of specimens intermediate between the

species and the variety are shown by circles on figure 18.

12. R. ODORATUM Wendland. R. aureum of Gray’s Manual,

ed. 7; not Pursh. Occasionally found persisting or escaping

from cultivation; it has been collected in St. Croix, Barron,

Waupaca, Outagamie, Vernon, Dane, Rock, Racine4 and Ke¬

nosha4 Counties.

4Wadmond, Trans. Wis. Acad. 16 : 842. 1909.

6 Russel, Bull. Wis. Nat. Hist. Soc. 5 : 200. 1907.

PRELIMINARY REPORTS ON THE FLORA OF

WISCONSIN. XX.

MALVALES

Alice M. Hagen

MALVACEAE — Mallow Family

Abutilon Theophrasti Medic. Velvet Leaf. (Fig. 1, dots) .

Occurs chiefly in waste places, fields and ditches, across south¬

ern Wisconsin. Introduced, but well established.

Althaea officinalis L. Marsh Mallow. (Fig. 1, cross).

Only one collection, from Lynxville, Crawford County. Intro¬

duced.

Malva rotundifolia L. Cheeses. (Fig. 2). Fairly com¬

mon in cultivated land and farm-yards in southern and western

Wisconsin. This species seems to show a very definite range,

and it will be interesting to note whether future collections will

extend the range throughout the eastern and northern parts

of the state. Naturalized from Europe.

M. borealis Walmm. ; Bergman, Geol. & Nat. Hist. Surv.

Minn. 4 : 437-440. 1916. (Fig. 3, triangles). Sepals with

long-ciliate margins; petals as long as or slightly longer than

the sepals; carpels 8-10, their backs conspicuously reticulated

and nearly or quite glabrous, their margins angled and some¬

times slightly toothed. M. rotundifolia , with which this has

been confused, is distinguished as follows : sepals not ciliate or

slightly so with short hairs ; petals 3-4 times the length of the

sepals ; carpels 12-15, their backs densely pubescent with short

hairs and but slightly if at all reticulated, and margins rounded.

While M. borealis is much less common than is M. rotundi¬

folia, it seems to occur farther to the northeast than does the

latter species.

M. crispa L. Curled Mallow. But one collection, from Ra¬

cine.

M. sylvestris L. High Mallow. (Fig. 3, crosses). An in¬

troduced species which has been collected three times in south¬

eastern Wisconsin.

248 Wisconsin Academy of Sciences , Arts , and Letters,

M. moschata L. Musk Mallow. (Fig. 3, dots). Natural¬

ized in southern and eastern Wisconsin.

Callirhoe triangulata (Leavenw.) Gray. Poppy Mallow.

(Fig. 4, dots). A native species, following dry prairies along

the Mississippi and Wisconsin Rivers.

Napaea dioica L. Glade Mallow. (Fig. 4, crosses). A na¬

tive plant which is locally abundant, mostly along streams, in a

small section in southwestern Wisconsin. This is mostly in

the Driftless Area, and to a small extent on ground covered

by the Illinoian but not the Wisconsin glaciation.

Hibiscus militaris Cav. Halberd-leaved Rose Mallow.

(Fig. 5, crosses). Bottomlands along the Mississippi and Wis¬

consin Rivers, in Grant County.

H. Trionum L. Flower-of-an-hour. (Fig. 5, dots). Found

chiefly in the southern portion of the state, and along the Wis¬

consin River, on waste land. A locally established plant

brought in from Europe. Its early introduction is shown by a

specimen from Marquette County (precise location not given)

collected by John Townley in 1861.

TILIACEAE — Linden Family

Tilia glabra Vent. An. Hist. Nat. 2 : 62. 1800; Sargent, Bot.

Gaz. 66 : 424. 1918; Bush, Bull. Torr. Bot. Club 54 : 235. 1927.

T, americana Am. Auct., but probably not L., or at least in very

small part. (Fig. 6). The large dots on the map represent

herbarium specimens, while the more complete range, shown

by the small dots, is taken from an unpublished map by L. S.

Cheney. Range very similar to that of the sugar maple.1

iSee Trans. Wis. Acad. 25 : 195-196. 1930.

249

Hagen — Reports on Flora of Wisconsin. XX.

* Malva borealis • Callirhoe triangulata

♦ Malva sylvestris + Napaea dloica

• Malva moschata

• Hibiscus Trionum Tilia glabra

+ Hibiscus militaris

STANDARDS FOR PREDICTING BASAL METABOLISM:

I. STANDARDS FOR GIRLS FROM 17 TO 21.1

Marian E. Stark

Contribution from the Department of Medicine and the Wisconsin General

Hospital , University of Wisconsin .

(Submitted for publication November 1, 1931.)

Contents

Introduction _ _ _ _ _ _ _ pp. 251-255

Experimental. Plan of study; collaborators; securing of normal ma¬

terial; standards of selection; tests; technique; measurements

made; tables of fundamental data _ _ _ _. _ pp. 255-267

Analysis of Data _ _ _ _ _ _ _ _ pp. 267-317

General studies of data on the standard series.

Geographical distribution of material; classification by colleges;

thyroid observations; variations between duplicate runs; pulse

rates _ _ _ _ _ _ _ _ _ _ _ _ pp. 267-277

Statistical analysis of data and the final prediction standard.

Formulas chosen; measurement of sitting height; the factor of

age; statistical constants for different age groups; final predic¬

tion equation and prediction table _ . _ _ pp. 277-289

Tests of fitness of the Wisconsin prediction standard and of pre¬

existing standards for girls of 17 to 21. Outline of study; pre¬

diction standards investigated; test series of normal controls

other than the Wisconsin standard series; tests of suitability of

prediction standards; comparative findings _ _ _ _pp. 289-317

Summary . _ _ _ _ _ _ _ _ _ pp. 317-318

Conclusions _ _ _ _ _ _ _ _ _pp. 318-319

Introduction

Anyone who has worked at all extensively in the field of basal

metabolism will have recognized that its two chief uncertainties

are (1) the securing of true “basal” conditions, — i. e., complete

muscular and also mental repose — on the part of the subjects ;

and (2) the proper choice of normal standards with which

clinical tests must be compared for interpretation.

1 The expense of printing this study has been in part defrayed through a grant

from the research funds of the Medical School of the University of Wisconsin.

252 Wisconsin Academy of Sciences , Arts , and Letters .

Since techniques have become so thoroughly simplified and

standardized and the physical conditions necessary for the

“basal” state in general well understood, the difficulties of the

first of these problems have been materially minimized. Though

the simplification of apparatus has resulted chiefly of course in

wider availability and greater accuracy of the determination it¬

self, it has also been reflected in lessened apprehension and

hence automatically better “basal” condition on the part of the

subjects. This, with the better understanding of the need for

mental as well as physical repose, is undoubtedly largely re¬

sponsible for the fact that the metabolic rates that are being

reported nowadays in connection with studies of normals in

various laboratories are definitely lower than they used to be

(1, 2); for mental unrest, as well as most forms of technical

error, lead to temporary actual, or apparent elevation of the de¬

termined rates. With all technical errors ruled out, the diffi¬

culty of controlling the mental state of the subjects must al¬

ways remain the chief variable in a determination which is so

peculiarly subject to psychic influences.

This is one of the reasons for the existence of the second

problem — namely, the difficulty of deciding upon adequate nor¬

mal standards. It is not surprising that most of the existing

standards, based on older data, are found too high to express

normal rates as they are being determined today. In clinical

work this is not such a striking handicap as in studies that

deal with normal controls, because of the mental hazard that is

bound to interfere more or less with the perfect relaxation of

subjects who are hospitalized for any cause whatever. How¬

ever, because of the unaccountability of this factor, most peo¬

ple prefer to have standards for clinical comparisons that are

based on “physiological” rather than “hospital” normals, and

to make such practical allowances for interpretation as are dic¬

tated by weighing of all the clinical evidence for any particular

case at hand.

The type of individual cooperation that must be obtained

from the subjects of metabolism experiments is undoubtedly

the largest single factor which has delayed for so long the com¬

piling of adequate standards of normality for all ages. After

acceptable data are obtained in sufficient quantity there comes

the choice of the proper method of analysis for building up

the prediction standards for this complex “physiological con-

Stark — Standards for Predicting Basal Metabolism . 253

stant” whose ultimate basis for constancy is not and perhaps

never can be fully understood, and which in practice is found

to have so many, often subtle, factors of variability.

Nevertheless, working standings of comparison had to be

built up before the science could advance very far. This was

made possible by the first large series of carefully controlled

data on normal subjects made available, that collected by Bene¬

dict and his collaborators of the Nutrition Laboratory at Bos¬

ton. It was natural that most of these early data should be on

adults and that the majority of these would be men. Upon

this body of data, including as Benedict offered it, 136 men and

103 women, are founded in whole or in large part all of the

major prediction standards that are in use today for adults of

both sexes. The two classical standards among these are of

course that of Aub and DuBois, the pioneer in the field in 1917,

and that of Harris and Benedict which appeared in 1919.

Theoretically the question of superior validity of one or the

other of the analytical methods involved in the two standards

or any of the modified forms that have been proposed for one

or the other, leaves so much to be said on both sides that it is

doubtful if a clean-cut choice can ever be made on purely rea¬

sonable grounds. Meanwhile, either of the original standards

has proved to work out with a high degree of practical satisfac¬

tion in dealing with adults. Of late, practical choice is perhaps

less often than formerly being left to precept or professional

“inheritance” since it is becoming increasingly evident that the

Harris-Benedict prediction, whose figures average throughout

lower than those of Aub and DuBois, affords by that fact a

better approximation to average normal findings under mod¬

ern test conditions.

It is not the purpose of this paper to discuss the adult stand¬

ards excepting as their consideration must bear directly upon

the subject at hand, namely the metabolism of girls and young

women. This sex and age group represents one of the least ade¬

quately surveyed portions of the whole field of practical basal

metabolism. In perhaps no other group is it more desirable for

the physician to have every reliable means possible at his dis¬

posal for dealing with metabolic difficulties.

Part of this inadequacy is undoubtedly due to comparative

dearth of data for this particular group during the time when

the classical reference standards for basal metabolism were be-

254 Wisconsin Academy of Sciences , Arts, and Letters .

in g established. Far more, however, would seem to be due to

the widely divergent conceptions which have been brought to

bear by men of similar authority in the held upon the choice of

material to be studied and the type of analysis to which it

should be submitted for establishing reference standards for

the transitional years between childhood and fully adult devel¬

opment. The standards made available have differed so seri¬

ously that the result has been utter confusion to the physician

who would try to interpret the results of basal metabolism tests

on his younger feminine patients.

Among the many metabolic studies that have dealt with nor¬

mal controls as the technique of metabolism measurements has

become more widely available there have been numerous ones

which included girls and young women as subjects; but while

many of these series have been used for extensive comparisons,

none of them in this country has been worked into a prediction

standard to take the place of those in the field up to 1924.

The standards proposed from authoritative sources in this

country for predicting the metabolism of girls under 21 have

been two : the lower ranges of the Aub-DuBois adult standard

(3), which give expected calories per square meter of body

surface in two-year age periods between 14 and 20; and the

standard recommended in 1924 by Benedict (U) specifically for

girls from 12 through 20, since the Harris-Benedict tables were

limited by their authors to ages over 20. These special tables

which are based upon the data obtained by Benedict and Hen¬

dry (5) three years earlier on Girl Scouts between 12 and 17,

give calories per kilogram of weight according to age. The

predictions to be obtained by the two standards disagree to a

startling extent, though the discrepancies do not run parallel

for all types of cases.

Such a state of affairs is particularly disconcerting in a me¬

tabolism laboratory such as that of the Wisconsin General Hos¬

pital, with which is associated the Student Health Department

of the State University, and in which consequently young wom¬

en of college age come in for a large share of the metabolic

studies that are carried on.

The help that the clinician feels justified in expecting from

the determination of the metabolic rate is by no means rea¬

lized in these younger subjects. And yet the demand for the

determination has continued to increase year by year, in spite

Stark — Standards for Predicting Basal Metabolism . 255

of disappointments,-— perhaps on the basis of occasional clear-

cut cases, but mostly, no doubt, because of the unquestioned

helpfulness of this laboratory adjunct in practice with adults.

For these reasons the laboratory undertook a few years ago

to gather enough normal data from the same source as its

clinical material to make possible either a decisive selection

between existing standards for these ages, or possibly to modify

one of these or to serve, if necessary, for the building of stand¬

ards of comparison of its own. The material to be offered in

this paper is the result of this study.

Experimental

Plan of Study. Our early systematic comparisons proved so

thoroughly to confirm our suspicion of the need for a new pre¬

diction that we formulated our plans to include this possibility

definitely. This realized on the basis of our first nearly 100

cases between the ages of 17 and 21, the rest of our study has

taken the form of comparisons in which our own and the other

available predictions have been '‘tried on” the individuals of

our group to ascertain not only average fitness in various re¬

spects, but whole ranges of individual variations, with which,

rather than averages, of course the clinician would be con¬

cerned in dealing with individual cases. Finally, it has been

possible to impose the really crucial test of making these vari¬

ous comparisons on a comparable number of normals other than

our own, on whom data have become available through various

studies made independently here, and collected from recent re¬

ports in the literature by workers in widely different localities.

The gathering together of these data and the formation of

concrete comparisons based on a significantly large body of

measurements, have been illuminating to us, and we offer them

in the hope that they may carry some practical suggestions to

others who are interested in the same problems, either on the

side of physiological normals in general, or for clinical applica¬

tion.

A study of this sort must be a cooperative venture. Any

description of the present one must fittingly begin with ac¬

knowledgment of essential collaboration not only of the sub¬

jects of the study themselves, but of personnel from three dif¬

ferent departments of the University and Medical School.

256 Wisconsin Academy of Sciences , Arts , and Letters.

Normal girls of the desired ages were obtained as subjects

through the cooperation of the Student Health Department of

the University, who were themselves of course particularly in¬

terested in the availability of suitable prediction standards for

this group. The members of the staff who thus made the work

possible were Dr. William A. Mowry, Dr. Sarah I. Morris and

Dr. Irma Backe, chief physician, and associate and assistant

physicians, respectively, of the department.

Our statistical advisor throughout the project has been Pro¬

fessor Mark Ingraham of the Department of Mathematics,

whose advice has been followed as to the type of prediction

standard built up after statistical analysis of the original data

on our group; and to whom we are indebted for helpful sug¬

gestions throughout the remainder of the studies and compari¬

sons that have been made. The extensive computations for

the original analyses and for the final prediction standard were

performed by Miss Beatrice Berberich, University Computer,

under the direction of Professor Ingraham.

The work as a whole was suggested in the beginning and has

gone forward under the continuous sponsorship of Dr. E. L.

Sevringhaus, Associate Professor of Medicine and in charge of

the metabolic department of the hospital. Without his support

and inspiration the project could neither have been begun nor

brought to conclusion.

Acknowledgment is due, finally, to Dr. E. F. DuBois for his

careful reading of the manuscript, and for the advantage of

stimulating suggestions and criticisms by which the report in

its final form has profited.

Policy in Securing of Subjects. In the favorable portions of

three school years data were collected on 97 normal girls of

ages 17 through 20, students in the University of Wisconsin.

As often as possible, more than one test was performed upon

each subject. 163 acceptable tests in all were secured upon the

individuals of the group.

The policy adopted was that of making metabolism appoint¬

ments for suitable entering students as a routine part of their

physical examinations. In this way as many cases as possible

were handled through the fall until the interference of exami¬

nation or vacation programs made it profitless to carry the rou¬

tine schedules any further. These were then supplemented by

Stark - — Standards for Predicting Basal Metabolism . 257

personal letters of request for cooperation, bearing brief but

frank explanation of the project, during various later portions

of each school year.

Of the students who were thus invited to cooperate, rather

than having appointments made for them as part of a more

or less required routine, a good proportion of course refused or

simply failed to appear. Thus it would seem that a different

type of selection had operated with these than with the ones

who were studied each fall. This is doubtless true to a certain

extent, but as a matter of fact, a good many of those sched¬

uled in the fall also failed to appear, with characteristic mod¬

ern independence. No effort was made to coerce the unwilling,

though at times follow-up letters were sent, or personal con¬

versations obtained in the effort to prevent rumors that might

lead to mistrust in the motives of the project or the spreading

of misconceptions as to the nature of the test itself. Either

undue coercion or much anxiety over the test, would, we felt,

operate against our getting the best results. On the other

hand, experience indicated that too much interest on the part

of the subject might do the same thing. We tried therefore

to steer a middle course between telling too little and too much.

Choice of Normal Material. The students were taken with¬

out any effort at selection (beyond our having considered the

age distribution we wanted) other than the requirement of a

grading of “A” in their physical and medical examinations. A

rating of “A” does not imply physical perfection, to be sure.

It merely means that by the type of routine examination pos¬

sible for such a group, no definite defects were judged to be

present and no contraindications in the past history or pres¬

ent findings to the student’s participation in the full University

program, including physical education. Such a criterion of

normality would at least seem to compare favorably with any

that have governed the choice of normal material for other

considerable groups forming the bases of metabolism studies.

In every case the subject was examined and interviewed by

the author in order to supplement certain points of the history

and physical observations that were subjects of special interest

in the investigation, and to rule out acute or accidental reasons

for unfitness. In this supplementary examination the girls were

questioned particularly as to menstrual history and the use

of goitre prophylaxis measures, and their necks were exam-

258 Wisconsin Academy of Sciences, Arts, and Letters .

ined by gross inspection and palpation of the thyroid gland. On

the basis of these findings and questionings a number of sub¬

jects on whom good tests had been secured had to be discarded.

One of the factors which make a careful study of this sort

so time-consuming is the variety of reasons that can disqualify

a subject even after she has come for the test. Some of the

students who were secured as subjects proved unsuitable for

such reasons as temporary indispositions (usually from respira¬

tory infections) ; fatigue from excessive school or social activi¬

ties ; occasional technical difficulties in the test itself ; and rarely

the subject’s failure to cooperate in the routine.

Experience showed it to be undesirable to attempt to get

satisfactory results at certain times of the school year, such as

the periods soon before or after vacations or examinations, or

prominent social events. Athletic exhibitions were found to

result in definitely disqualifying fatigue in the cases of some

of the students from the Physical Education Department, which

furnished us otherwise some of our most satisfactory and co¬

operative subjects.

A considerable group of subjects were discarded because of a

history of menstrual difficulties or irregularities which some¬

times had arisen since the girl had entered the University, or

if present before, had not been counted against her in the physi¬

cal examination, either because she had failed to mention it, or

because it was apparently nothing that should interfere with

ordinary University activities. These subjects were discarded

nevertheless without regard to the tests they gave because the

relation between metabolic rate and menstrual difficulties in

general is still so poorly understood. This was indeed one of

the phases of the problem in which we were specifically inter¬

ested.

The standards which governed us in this choice were arrived

at after consultation with several of the clinical staff had indi¬

cated what could safely be taken as the borderline between nor¬

mal variations and the suspicion of pathology. Dysmenorrhea

that was more than slight or moderate, — i. e., incapacitating, —

or irregularity in time of periods of more than one week were

classed as disqualifying the subjects. The mere presence of

catamenia was not allowed to rule out a test. The large amount

of work that is recorded on this subject, as well as general ob¬

servation, have led us to the opinion that unless actual discom-

Stark— Standards for Predicting Basal Metabolism . 259

fort is present during the test, the metabolic rate in normal

women is to be found during the menstrual period within the

limits of incidental fluctuations to be expected at any time dur¬

ing the cycle. Of course active discomfort from menstruation,

as from any other source, would rule out the results of a test.

A record was kept with the majority of the tests, of the time of

onset of the nearest menstrual period in case any correlations of

interest could later be made.

Since Wisconsin is in a goitre belt region, we were particu¬

larly interested in learning whatever we could about any dem¬

onstrable relation between thyroid development and metabolic

rate of subjects in apparently normal health. — It has been ob¬

served that some degree of thyroid enlargement is so common

in our Middle States that even the perfect young women repro¬

duced on magazine covers are apt to show slight goitres ! Cer¬

tainly it would have been impossible to assemble a significant

sized group of girls in this part of the country if one were to

exclude those with thyroid enlargement. As long as our other

requirements for normality were met, we could not discrimi¬

nate against even slight or moderate “goitres’1. Of course no

subject was accepted in whom symptoms were admitted or

could be suspected from the history or questioning.

We also could not discriminate against those who had used

iodine for prophylaxis ; but we do not consider that the medica¬

tions used were ever of a character that could conceivably set

this group apart from others from regions where the natural

ration can be counted on to furnish the necessary iodine.

In order to make later comparisons possible, we kept records

in each case that would allow grouping of the subjects as to

geographical source and thyroid development in relation to me¬

tabolic rate. These results will be given with the other data.

The Tests, The tests were made in the metabolism labora¬

tory of the Wisconsin General Hospital. The experimental sub¬

jects were purposely studied under the same general conditions

as the regular patients, but in the interests of unity and con¬

trol in the series, were all handled exclusively by the author,

with very rare exceptions when one of the hospital operators

filled in in case of need.

Each student who was given an appointment was provided

with the instructions for preparation and brief description of

260 Wisconsin Academy of Sciences, Arts, and Letters .

the routine which the hospital furnishes to out-patients who

have had no previous experience with the test. Though a few

of the subjects proved to have had similar tests before in dif¬

ferent capacities, by far the largest majority were first tests.

No practice runs were made, for a large proportion of strictly

first tests in the series was felt to be desirable in making it

comparable with the general run of out-patients for whom

standards of comparison were being sought.

The usual requirements for preparation were followed, — i. e.,

the subjects came without breakfast, and were required to lie

quietly, with enough covering to be comfortable, for 30 minutes

before the test. The appointments were all between 7 :30 and

8:30 a. m., and the students excused from University exercises

for the necessary periods.

Technique. A closed circuit type of apparatus was used in

all the experiments, — in an occasional test at the beginning of

the series the “Sanborn Grafic” was used, but for all the rest,

the Benedict Portable model with motor-driven circulation.

With this equipment of course it is not possible to determine

the respiratory quotient. This we do not feel to be a disad¬

vantage for this kind of study, however, in that the most ex¬

treme variations that might conceivably be encountered in R.

Q. of normal subjects under the standardized conditions could

make only about a 3% difference in the figured metabolic rates.

This is well within the range of variation of the latter to be

expected for determinations on the same individual at different

times, or even between immediately consecutive short runs

within a single test. Since there seems to be no good reason

in the present state of our knowledge to assume a very differ¬

ent probability for variations above or below the commonly as¬

sumed average of 0.82, 2 the errors incurred by assuming the

latter can very well compensate each other for all practical pur¬

poses. It is probably true moreover that the variations in re-

2 A caloric equivalent of 4.825 per liter of oxygen has been used, as is custo¬

mary in calculating the heat-production from the measurement of the oxygen

alone. The actual calculations have used Carpenter’s tables, based on the values

given by Lusk’s modification of Zuntz and Schumberg for a non-protein R. Q. of

0.82. (Ref: Carpenter, T. M., : Tables, Factors and Formulas for Computing

Respiratory Exchange and Biological Transformations of Energy, Carnegie Inst,

of Wash. Publ. 303A, 1924.) Dr. BuBois (personal communication, 1931) states

that this value of oxygen corresponds more nearly to an average R. Q. of 0.86, if

we take the protein factor into consideration. The validity of the assumption

is not affected.

Stark — Standards for Predicting Basal Metabolism . 261

ported R. Q. due to experimental error greatly exceed its true

physiological variability. In the hands of all but a very few

workers, then, the substituting of the technique of gas analysis

for that of the simple oxygen determination, we feel is apt to

result in a net loss, rather than gain, in the dependability of

the final results.

Single runs were customarily of 8-minutes’ duration, and at

least two acceptable runs were required for each test. The low¬

est run was accepted as determining the rate for any given test,

but all accepted runs were figured separately for the informa¬

tion to be gained by their comparison. This is our routine prac¬

tice, since it would hardly seem as though the subject’s actual

metabolical level would change materially within a short basal

period, whereas whatever technical errors could conceivably in¬

fluence the results with the technique we use, as well as the

less readily ruled-out failure of the subject to relax, would tend

to increase, rather than decrease the readings.

Readings of the pulse-rate were made near the beginning and

end of each run for supplementary evidence as to the degree of

relaxation of the subject. The machines were of course main¬

tained in good condition and kept at all times free from leaks

or other defects. During each run a confirmatory leak-test

was made by Benedict’s expedient of placing a weight of about

50 gm. on the spirometer midway through the run, in which

case a leak in any of the connections would be detected by a

change in slope of the respiratory tracing from the point where

the weight was added.

The criterion for acceptability of a given test was not any

pre-conceived standard of closeness of agreement between the

constituent runs within the test nor, within any reasonable

limits, of consistency of results from one test to another or with

any of the existing standards of normality. Rather, the stand

was taken that unless there was demonstrated some definite

and convincing reason for disqualifying the subject or discard¬

ing the test, the latter must be retained ; and that the first ac¬

cepted test on each of the subjects was to be used as the basis

of the final analyses, regardless of whether subsequent tests on

the same individuals turned out to be better or worse in any

detail. This plan was adopted rather than either of the two

obvious alternatives that had to be considered,-— i. e., of averag¬

ing all the available tests for each subject, or choosing there-

262 Wisconsin Academy of Sciences , Arts, and Letters .

from the lowest in each case as probably representing more

nearly true “basal” conditions. The idea that underlay this

choice was that we were interested primarily in arriving at

some concrete ideas, not of ideal rates of heat-production, and

not only of their average probable level ; but rather in the whole

normal range of variability within which individual determina¬

tions might reasonably be expected to fall for subjects of the

general types we have to deal with and to study under compar¬

able well-controlled practical conditions.

Measurements that were made. Our tentative goal before

undertaking any extensive analyses was 100 cases, to be dis¬

tributed equally among the four years from 17 through 20, for

whom the need of adequate standards has been most keenly

felt. It chanced that the collaboration of the newly-organized

statistical service of the Department of Mathematics was placed

at our disposal when we were within a short distance of this

goal, and the final series submitted to the analyses upon which

our prediction standard was based included 97 individuals,

among whom ages and accepted tests were distributed as fol¬

lows:

The individual measurements which were made upon this group

are given in Table I.

A few additional subjects who were accepted after the analy¬

ses were under way or after the quotas for their year had been

filled are included in the supplementary series of Wisconsin

subjects who were used for later comparisons.

Of the 97 subjects in the standard series, 58 were accepted

twice and 6 three times. Intervals between tests were 3 weeks

in 35, and 7 to 55 days in the remainder. The range of total

intervals in cases taken three times was 26 days to 191/2

months.

Four of the girls appear in two different age-groups, and so

are each counted as two different individuals in the analyses.

Three of these were drawn the second time in response to a

special appeal to their Physiology class to help fill the 19 and

Stark — Standards for Predicting Basal Metabolism . 263

20 year old groups. They were Physical Education students

who as usual were interested and ready to help. The fourth

was called in for a third test because she had appeared poorly

relaxed in her second, although her first had been entirely ac¬

ceptable. By the time she was obtained the third time more

than 15 months had passed since her second test (and 16 since

her first), so that she now belonged in the next higher age

group. Since there was no valid objection to either her first

or third test, it was felt that both should be retained. Since

16 months was the shortest interval between any of the first

tests at the different ages it was held legitimate to consider

them as representing different individuals for purposes of

analysis in these few cases. Furthermore, of course most of

the measurements on the girls had changed during the inter¬

vals.

Age was taken to the nearest birthday. The superscripts

with age given in the data tables refer to the 1st or 2nd half

of the designated year, — i. e., the subjects marked 162 or 171

would all have been considered as 17.

Weight was calculated without clothing. The subjects were

instructed to undress and lie down for the rest-period in gowns

that were furnished, and the known weight of these gowns was

allowed for in recording the weight of the subjects.

Height was measured both standing and sitting for almost

all of the subjects, with the idea that perhaps, in view of the

emphasis that has been placed upon the value of the sitting

height by various observers, it might prove a more represen¬

tative measurement for use in predicting the metabolism than

the standing height usually employed. The results of compari¬

sons in this regard will be mentioned in connection with the

statistical analysis (p. 278).

Calories per 24 hours represent the “basal” heat-production

calculated from the oxygen consumption as measured during

the lowest run within any given test, by the usual assumption

of the average post-absorptive respiratory quotient for which

the calorific equivalent of oxygen is 0.004825 per cc. (See

foot-note, p. 260.)

In calculating Calories per Square Meter per hour, the sur¬

face area is determined by the height-weight chart of DuBois

and DuBois.

264 Wisconsin Academy of Sciences, Arts, and Letters,

Table I. Measurements on 97 subjects used for the University of Wisconsin

standard. In the second column, headed “Age”, the small numbers printed

,as superscripts refer to the first or second half of the designated year.

Stark— -Standards for Predicting Basal Metabolism . 265

Table I. Continued

266 Wisconsin Academy of Sciences, Arts, and Letters ,

Table I. Continued

Stark— Standards for Predicting Basal Metabolism . 267

Table I. Continued

Metabolic Rates represent the customary percentage devia¬

tion of the measured basal heat-production from a given pre¬

diction for the individual. The comparisons based on their

calculation according to the various available predictions are

reserved for the final section on analysis of data.

Analysis of Data

The analyses which we have carried out may be presented

under three general heads:

I. Generalizations and comparisons among the data on the

original 97 individuals upon which our prediction standard

was built.

II. Statistical analysis and the final prediction standard

based on the data from this group.

III. Testing of this standard in comparison with the oth¬

ers that are available for this age-range:

1) Applied to the individuals of our own standard

series; and finally

2) Applied to a comparable series of different individu¬

als upon whom data have become available partly

268 Wisconsin Academy of Sciences , Arts, and Letters .

from additional studies here, and partly from data

given in the recent literature by other investigators

from widely different localities.

The various comparisons will be offered wherever possible in

the form of summary tables, with only brief additional discus¬

sion. When metabolic rates are given in these general com¬

parisons, they are figured according to our own prediction

(based on this group), and refer unless otherwise stated to the

lowest value found for each individual.

I. General Studies of the Data within the Wisconsin Series.

Geographical Distribution of Material. Most of the students,

(80%) as has been indicated were drafted from newly entering

classes, so that their prevailing living conditions can be taken

as determined by the places which they called their homes.

Wisconsin material of course predominated very largely (58 of

the 97) ; but it was interesting to note that the rest of the

group represented 16 other states. Of these, as would be ex¬

pected, other mid-western states were in the majority. Illi¬

nois furnished 11, Ohio 8, Indiana 3, Iowa and Michigan each 2.

Of the rest, 2 were from Pennsylvania, and one each from Mis¬

sissippi, Missouri, Kansas, Alabama, New Jersey, New York,

Idaho, New Hampshire, Virginia and California. An effort was

made to subdivide these into goitrous and non-goitrous sources,

using the maps of McClendon (6) as guides. The number of

subjects from the former so far exceeded the latter that no fair

comparisons could be made. The comparative showings, how¬

ever, are appended in connection with the thyroid observations.

Classification by Colleges was made chiefly to find how our

group would compare with the student body at large, and fur¬

ther, how large a percentage of the total was represented by

Physical Education students, since these had seemed to be

easier to obtain as subjects than other girls in general. This

is the only feature of the classification which would seem to

merit quoting. The Physical Education group was found to

have constituted just 1/3 (32 out of 96 who could be classified)

of the total, which is three times the proportion in a represen¬

tative Wisconsin freshman cl&ss, and six times that among all

women registered at the time of the study. This preponderance

can be attributed to several factors: the physical fitness of

Stark — Standards for Predicting Basal Metabolism . 269

these girls, their early registration for their physical examina¬

tions, and their consistent willingness to cooperate. We were,

interested to know, in view of the various comparisons that

have been recorded which show definite influence on the me¬

tabolic level of such factors as muscular development and de¬

gree of activity, whether these girls as a group would show any

such distinctions from the relatively non-athletically inclined

students.

A very general comparison in the matter of heat-production

which was all that was felt warranted, showed practically the

same percentage of plus and minus rates among the athletic

and non-athletic groups. It is true, of course, that these girls

were really just starting their intensive athletic careers, though

they must have inclined naturally to more vigorous lives than

the average. Some investigators have maintained that indi¬

viduals who lead a very active life show greater variability in

metabolic rate from day to day than do those of a more seden¬

tary occupation. In a study of this kind no such tendency could

be detected, and it might well be the case that in comparison

with other untrained subjects it would be more than counter¬

acted in such a group as this by the stabilizing effect of su¬

perior cooperativeness and understanding of the aims of the

experiments.

Thyroid Observations. A survey of the notes made on cervi¬

cal inspection and palpation of the subjects in our standard

series resulted in their grouping under 4 rough classifications

as to thyroid condition. The groupings have been designated

as follows: “0” indicates that neither isthmus nor lobes were

palpated; “1” that lobes or (and) isthmus were just palpated

but not seen; that either or both were considered both

palpable and slightly visible; and “3” that there was definite

prominence of the gland, amounting to what could be consid¬

ered slight or moderate degree of goitre. These observations

are not meant to indicate that any sharp classifications were or

could be attempted under the conditions of the examinations.

They are grouped and presented merely for any general interest

they may suggest, or comparisons for future studies.

Groupings have been made in two ways: Table II gives dis¬

tributions according to geographic sources of subjects, together

with the types of metabolic rates found in these groups. Table

270 Wisconsin Academy of Sciences, Arts, and Letters .

Ill gives the relation between thyroid observations and (a)

ages; and (b) metabolic rates in the group as a whole.

Reference to Table II will show that the slighter degrees of

prominence of the gland are not confined to subjects from the

so-called goitre regions. To be sure, it was not possible by the

manner in which the survey was undertaken to establish how

long the reported residences had been in effect. Comparisons

with a group of girls of the same age who have grown up in

strictly non-goitrous regions would be interesting. That both

Table II Thyroid observations according to geographical source of sub¬

jects. Wisconsin Standard Series.

♦Michigan: reported Iodine in drinking water.

fPennsyl vania: 1; Illinois: 1.

the lowest and the highest rates observed for the whole group

occurred in subjects from the goitrous areas may or may not

be entirely coincidental with the greater number of subjects

from this group of states.

The most striking feature shown in Table III is the exceed¬

ing rarity of entirely negative thyroids, — i. e., cases in which

neither isthmus nor lobes were found visible or palpable. There

were only two such cases in our whole series of 97 girls.

Stark — Standards for Predicting Basal Metabolism. 271

Table III Thyroid observations according to ( a ) age ; and (b) metabolic rates.

Wisconsin Standard Series.

Total =92. (One not classified. Three are individuals who appear in two

different age-groups, and hence are only classified once)

Nearly 1/3 of these subjects had visible glands. More than

1/4 showed what could be called slight or moderate goitre.

They were of course as far as could be ascertained free from

any suspicion of thyroid symptoms, else they would not have

been accepted as normal subjects for the study.

Plus and minus metabolic rates are quite equally distributed

in groups “2” and '‘3”. There are definitely more minus than

plus rates among the negative and practically negative groups

(“0” and T).

Variations between Duplicate Runs in the Same Test and be¬

tween Different Tests on the Same Individuals: (a) Duplicate

Runs within Tests . The closeness of agreement between dupli¬

cate runs in a given test is an index of the technical accepta-

272 Wisconsin Academy of Sciences , Arts, and Letters .

bility of the test and of the success with which relaxation has

been maintained by the subject. Anyone who works in the field

of course realizes that incidental and wholly unaccountable fac¬

tors of variation make exact agreement between duplicate runs

rare. Many laboratories specify agreement within 5% of the

duplicate runs within a test as constituting a satisfactory test.

In dealing with the general run of sick patients the practical

allowance is usually extended to 10%. We purposely refrained

from specifying any such arbitrary standards of acceptability

in advance in order to test by impartial comparisons the fre¬

quency with which these expectations built up in hospital prac¬

tice would be fulfilled in our series of untrained subjects in nor¬

mal health.

In Table IV are summarized comparisons between successive

runs in all of the accepted tests on the individuals of our stand¬

ard series plus the seven extra subjects from the same group.

There is doubtless a larger proportion of perfect agreement

and small differences in this series than one could expect in any

clinical group of comparable size; but our general impression

gained from experience is fulfilled, namely that a large majority

of well-controlled tests would agree within 5%, while there

would always be a few larger differences. To sum up the table

Table IV. Agreement of duplicate runs within tests*. Wisconsin Standard Series

+7 extra subjects from the same group.

♦The percentage deviations are figured in each case on the basis of the low run.

They are therefore larger than if they had been given as deviations from their

mean.

Stark — Standards for Predicting Basal Metabolism . 273

itself: it is seen that all of the duplicates agree within 10%,

about 9/10 within 5%, and about 3/4 within 3%.

It seems rather surprising that notably more cases showed

the lowest values in the first runs. One might perhaps con¬

clude from this that on the whole the preliminary rest-period

had succeeded in building up a good degree of relaxation and

that there was seldom initial tenseness because of the strange¬

ness of the procedure. It is to be remembered that first runs

for most of these subjects meant absolutely the first experi¬

ence with the test. It is to be noted further, that most of the

largest discrepancies (5—10% difference between successive

runs) were among the group who gave the lowest readings in

the first runs. Perhaps these were the lively or excitable types

who found it difficult to remain perfectly quiet for long at a

time. They would correspond to the class of irritable and easily

fatigued patients with whom nothing is to be gained by too

persistent striving for good results at any one appointment.

(b) Repeated Tests on Same Individuals. For intelligent

practical use of the metabolism test it is essential to have some

working conception of the range of variability of rates that one

can reasonably expect to find at different times in the same in¬

dividual. This tendency to vary evidently entirely independent¬

ly of observational errors, is of course itself extremely variable

in subjects or groups of subjects of different types, degrees of

training, or in different circumstances.

DuBois considers (7, p. 130 f.) that one may expect differ¬

ences of about the same order of magnitude in almost any large

series of tests made on the same man over a long period of

time as one finds in a large group of normal men of the same

age and size,— i. e., almost all of the determinations will fall

within 10% of the average, with a few variations between 10

and 15%. Some of the records of individual series which he

quotes showed striking and consistent uniformity, though oc¬

casional rates that deviated to an extreme degree from the

mean of the series without apparent reason are mentioned in

at least two of the classical series. He quotes Magnus-Levy as

pointing out the significance of this, namely that we cannot

trust too much in the tests of any one day.

Griffith et al (8) in a recent extensive study of 5 normals

list the extreme “intra-individual variabilities” that they found

as falling between 8.6 and 11.3% of the respective yearly

274 Wisconsin Academy of Sciences, Arts, and Letters.

means. They made 55-246 determinations apiece on the 5 sub¬

jects, over periods of 1-2 years.

Comparable conclusions cannot be drawn from studies such

as ours, of course, in which each individual was examined only

two or three times. However the summary of such a study of

a significantly large and representative group of untrained sub¬

jects should offer a fairly logical basis of comparison for an

average group of untrained patients of the same types for

which the series is designed to serve as standard.

Table V summarizes the findings in our standard series.

Table V. Agreement between repeated tests on the same individuals * Wisconsin

Standard Series.

♦Differences are figured on the basis of the calories calculated in the first ac¬

cepted test for each individual, i. e., they represent successive changes in measured

heat-production for the individual, and not differences from a mean.

The total number represents the changes in 6 third tests, in addition to tests

repeated once.

That there is a majority tendency toward lower rates in tests

subsequent to the first one undoubtedly indicates in part the

true effect of experience with the routine. The significance of

this factor is minimized for the present group, at least, by the

smallness of the majority that showed downward changes, and

by the similar magnitudes and distribution of the changes in

the two groups. This does not hold true, however, for the

larger changes (greater than 10%), which were found with

Stark — Standards for Predicting Basal Metabolism. 275

one unimportant exception in the group whose rates decreased

with practice. These larger changes no doubt occurred in the

more nervous type of subject, and in at least one or two cases

where there was a definite suggestion that the girl had been

worried the first time by a suspicion that she had been singled

out for the test because she wasn’t as normal as she was told

she was ! This of course is a mild example of the “mental haz¬

ards” that operate to elevate the rates in hospital cases. It

suggests again the need for caution in judging borderline rates.

In summarizing mfer-individual variations of rates in the

final section of the present study (Table XVI) it is found

that 87 % of the rates in first tests of this same group fell with¬

in plus or minus 10% of the average normal prediction by our

standard and 100% within plus or minus 15%. Correspond¬

ing figures for a group of other normal girls of the same ages

collected for comparisons are 86 and 96% respectively. Al¬

lowing for the different ways in which the two types of devia¬

tions have been calculated, it appears that for a normal group

at least it is not necessary to expect quite as large a range of

intra- as inter-individual variability in making practical use

of the test. If observations are repeated frequently enough,

occasional larger deviations can be expected.

Pulse-Rates. Readings of the individual pulse-rates have

not been included in our tables of fundamental data because

the lability of this measurement and the fact that it varies only

roughly parallel with the metabolic rates from one individual

to another, are matters of common observation. For any given

individual of course the general type of pulse and any observed

tendency for it to change markedly during a metabolism test

are valuable supplementary data for the operator and the clini¬

cian. For the present study it has seemed of interest only to

present a summary of the average and extreme pulse-rates that

were noted in these normal girls, for general comparison with

other groups that have been studied. This summary is given

in Table VI.

It is observed that the group averages for pulse-rate through¬

out are only slightly lower or remain the same in tests after

the original ones. (The pulse-rates for first tests were aver¬

aged for this comparison of course only for those subjects who

had repeat tests.) This together with the low range of rates

276 Wisconsin Academy of Sciences , Arts , and Letters.

Table VI. Average and extreme pulse-rates by groups. All tests on

Wisconsin Standard Series. Each rate represents the average of counts

made near the beginning and near the end of each run , and refers to the

lowest run (or the average in two runs in which the metabolic rates may

have agreed exactly) for a given test.

can be taken as further evidence of a satisfactory degree of

repose on the part of the subjects as a whole.

As was the expectation for given individuals, deviations in

pulse-rate did not by any means always occur parallel in di¬

rection with changes in metabolic rate. The lowest pulse-rates

were not found invariably to be associated with the lowest me¬

tabolic rates, either with the same subjects in repeated tests,

or moreover in the series as a whole. Though different com¬

binations of forces are evidently effective to different degrees

in causing temporary fluctuations (within normal limits) in

these two physiological functions, we know of course that their

correlation is much more instructive when we come to deal with

cases of pathological metabolism.

Stark — Standards for Predicting Basal Metabolism . 277

A further point is of passing interest, — namely, the preva¬

lence of low pulse rates throughout our group in comparison

with what is generally considered as “normal” for a girl or

woman. Though the breathing of pure oxygen may have a

tendency to lower the pulse-rate, personal observations suggest

that this is probably not a factor of significance in such short-

period experiments and with normal individuals. A more logi¬

cal explanation would seem to be the fact that our common con¬

ceptions of “normality” are founded upon observations made

in non-basal conditions, whereas not enough truly basal meas¬

urements have been recorded to impress themselves on our

manner of thinking.

It is interesting to note that the minimum which we found

in three of our four groups of healthy American girls is lower

than the minimum (54) found by MacLeod, Crofts and Bene¬

dict (9) in their study of 9 Oriental young women who had

lived in this country for from 15 months to 4^2 years. The

unmistakable average tendency for what we consider low lev¬

els of such vital functions as pulse and metabolism in Orientals,

even after prolonged exposure to our conditions of living, is

often referred to as an indication of a constitutionally superior

capacity for repose on the part of these races, rather than an

essential difference in metabolic processes. It would appear,

then, that a good many of our girls were quite comparable to

the Orientals in this respect.

The more formal analyses upon which our prediction stand¬

ard was based are summarized in the following section.

II. Statistical Analyses of Data and the Final Predic¬

tion Standard.

It was decided to use the first accepted test only, on each

subject for the standard, so only these were submitted to the

formal analyses. Our reasons for this course have been suffi¬

ciently indicated in the foregoing discussion, but it is well to

keep the fact in mind throughout any comparisons that are

made.

A survey of the data on our first two groups of 25 each that

were completed — the 18 and 20 year groups — convinced Pro¬

fessor Ingraham that 25 subjects of any single age were suffi¬

cient to justify some rather extensive preliminary analyses.

278 Wisconsin Academy of Sciences , Arts , and Letters.

These preliminary analyses, together with consideration of

what has already been accomplished in the field, and of the

nature and needs of the specific problem at hand, indicated

quite definitely what types of study would be profitable to con¬

tinue and what should be dropped. The chief decisions thus

arrived at were three: (1) That the method of ordinary

linear correlation was suitable for expressing the relations in¬

dicated between metabolism and body measurements for the

groups studied. (2) That the use of the sitting height meas¬

urements did not add anything of value to the study. And

(3) That the volume of data on hand, though decidedly large

as studies of this sort go, and of definite statistical significance,

was still not by any means extensive enough to define with cer¬

tainty the age-factor within the narrow range of ages of the

present study. These points may bear some further brief con¬

sideration :

1. The Method of analysis. Two general types of formulas

that are suggested for studies of this sort were investigated, —

In one of the “normal” metabolism was expressed as a linear

function of the age, weight and height, and in the other the

logarithm of the metabolism was a linear function of the lo¬

garithms of the age, weight and height. Within the compara¬

tively small range of values here available for study, the two

gave such closely similar results (i. e., the curves were sensibly

alike) that in the absence of apparent theoretical reasons for

preferring the one rather than the other, the logarithmic for¬

mula was dropped in favor of the simpler method of calcula¬

tion.

2. The use of the measurement of sitting height. For the

two separate age-groups submitted to the preliminary analyses,

equations were formulated in which metabolism was correlated

with (a) weight and sitting-height alone; (b) weight and

standing height alone; and (c) weight and both sitting height

and standing height. It was immediately apparent that the

inclusion of both measurements for height did not increase the

accuracy of the prediction, so that (c) was dropped. In the

list of statistical constants figured for all four age-groups given

in Table VII will be found the standard errors of estimating

the metabolism by the first two forms of equation, the one in¬

cluding the sitting height and the other the standing height

measurement. These figures show the average expectations to

Stark— Standards for Predicting Basal Metabolism . 279

be the same in one group, and actually very slightly in favor

of the equations using standing heights in the other three. Ac¬

tual figuring of the individual metabolic rates by the two equa¬

tions for half the individuals of the total series showed a neg¬

ligible advantage in favor of the sitting-height equation in the

matter of extreme range of rates. There is the further prac¬

tical consideration in favor of the more common measurement,

namely, that the greater errors inherent in the making of the

smaller measurement, that of sitting height, would probably in

practice fully outweigh any theoretical advantage of using that

measurement as possibly being physiologically more represen¬

tative.

3. The age-factor. A problem of fundamental interest in

the study of heat-production from the physiological stand-point

is the general picture to be gained of the changes of metabolism

with changes in age throughout life. The status of information

along these lines, based upon extensive studies of composite

data on record up to about 1924, is instructively summarized

by DuBois in the second edition of his book, “Basal Metabolism

in Health and Disease” (7) in the chapter dealing with factors

which influence the normal basal metabolism. These composite

studies agree in indicating unmistakably that the metabolism

for both sexes when referred to height, weight or surface-area,

is relatively high in childhood and that some time around the

age of maturity the curves flatten out and henceforth show a

more or less gradual decline throughout adult life. Certain in¬

teresting details in the configuration of these curves still await

better definition, and with it, a better understanding of the

physiological processes effective at particular periods of devel¬

opment. Notably these are the disputed “jog” in the curve

just before or at the age of puberty, necessarily difficult to

confirm and fully define; and the exact point where the age-

curve flattens out as maturity becomes established. Lack of

statistically significant volumes of data for these transitional

years has been keenly felt by those interested in studying the

age-curves from either a practical or a theoretical point of view.

It was reasonable to hope that the range and relatively large

volume of homogeneous data of the present study could throw

light on the second of these two questions. That this hope

must remain incompletely realized until similar studies are ex-

280 Wisconsin Academy of Sciences , Arts, and Letters.

*20 individuals, only, measured for sitting height.

Table VIII. Averages , standard deviations and coefficients of variability of measurements', Wisconsin Standard Series by age-groups. All

first tests as used for standard.

Stark — Standards for Predicting Basal Metabolism. 281

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282 Wisconsin Academy of Sciences , Arts , and Letters,

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Stark— Standards for Predicting Basal Metabolism . 283

tended to cover a larger range of years becomes apparent when

we examine some of the data of the following tables.

In the summary of the more important of the statistical con¬

stants which were figured in the preparation of our prediction

(Table VIII) it is to be noted that the figures for average ca¬

lories per 24 hours in first tests for the separate ages, 17, 18,

19 and 20, do not show a consistent trend. The inconsistency

appears moreover not only in the figures for group averages,

but also in the slopes of the regression lines of metabolism on

weight or height as plotted for the separate years in the course

of our preliminary analyses (not here reproduced). Accidental

fluctuations can readily obscure the picture of the relatively

small changes in metabolism intrinsically due to changes of age

over these short periods, unless an impracticably large number

of cases could perhaps illustrate mass tendencies by smoothing

out the incidental irregularities.

A general trend downward with age is however indicated

when the series is considered in two two-year groups, 17-18 and

19-20 years, by which larger grouping some of the smoothing

out is accomplished. In the supplementary calculations of

Table Villa, this point is examined in greater detail. For bet¬

ter comparison the heat-production is now calculated as calories

per square meter per hour, as well as being given in the abso¬

lute figures for calories in 24 hours, and the two-year age-

groups are considered from various angles. The average body

measurements are shown to illustrate the average similarity

of the different groups. The following points are noted:

The downward tendency with increasing age is evident

throughout the data, whether we average all tests in each group,

or consider first and later tests separately for those subjects

who were observed more than once. In these latter sub-groups

the average differences with age, considering either first or

later tests, are in all cases larger than the differences within

the same age-groups between first and subsequent tests, wheth¬

er the heat-production is expressed as total calories or as ca¬

lories per square meter of body surface. Somewhat closer

analysis of the table suggests however that some of this differ¬

ence between the younger and the older girls in the matter of

metabolic level is apparent and due to better relaxation on the

average among the older girls. Looking at the figures for ca¬

lories per square meter per hour, it is noted that the older

284 Wisconsin Academy of Sciences, Arts, and Letters .

girls show a considerably smaller drop from first to later tests

than do the younger group. This may well be a type of pitfall

which makes valid generalizations admittedly so much more

difficult to make as we deal with less mature subjects.

That the data at hand justify the conclusion that there is a

definite downward trend of the metabolic level in girls between

17-18 and 19-20 years, however, is hardly to be questioned.

The same indication is seen in the larger composite series of

controls assembled for the comparisons of the last part of our

present study. Though the age-distribution among these 119

subjects is not regular enough to justify formal comparisons,

it is nevertheless interesting to find that the calories per square

meter per hour for the 31 subjects who were 17-18 years old

average 35.04, while the corresponding average is 33.94 for the

88 who were from 19 to 20 years old. The more definite out¬

lining of this portion of the age-curve can be looked forward

to with some degree of confidence when similarly large groups

of younger and perhaps some older subjects are studied in ex¬

actly comparable fashion. With this in mind we have made a

beginning upon a study similar to the present one of boys and

girls from the upper grammar school through high school ages.

Meanwhile we have met our pressing need for a workable

practical standard for predicting the metabolism of girls of

college ages not covered by the adult standards, by lumping

the data on our girls of 17, 18, 19 and 20, leaving the small

age-factor for the time being out of consideration. This pre¬

diction table is presented (Table X) in the hope that it may

also prove serviceable to others who have to meet problems

similar to our own in the clinical application of the basal me¬

tabolism test.

Table IX. Extreme variations in individual measurements Wisconsin Standard

Series. First tests, as used for standard.

S tark— Standards for Predicting Basal Metabolism . 285

Various data assembled in the preparation of the final stand¬

ard, which may be of interest for future comparisons, are sum¬

marized in Tables VII through IX*

In Table VII, in addition to the standard errors of estimat¬

ing the metabolism by equations using sitting and those using

standing-height, already referred to, are recorded the correla¬

tion coefficients obtained between the various measurements

made on our subjects. Table VIII gives the variabilities en¬

countered in the different measurements. These variabilities

have been indicated in the two ways which are apt to convey

the most meaning to the statistically-inclined reader: (1) in

terms of the Standard Deviation3; and (2) as “Coefficients of

AYW.

Variability” which express the percentage of the group average

in question represented by each standard deviation.

For the best determination of the relation of one variable,

such as level of heat-production, to one or more others with

which it is being correlated, it is desirable to analyze groups

in which the variables can be established at both extremes of

the normal ranges of measurements, rather than to have the

groups so homogeneous that all of the individual measure¬

ments fall within a narrow range of variation. Predictions

based on groups composed almost entirely of individuals of

average build would be bound to describe the average individual

well, but might or might not fit individuals who deviated mark¬

edly from the general average of a similar population in one

or another body measurement.

Our four groups were noticeably unequal in the matter of

homogeneity. Weight is of course the most significant variable

in influencing the level of the metabolism in groups of other¬

wise fairly similar individuals. While two of our groups in¬

cluded a satisfactory number of lighter and heavier individuals

to approximate the relationship between weight and heat-pro¬

duction fairly satisfactorily, the other two contained mainly

3 Standard Deviation, “a”, of a group of measurements (the usual root-mean-

square measure of dispersal) is equal to the square root of the average of the

squares of the deviations of individual measurements from their group average.

Statisticians us© the standard deviation because of considerations based on the

theory of dispersion of samples. It is worth pointing out that in choosing a pre¬

diction standard which minimizes the standard deviation of the error of estimate

rather than the average of their magnitudes, the larger deviations are weighted

more heavily than the smaller.

286 Wisconsin Academy of Sciences , Arts , and Letters .

individuals of average build. This indicates the most plausible

reason for the inconsistent slopes of the regression lines for

the separate years, noted above.

It is observed from Table VIII that statistically the 18 and 19

year groups showed considerably smaller variability in the fac¬

tor of weight than did the 17 and 20 year ones. While the

groups did not show the same order of differences in the con¬

stants which measure the average dispersion of standing or

sitting height measurements, Table IX brings out the fact that

extreme individual measurements for height4 were again found

only in the two outside groups, whereas the two middle ones

departed less in either direction from the general averages for

height, as well as weight.

The separate correlation coefficients (Table VII) are higher

in every case for the less homogeneous groups. The fact that

the standard error of estimate is proportionally lower for the

most homogeneous group of all, that of 19 years, merely ex¬

presses the fact that a prediction based on a group of indi¬

viduals who are nearly all alike is bound to describe closely

any individual whose measurements are similar to those of the

group average. This constant does not include any measure

of the success with which such a prediction would fit individu¬

als who differed from this group average in one or more re¬

spects.

Since body-measurements, and particularly weight, are apt

to be changing somewhat rapidly at these ages and under these

conditions of living, one would not, perhaps, expect the various

correlations to be as high as could be found in a group of

adults. Comparing the figures for our total group with those

published in the classical biometric study of Harris and Bene¬

dict (10), we find that in every instance our coefficients are

lower than those found for their group of 186 men, but are

higher than the average of their two series of women (total

number = 103).

These 103 women, on whom the Harris-Benedict prediction

for females is based, included 12 girls from the ages of 15

through 20 years, whose measurements were apparently in¬

cluded in all the general analyses, though the final prediction

4 Standing height, only, observed in this respect, since the desirability of the

sitting height measurement had by this time been decided against.

Stark— Standards for Predicting Basal Metabolism . 287

tables begin with 21 years, and the authors specify that their

equations should not be used for younger individuals.

Our objective findings throughout in connection with our

relatively large group of girls from 17 through 20 confirm the

feeling which we have entertained all along, — namely, that it

would have saved a great deal of confusion in the field of prac¬

tical metabolism if Benedict had considered these girls along

with his adult women, rather than thinking of them as more

safely classed physiologically with the younger Girl Scouts

whom he studied later in an entirely different way, and whose

curves he extrapolated upwards to include the years 17-20.

That the Harris-Benedict equation for adult women, used

without any modification, affords the best fit of any of the older

standards for both ours and other normals of the same ages

which we have been able to collect, we shall bring out in the

final section of this report. We have also extended the study

to indicate the amount by which the original standard, now

too high, may be lowered to “center” it for these groups, which

procedure then yields for the total series, by every criterion

which we have been able to apply, a fit that is almost as good

as that of our own standard, which was worked out by similar

methods, but specifically for these ages.

These various considerations, and the character of the sta¬

tistical constants which we have found in general, strengthen

us in our early impression that our four groups together con¬

stitute a series of representative significance for a study of

this sort.

The Wisconsin Prediction for Girls from 17 through 20. The

final equation arrived at by the standard methods of multiple

correlation for predicting the metabolism of these ages was :

h = 10.63 w + 3.23 s + 184.61

where h = heat-production in calories per 24 hours ; w = body-

weight in kilograms; and s = standing height in centimeters.

The predictions which this yields for a large range of body

measurements is embodied in the accompanying standard pre¬

diction table (Table X).

288 Wisconsin Academy of Sciences , Arts , and Letters,

Table X. University of Wisconsin

PREDICTION TABLE FOR BASAL METABOLISM OF YOUNG WOMEN

AGES 17 - 21

Height in Centimeters

Stark— Standards for Predicting Basal Metabolism . 289

III. Tests of Fit of the Available Prediction Standards

for Girls between 17 and 21.

Outline of Study. Four prediction standards, including our

own, have been subjected to comparative studies as applied to

the Wisconsin and other normals which we have collected.

These are:

1. The original Aub-DuBois tables.

2. The Harris-Benedict prediction for adult women, ex¬

trapolated for these ages under 21.

8. Benedict's special prediction for girls from 12 through

20; and

4. The University of Wisconsin prediction just de¬

scribed for girls from 17 through 20.

Based on the average showings of the rates as figured for

our original series, the first three predictions have been modi¬

fied by applying a constant percentage correction such as would

center them for our standard group. On this more legitimate

basis, all of the comparisons have been repeated, not only for

our series, but also the other normals tabulated, and the com¬

parative findings summarized.

The Standards that were Investigated: The Aub-DuBois.

A brief mention of the history of the Aub-DuBois prediction

as it bears directly on its application to the restricted group

at hand may not be amiss as a matter of interesting back¬

ground. The figures given for boys of these ages were arrived

at by DuBois with his collaborators by tentative development

of their composite curve for changes of metabolism for all

ages, over the admittedly dubious area between childhood and

21 years. DuBois studied a group of 8 Boy Scouts of 12 and

13 years (11) to try to define better the trend of his curve

through the pre-adolescent period. How nearly a truly “basal”

condition could be enforced upon these boys under the condi¬

tions of the tests is a matter of question which is left more or

less open by the author himself (12, p. 18b). Other data on

boys that were included were taken from Magnus-Levy and

Falk, whose measurements on adults were also used. Analyses

by Harris and Benedict show that in comparison with their own

and other then available data practically all of the measure¬

ments of the German investigators run high (10, pp. 232,

290 Wisconsin Academy of Sciences , Arts, and Letters .

236, 239, 2U2). This fact is probably important in account¬

ing for the observation that the Aub-DuBois predictions are

higher in general than the other American standards. To ob¬

tain the figures for girls, approximately 7% was subtracted

from each of the constants for the males of the same ages, as

was done throughout the table in the case of females. This

procedure was specified by DuBois to be entirely tentative (7,

V . 169).

The Harris-Benedict Adult Prediction, extrap . Our reasons

for trying out the predictions obtained by extrapolating the

Harris-Benedict tables for adults for these younger subjects

have already been referred to in connection with the statistical

comparisons among our own data, p. 287.

Benedict's Prediction for Girls. The Girl Scouts upon whom

this prediction is based were all from 12 to 17 years old, and

the figures from 17 to 21 were obtained by extrapolating the

curve that had been sketched through the data for the younger

girls. Benedict himself did not claim that these standards

were ideal or even entirely logical. But in 1924 when he found

it desirable to record recommendations for predicting the basal

metabolism of girls of various ages from then available data

(U), he reasoned that in the absence of sufficient evidence to

justify classing girls of these ages with adults in the matter

of metabolic stability, it was reasonable to assume that they

were still in the developmental period physiologically, and

hence to be considered with the younger girls.

The hope of coming to some more definite conclusions as to

the validity of this logic was not improved by the unusual way

in which the predictions for these younger girls, who set the

standards, were arrived at. To overcome the seemingly pro¬

hibitive difficulties at that time of trying to secure individual

cooperation in a significantly large number of such young sub¬

jects, Benedict and Hendry resorted to two unusual expedients :

(1) the group method of study; and (2) the recording of the

lowest rate of metabolism found during the night, while the

groups of girls slept in the large calorimeter chamber. While

Benedict maintained that the effect of sleep on the metabolic

rate was too much in doubt to invalidate the results of such a

study for comparison with waking metabolism under good

“basal” conditions, the fact remains that the figures found in

Stark — Standards for Predicting Basal Metabolism. 291

his tables are almost universally considered much too low for

practical use in the majority of cases.

One of the things that started us on the present study was

the disconcerting observation that the discrepancies between

the standards available for these ages from equally authorita¬

tive sources were so inconsistent, and, to us, unpredictable.

Thus it was not unusual to have to report to a clinician that the

metabolic rate of his patient had turned out to be something

like +12 or +13% according to Benedict’s standard, whereas

according to DuBois’ the rate found had been — 12 or — 13% !

Occasional cases without just suspicion of hyperthyroidism

might show rates as high as +40% by the Benedict predic¬

tion, which could hardly be put down to nervousness entirely,

since according to DuBois the rates were normal. On the other

hand, occasional subjects would show practically the same rate

by both the predictions. Feeling incapable of judging of the

superior merits of one or the other of these standards on theo¬

retical grounds, in the face of apparently better satisfaction

in practice now from one and now from the other, we decided

to leave the choice to a thoroughgoing series of comparisons on

normal subjects of the same ages as the patients who were

proving so baffling.

From time to time in the course of assembling the amount

of data which we set out to compare, we had transitory hopes

of finding our problem already solved for us by one or another

modification or adaptation of one of the major standards that

was apparently giving satisfaction in other quarters. The most

widely-circulated of these should be mentioned in this connec¬

tion:

Krogh’s Changes in the Aub-DuBois Tables . In 1925 there

appeared in connection with Krogh’s respiration apparatus

some tables in which the Aub-DuBois predictions were modi¬

fied by a uniform reduction of 6%, to meet more accurately

the unusually rigid conditions which Krogh exacted for the

performance of the standard “basal” test (13). Even had we

given attention to this proposal earlier than we did, we would

hardly have felt that Krogh’s recommendations, proposed un¬

der working conditions quite different from our own, would

justify our acceptance without a thorough trial. By this time,

moreover, we had become interested in making other compari-

292 Wisconsin Academy of Sciences , Arts , and Letters .

sons than merely the success with which a given standard would

predict the average absolute level of the metabolism of groups

of normal subjects like our own. Hence the Krogh tables need

not be considered as a separate standard.

Boothby and Sandiford’s Proposed Tables . More recently

Boothby and Saniford (14) have proposed more fundamental

revisions of the DuBois constants on the basis of the large

number of “hospital normals” whom they had studied prior to

1929 at the Mayo Clinic. Though their proposed constant are

somewhat lower than DuBois’ for girls of the ages in which we

are interested, they are still far too high to describe our normal

subjects. For this reason, in addition to our theoretical pref¬

erence for standards based on physiological rather than “hos¬

pital” normals, we have not considered their standard.

The Sanborn Tables . We have also not at any time consid¬

ered the widely-used predictions put out by Sanborn (15) with

the Benedict type of machine which he markets, since his modi¬

fied tables have been found inaccurate in some respects and

their manner of origin stigmatized as unscientific (16).

Kestner and Knipping’s Predictions. In Germany, Kestner

and Knipping (17) have extended and adapted the Harris-

Benedict prediction tables to fit all ages of both sexes. Though

their adaptations evidently afford an excellent fit for the Ger¬

man cases which they handle, they have not proved the solu¬

tion for our problems, since the original formulas have been

changed in the direction of raising the predictions for girls

under 21, whereas the whole tendency of our data is to lower

them. Hence these tables, also, have been left out of consid¬

eration in the comparisons which we have made.

Test Series of Normals other than the Wisconsin Standard

Series. A total series of data on 119 normal controls has been

collected for concrete comparisons as to the suitability of the

different prediction standards. The 119 individuals include

those subjects within the age-range of our original series fur¬

nished by the following sources:

Wisconsin Supplementary Series. (1) The 7 extra subjects

who were studied by the author after the data for the Wis¬

consin Standard were under analysis. (2) A series of 21 pre¬

sumably normal sorority girls studied by Jean Fish in 1928

in connection with a thesis in psychology for the degree of

Stark — Standards for Predicting Basal Metabolism, 293

Table XI. Subjects within the range of the University of Wisconsin prediction ,

reported by Remington and Culp (1931)*; student nurses , Medical College of the State

of South Carolina. Rates given refer to the first test for each subject.

♦Remington, Roe E., and Culp, F. B., Basal Metabolic Rate of Medical Students and Nurses in

Training in Charleston , S. C., Arch. Int. Med., 1931 XLVII, 366.

tRecalculated from Calories /sq. m./hr.

jAdded to data of authors.

294 Wisconsin Academy of Sciences , Arts, and Letters ,

Table XI. Continued

t Recalculated from Calories /sq. m./hr.

JAdded to data of authors.

§Omitted from calculations.

Stark— -Standards for Predicting Basal Metabolism . 295

Table XI. Continued

tRecalculated from Calories /sq. m./hr.

t Added to data of authors.

296 Wisconsin Academy of Sciences , Arts , and Letters,

Table XII. Subjects within the range of the University of Wisconsin prediction,

reported by Jennie Tilt (1980)* students at the State College for Women , Tallahassee,

Florida.

*Tilt, Jennie, J. Biol. Chem. 1930 LXXXVI, 635.

fRepresent averages of two or more determinations.

jRefigured from measurements given.

§Added to data of author.

Stark — Standards for Predicting Basal Metabolism . 297

Table XIII. Supplementary data from the University of Wisconsin

(a) Subjects by Jean Fish (1927-28, thesis in psychology).

(b) Subjects by Shipley (1930, thesis in psychology).

(c) Additional subjects by M. E. Stark (not included in U. W. Standard)

298 Wisconsin Academy of Sciences, Arts, and Letters .

Table XIV. Subjects within the range of the University of Wisconsin

; prediction , reported by Benedict (1928)*,. Supplementary series of nor¬

mals from the Nutrition Laboratory of Boston.

♦Benedict, F. G., “Basal Metabolism E)ata on Normal Men and Women (Series II) with

Some Considerations on the Use of Prediction Standards” — Am. J. Physiol., 1928, LXXXV, 607

t Average of several runs on 1-4 days.

jAdded to data of Author.

Master of Arts. (8) A series of 9 presumably normal stu¬

dents investigated by W. C. Shipley in 1930 in connection with

a problem in psychology.

These three groups together constitute a supplementary Wis¬

consin series of 37 cases which we were glad to have available

for comparisons. The two phychology studies were made inde¬

pendent of the hospital under carefully controlled experimental

conditions, for the physiological comparisons which they might

afford. Miss Fish and Mr. Shipley were both instructed in the

technique and fundamental principles of the metabolism meas¬

urements by the author.

Other Series: The Florida College for Women. For the

most crucial test of all we were fortunate in finding two large

series of normal controls that have been published in 1930 and

1931 : (4) A study of the basal metabolism of young college

women made by Jennie Tilt (18) at the State College for Wom¬

en at Tallahassee, Florida was published in the Journal of

Biological Chemistry for April, 1930. Among her 52 subjects

whose ages ranged from 17 to 25, there were 29 girls between

17 and 20 V2 years, so that they fell within the limits of our

group. Miss Tilt used technique similar to ours and calculated

the rates of her subjects by both the Harris-Benedict and the

Aub-DuBois predictions. By both standards the rates of most

of her subjects were decidedly low, and she interpreted her find¬

ings as evidence that the basal metabolism of young women in

Florida tends to be significantly lower than that predicted for

young women of the same ages and living in the north.

Stark — Standards for Predicting Basal Metabolism. 299

Student Nurses in S. Carolina. (5) More recently still, there

appeared a study by Remington and Culp (19) of basal me¬

tabolic rates of medical students and nurses in training at

Charleston, South Carolina. Of the 93 student nurses, 48 were

within our age-range, and thus afforded another large homo¬

geneous group for comparison. The technique used was simi¬

lar in principle to ours, but only one standard, the Aub-DuBois,

was used for prediction. The study was evidently a result of

Remington’s general interest in problems connected with differ¬

ences in iodine distribution. He felt that since most of the

metabolic studies made so far have been in regions where there

is iodine deficiency, and goitre is common, it might add some¬

thing of interest to the subject to make such a study in a region

where neither of these conditions is found.

Remington and Culp noted the prevalence of low rates ac¬

cording to the DuBois standards, for both sexes, but having

noted that both lower and higher figures have been reported

from studies made in northern states, concluded that the low

values could not be regional. They suggest nutritional level as

a possible factor in trend of metabolic rates, though they feel

that this factor must be slight in groups of normal individuals.

When their subjects were grouped as to their deviations from

standard weight tables, the low-weight classes showed the low¬

est average metabolic rates. They did not consider the possible

bearing which the choice of standards might have on warranted

interpretations.

Supplementary Normals from the Nutrition Laboratory of

Boston. (6) We were interested to see how our data compared

with the more modern studies of normals from the Nutrition

Laboratory of Boston, but found just 5 subjects within range of

our study in Benedict’s 1928 series of normals (1). Benedict

made a definite effort in this later series to secure a wider range

of body-builds in his subjects than had been the case in the

original Nutrition Laboratory series, and three of these five

girls are heavy in proportion to their heights. As a separate

group the small series is therefore unbalanced; but for this

very reason, as well as its source, it offers some interesting

comparisons.

The Test Data. The measurements reported on all of these

subjects are reproduced herein. We have indicated with each

300 Wisconsin Academy of Sciences , Arts , and Letters .

table the recalculations we have made and the derived data that

have been added to afford consistent comparisons.

Tilt’s data are found in Table XII. Her results were pre¬

sented as averages of all the observations made on each sub¬

ject, representing from 2-8 consecutive or scattered tests in

each. All of the calculations of metabolic rates are our own,

since many of those given by Tilt failed to fit in with our meth¬

ods of calculation based on the reported body measurements.

The data on the 5 subjects from Benedict (Table XIV) also

represent, as is the practice at the Nutrition Laboratory in

seeking the most representative normal values, the average of

several basal periods, on from 1 to several days.

The data from Remington and Culp are found in Table XL

Their plan of study of making two or usually three determi¬

nations per subject on consecutive days allowed of the logical

averaging of the body measurements taken on the different oc¬

casions, while the results of each metabolism test are given

separately. Hence this series is more strictly comparable with

our own in one respect while the wide difference in locality and

also the fact that the subjects are student nurses rather than

college students, as are almost all the rest, lends breadth to

the discussion.

Tests of Fitness of Prediction Standards: DuBois (2) re¬

marks that selection can operate so as to make a group prove

any point with regard to trends in metabolic rate ! Difference

in such factors as repose, physical condition, manner of living,

and experience with the test on the part of the subjects investi¬

gated can combine in different ways to determine real or ap¬

parent group differences. The investigator adds his share of

variabilities in the matter of what constitutes “normality” and

a suitable standard of comparison for any group in question,

which will naturally depend to some extent upon the aims of

the investigation.

Preconceived criteria and prejudicial selection have been

avoided as far as possible in the present study. The aim has

been to make possible a practical choice primarily for the so¬

lution of a definite problem of our own in clinical metabolic

work. This problem is not confined to us. We have undoubt¬

edly had exceptional opportunities, however, in the way of

sources of material and cooperative effort in the search for a

Stark— Standards for Predicting Basal Metabolism. 301

solution. We believe that the nature of our material, of the

criteria we have set ourselves, and the results of the extensive

comparisons we have been, able to make with our own and a

significant number of data from widely separate sources are

such as would make our findings of interest to others who may

be concerned with the availability of adequate normal data for

physiological or clinical comparisons in this small but important

age and sex-group for whom the question of normal standards

has been baffiingly unsettled.

BASAL METABOLIC RATES

OF 97 NORMAL GIRLS 17 THRU 20 YEARS AT TNL UNIVERSITY OF

WISCONSIN CALCULATED BY DIFFERENT ’ NORMAL ’ PREDICTIONS

Arrows indicate Averaged

Fig. l.

The criterion of fitness of a standard that is most universally

applied is of course the average of the metabolic rates figured

for a given group according to that standard. This is instruc¬

tive, but it is only a part of the information which we need to

be able to judge the comparative merits of different predictions

on which to base standards of individual normality. The clini¬

cian is not interested in the average metabolic rate of a group

302 Wisconsin Academy of Sciences , Arts, and Letters .

of individuals, nor does it help him materially if, out of the

borderline cases with the same general type of history in whom

he is anxious to rule out hyper- or hypo-thyroidism, one is re¬

ported to show a metabolic rate of +20% and the other of

— 20%, thus yielding a perfect average fit with a “nor mar’

standard of reference.

Therefore we have compared not only average, but individual

ranges of showings in various salient respects of our different

groups and standards. The results of the different comparisons

have been summarized in the accompanying tables and graphs.

BASAL METABOLIC RATES

OF LATER TESTS ON 60 OF THE WISCONSIN STANDARD

GROUP OF SUBJECTS

Arrow indicate averages

Fig. 2.

The Comparisons: For general comparisons among our six

groups we have calculated average body measurements, aver¬

age determined heat-production as calories per 24 hours and

as calories per square meter of body surface per hour; and

average metabolic rates according to the four prediction stand¬

ards which we are comparing. These figures are collected in

Stark — Standards for Predicting Basal Metabolism . 303

BA5AL METABOLIC RATES

Or lid NORMAL GIRLS 17 THRU £0 YEARS FROM WISCONSIN, FLORIDA,

SOUTH CAROLINA AND BOSTON CALCULATED BY DIFFERENT ’NORMAL'

PREDICTIONS. DATA NOT INCLUDED IN THE WISCONSIN STANDARD

| Remington and (Mp 193/ § Tilt 1930 £j University of Wixmin [Not included in drama Oroop) Q Benedict @23

Table XV. The actual ranges and scatter of individual rates

by the four standards are charted in Figures 1, 2 and 3, which

bring out the general similarity of the showings by the four

304 Wisconsin Academy of Sciences , Arts , and Letters .

predictions when applied separately to the two groups of con¬

trols represented by our own standard subjects and the other

series combined. Separate charts have been made for first and

repeated tests for our 97 subjects (Figs. 1 and 2) for the em¬

phasizing of their essential similarities, with the slight average

tendency that would be expected toward lower values in re¬

peated tests. A single chart (Fig. 3) was made to give average

determined rates throughout for the 119 individuals of the

combined comparison series, since only part of these allowed

of the separation of first from later tests. The general simi¬

larity of the three pictures is striking. They make it obvious

at a glance that both the Aub-DuBois and the Harris-Benedict

standards are too high, while the Benedict prediction is almost

equally too low, to describe with any degree of accuracy either

these groups of normal girls examined in the north, or those

in the south.

In the first two cases the suggestion immediately offers it¬

self that it should be a simple matter to adapt either predic¬

tion, as it stands, by merely subtracting a uniform percentage

from the original constants, as Krogh suggested for the Aub-

DuBois tables (see page 291). In the case of the Benedict pre¬

diction, however, the excessive scatter leaves no room for hope

from the mere expedient of centering.

Leaving out of consideration the fact that the Benedict stand¬

ard gives values that are too low to represent the waking me¬

tabolism of the average girl, even under well controlled basal

conditions, it proves inferior to the other three by every stand¬

ard of comparison which we have been able to apply. The rea¬

son for this is undoubtedly the group method of study on which

it is based, which smoothed out individual variabilities in body

measurements and left only average relationships to be defined.

The metabolism was expressed as a simple function of the

weight (obviously the most important single variable) with

slight changes for age, leaving height entirely out of consid¬

eration. When we try to use this prediction for individuals

whose weight varies out of the average proportion to their

height, the results are bizarre, as our further actual compari¬

sons will bring out. Meanwhile it is interesting to note in

Table XV that the only one of the six series for which the Bene¬

dict prediction offers the best average “fit’” is Benedict’s own

small group of normals ! The averages by the other predictions

Table XV. Average Measurements and Calculated Metabolic Rates in Different Groups of Normal controls. All tests made on each sub¬

ject are included :

Stark— -Standards for Predicting Basal Metabolism . 305

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*Each test reported = average of several observations.

306 Wisconsin Academy of Sciences , Arts , and Letters .

do not indicate any characteristic difference in rate of heat pro¬

duction for this group compared with the others ; but these girls

happen to be predominantly overweight.

Table XV shows a remarkably close agreement between the

six groups when heat-production is calculated as calories per

square meter of body surface — the most logical single unit

of reference we know of for comparing levels of heat-produc¬

tion among different individuals or groups. There is no in¬

dication of a lower metabolic level predominating in the south¬

ern girls. Rather, the highest average figure is noted for the

large series reported by Remington and Culp from South Caro¬

lina. While the different type of life led by the student nurse

in comparison with that of college students may have some¬

thing to do with this finding, the difference is not large enough

to suggest this as a definite conclusion.

For the more exact comparisons summarized in the last two

tables we have used from the total data only those series which

allow the separation of first from later tests on individuals.

This has left us for logical comparison with our own stand¬

ard, which was based on first tests only, a group of 85 sub¬

jects consisting of the 48 by Remington and Culp and the 37

embraced by the three Wisconsin Supplementary series.

The following comparisons have been made within these two

groupings for the four predictions being tested: (Table XVI) :

Column 1: Algebraic averages of individual rates.

Column 2: “Standard Errors” of estimating rates by the

different predictions, i. e., the root-mean-square measure of dis¬

persal of the individually calculated rates, thus analagous to

the standard deviation method of evaluating dispersal among

the original measurements.

Column 3: Percentiles falling within the range convention¬

ally specified as strictly “normal” (±10%) in clinical metabolic

studies with adults.

Column 4: Percentiles falling within the range customarily

accepted as normal for practical purposes.

Column 5: Extreme ranges of rates outside plus or minus

10%.

Special Comparisons of Fit of Standards for Individuals of

Unusual Body Build. A traditional argument of those who

prefer the DuBois prediction to the Harris-Benedict has been

that while either standard may be relied upon to afford a fairly

Stark — Standards for Predicting Basal Metabolism . 307

good fit for individuals of average dimensions, the DuBois pre¬

diction alone, because it is based on the universally applicable

surface-area concept, is apt to prove suitable for reference in

the case of subjects of unusual bodily configuration. For this

a priori opinion the classical dictum of Krogh (20) may have

been not a little responsible. This rationalizing of a preference

is not at all the reasoning of DuBois himself, who believes that

some form of standard which relates the heat-production to

surface area is the most desirable because of the general biolo¬

gical significance of the comparisons which this permits, even

at the expense of some degree of accuracy in predicting for

individuals, as is inevitable when in practice we estimate the

surface-area from the simple measurements of height and

weight.

Among those who lean toward the statistical method of pre¬

dicting directly from the body-measurements without introduc¬

ing the intermediate estimation of the surface-area some have

ascribed superiority in predicting for people of unusual build

to the Harris-Benedict, and some to the formulas of Dreyer5.

Benedict himself wrote in 1928 (1) that the latter point of

view was gaining ground. He did not feel that this was borne

out however in his group of 27 men who had been chosen defi¬

nitely to include wide differences in both age and configuration.

For this group the average deviation of measured from pre¬

dicted heat-production was least by the Harris-Benedict stand¬

ard. Large discrepancies for certain individual rates accord¬

ing to the three predictions that were compared led him to

conclude that though the prediction for even a small group may

be made with surprising accuracy, the prediction of the basal

metabolism of individuals of unusual configuration is very

uncertain. It is clear that different criteria of suitability have

been applied by the different observers who have formed these

diverging opinions.

We were interested in finding as a final test of the various

standards of our study, which would prove by actual compar¬

isons to afford the best all-round “fit” for the least typical indi¬

viduals of the various groups of controls at hand. We decided

5 Actual comparisons based on the Dreyer prediction, which is no doubt too little known

in America, would have been interesting in the present study. Regrettably it was not con¬

sidered, after noting that its figures are too high for these girls. Ref. : Stoner, W. H.,

(Tabulation of Dreyer formulas) : Dost. Med. & Surg. J., 1923, clxxxix, 239.

308 Wisconsin Academy of Sciences , Arts , and Letters,

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Stark-— Standards for Predicting Basal Metabolism . 809

upon what constituted atypical build among essentially normal

subjects by the graphic method, as follows, in preference to

judging what should be considered over- or under-weight by

standards worked out on other groups :

Using the data of our original standard series as basis of

reference, we plotted heights against weights of the 97 sub¬

jects. Through this map we then drew the regression line of

weight on height, calculated from the statistical constants at

310 Wisconsin Academy of Sciences , Arts, and Letters.

hand for this group, and drew other lines on either side of this

average at distances equal to the standard deviation for weight.

This set apart 17 subjects outside (or just at) these limits as

being significantly below or above the group averages in weight

for height, and these subjects were used as the special group for

comparisons from our series. By fitting the data from the other

series to the curve, 21 individuals of unusual builds were picked

out in the same way. The height-weight chart is reproduced as

Figure 4, for the graphic representation of the distribution of

measurements that obtained in our standard group of subjects.

The list of those of unusual build picked out as described in¬

clude the following:

From the Wisconsin Standard Series (Data pp. 264-267) : Over¬

weight subjects b, d, f, k, y, age 17; subjects 14, 20, 22, age 18; and

subjects III and XXV, age 20. Underweight subjects i and x, age 17;

subjects 9, 24 and 25, age 18; and subjects 19 and 21, age 19.

From the series by Remington and Culp (Data pp. 293-295) : Over¬

weight subjects 150, 245, 250, 259, 278 and 888. Underweight subjects

158, 213, 239 and 395.

From Benedict’s series (Data p. 298) : Overweight subjects 132,

135 and 136.

From the series by Tilt (Data p. 296) : Overweight subjects 3 and 4.

From the Wisconsin total supplementary series (Data p. 297) :

Overweight subjects 1 and 7 by Shipley; and 7 and 12 by Fish. Under¬

weight subjects 3 and 20 by Fish.

To check our selections by a more conventional standard, we

calculated the degree of over- or underweight of each of the

selected subjects according to Bardeen’s height-weight tables

(21) which are our standards for clinical reference. Measure¬

ments taken at different times on the same subject were aver¬

aged for these comparisons. It was interesting to us to find how

closely similar were the weights predicted by Bardeen and those

predicted from our own regression equation for the individuals

throughout both series. The two predictions were within 1 or 2

pounds in over half of the cases, and in only one case was the

difference over 5 pounds. We cite this as evidence that our group

was anthropometrically representative.

Of the subjects picked out as unusual, 25 were overweight

for their heights by from 15 to 47 pounds according to our

standards, and from 12 to 40 pounds by Bardeen. The other

13 were underweight by 12 to 34 pounds by our formula, and

by 13 to 35 pounds by Bardeen. The average showings of

Stark— Standards for Predicting Basal Metabolism . 311

these four groups as to measurements and metabolic rates are

given in Tables XVII and XVIII. The groups were not large

enough to justify the calculation of any statistical constants,

but are sufficiently large for some illuminating general com¬

parisons. Average rates have been used for the individuals of

these series since the groups of Benedict and Tilt were brought

in for the comparisons.

A natural question to ask ourselves in making comparisons

like this is whether these girls who are so much larger or

smaller than the great majority of their group, though pre¬

sumably in good health, should be expected to show essentially

“normal” metabolic rates by the same standards used to judge

their fellows. This question cannot of course be answered in

our present state of ignorance about the factors which deter¬

mine body build. It might help us to gain some impression

about the essential normality of this particular atypical group,

however, if we pick out some of the most extreme examples and

compare their measured rates of metabolism in calories per

square meter of surface area with the average for the group at

large.

Thus on Figure 4, the four most strikingly aberrant points

are found to correspond to subjects III, age 20, and f, age 17,

who are both exceptionally tall and exceptionally heavy ; subject

20, age 18, who is the heaviest of the short subjects; and sub¬

ject x, age 17, who is very thin for her height. In addition we

might look at the two tiniest of all the subjects, albeit the pro¬

portion of their measurements does not bring them outside the

standard lines. Consulting the data tables for the Wisconsin

series we find the following relationships :

32.42

Average _ _ _ _ _ _ _ _ _ _ _ _ 34.07

312 Wisconsin Academy of Sciences , Arts, and Letters .

While they do not rule out the possibility that some of our

atypical girls may not have been strictly normal, the figures in

the last column certainly do not give us any reason for feeling

that accurate prediction for these girls should not be a reason¬

able requirement for a good working standard.

If we calculate the rates, then, of the underweight and over¬

weight subjects of our two large series according to the 4 pre¬

diction standards in their present forms, we see in Table XVIII

that the best average prediction is afforded in all 4 cases by

the University of Wisconsin standard, with Benedict’s predic¬

tion for girls, which fell so far behind for the groups as a

whole, almost as good for the overweight subjects. For this

group the two standards are about equally satisfactory also in

the matter of extreme range of rates. — But when we come

to the underweight subjects, the Benedict prediction is start¬

lingly aberrant by both criteria. The fact that the two types

of subject show a rather close agreement by all the other

criteria applied is against a reasonable suspicion that intrinsic

differences in metabolic levels are involved. It seems surpris¬

ing that the average calories per square meter are so similar in

all four groups.

There is no suggestion so far of the superiority of the Aub-

DuBois prediction in describing these groups of subjects who

are at the extreme ranges of body measurements for the popu¬

lations from which they were selected.

Table XVII. Average measurements of underweight and overweight individuals

taken from various series of normal controls. The figures given include all the tests on

each individual.

Stark— Standards for Predicting Basal Metabolism . 313

Table XVIII Comparisons of “fit” of different “normal” predictions of metabolic

rates for overweight and, underweight individuals from various series of normal controls.

The figures include all tests made on each individual.

♦Overweight subjects: U. W. total Supplementary Series: 4; Tilt: 2; Remington & Culp: 6;

Benedict ’28: 3.

Underweight subjects: U. W. total Supplementary Series: 2; Remington & Culp: 4.

But really legitimate comparisons are not possible between

standards that are “off center” for any given group to such dif¬

ferent degrees as are the three original standards which we are

trying to compare, and hence has undoubtedly arisen much dis¬

agreement in judgment. Each of these predictions as they

stand will score for rates in different ranges, but they will not

be the same ranges over any considerable series of trials. Hence

the evidence can be prejudiced in either direction, depending on

what one may be looking for. For example, with all of the ob¬

vious shortcomings to disqualify the Benedict standard for girls

for average usefulness, it has just been proven to afford an ex¬

cellent “fit” for sufficiently overweight girls !

The Centering of the Original Prediction Standards for Le¬

gitimate Comparisons. Hence for fairness and our own satis¬

faction, we have applied such constant percentage corrections

to the Harris-Benedict, the Aub-DuBois, and also the Benedict

predictions as would center them for our standard series, and

then have repeated our previous comparisons on the now com¬

parable bases for judgment. These comparisons based on the

314 Wisconsin Academy of Sciences , Arts , and Letters.

revised predictions, together with the original showings of the

University of Wisconsin prediction, have been collected in

Tables XIX and XX.

Observations on the Final Comparisons. If the Harris-Bene-

dict predictions are lowered by 7%, and the Aub-DuBois by

12 %6, either one affords a satisfactory fit for the two series of

normal controls that have been studied in the search of suitable

normal standards for girls under 21.

The constants in Table XIX show that by all four criteria

applied, — average prediction of rates, standard error of esti¬

mating the rates, percentiles within “normal” limits, and ex¬

treme ranges of rates, — the fit afforded both for groups and

individuals is somewhat better when the Harris-Benedict pre¬

diction is used than is the case with the Aub-DuBois.

The standard developed at the University of Wisconsin spe¬

cifically for girls between 17 and 21 affords a slightly better

fit than either of the others. This is supported not only by the

comparisons on the series of 97 girls upon whom the Wiscon¬

sin standard itself was based — which would be expected —

but the same order holds throughout similar comparisons in the

composite series of 85 normal controls of the same ages from

other sources. This with the various general comparisons

which we have been able to make leads us to believe that our

material can have more than local significance.

A small series of special tests to try to determine objectively

which of the various standards would most closely predict the

metabolism for individuals presumably normal but of unusual

bodily proportons showed for the 38 over- and underweight

girls of the above two and two other series of normal controls,

no consistent superiority of the Aub-DuBois, the Harris-Bene¬

dict or the University of Wisconsin standard. The total spread

of individual rates was somewhat smallest by the Wisconsin

standard.

The Benedict prediction for girls from 12 to 20 is improved

by centering only to the extent that having offered a passibly

8 This necessary correction should be nearer 10% according to the observation

of Mrs. Bradfield (Amer. J. Physiol., 1927, LXXXII, 570) that the DuBois

Height- Weight formula for estimating the surface-area gives figures that aver¬

age about 2% too high when applied to women. This finding does not of course

affect any of the comparisons between the different standards which we consider

fundamental, — i. e., those which are made after all the predictions are centered

for the same body of data.

Table XIX. Comparisons of “fit” of different “normal” predictions of metabolic rates when older standards are centered to fit average

University of Wisconsin data. Only the first tests on each individual are included in the figures.

Stark — Standards for Predicting Basal Metabolism . 315

*By adding a constant correction of — 7% to center for the University of Wisconsin Standard Series.

fBy adding a constant correction of — 12% to center for the University of Wisconsin Standard Series (see footnote, p. 000).

}By adding a constant correction of +8% to center for the University of Wisconsin Standard Series.

§See Table XVI, footnote.

316 Wisconsin Academy of Sciences, Arts, and Letters ,

Table XX. Comparisons of “fit” of different “ normal ” predictions of metabolic

rates for overweight and underweight individuals from various series of normal controls ,

when older standards are centered to fit average University of Wisconsin data. All the

tests made on each individual are included in the figures.

♦Corrections given in Table XIX.

fSee Table XVIII, footnote.

good fit before for overweight girls, and an extremely bad one

for the underweight, the latter disadvantage is now about equal¬

ly divided among both atypical groups. There is no need to

consider any longer the Benedict standard for girls in the pres¬

ence of three other predictions that are actually, or by simply

translating their zero-points, superior to it in every respect.

The older standards have the benefit of an age-factor, which

as yet we cannot define for our group, but we hope eventually

to do so to its further improvement after our observations shall

have been extended to cover a wider range of ages.

Meanwhile since there is slightly better prediction by our

present simple table embodying the equation h = 10.63 w +

3.23 s + 184.61 (Table X) than is possible by modifying any

of the pre-existing standards — a procedure preferably to be

avoided until it can be done by general agreement, — we pro¬

pose to use that for clinical prediction in girls from 17 to 20,

inclusive.

Stark — Standards for Predicting Basal Metabolism . 317

Summary

On the basis of practical experience that the standards so

far available for predicting the basal metabolism of girls under

21 are contradictory and in no wise satisfactory in clinical ap¬

plication, a possible solution of the problem was sought by the

collection of a statistically significant number of data on nor¬

mal girls between 17 and 21. This is an age and sex group of

particular importance in an institution which handles student

health problems.

Basal metabolic rates were determined in 163 tests on 97

girls, of ages quite uniformly divided between the 4 years 17,

18, 19 and 20. These were students at the University of Wiscon¬

sin who were judged to be Grade A in their medical and physi¬

cal examinations and who were determined by special examina¬

tion and questioning at the time of the tests to be free from

defects that should disqualify them from serving as physiologi¬

cal normals.

The tests were performed by the author between 7 :30 and

8:30 a. m. in the metabolism laboratory of the Wisconsin Gen¬

eral Hospital, under the same general conditions of technique

and control used for hospital cases. All tests were accepted for

analysis unless specific grounds for their rejection were appar¬

ent in technical flaw, lack of cooperation, or state of health of

the subject at the time of the test. The aim was in this way to

establish reasonable ranges of variation rather than strictly

minimum or average ideal levels of “basal” metabolism,

The results of the tests were subjected to various compara¬

tive analyses, and the first accepted tests on the 97 subjects

were made the basis of a prediction standard which was based

on the equation :

Heat Production in Calories per 24 hours = 10.63 X Weight

in Kilograms + 3.23 X Height in Centimeters + 184.61.

The equation was arrived at by standard methods of multiple

correlation. A factor for age, not yet possible to include be¬

cause of the narrow range of years observed, we shall hope to

add in time when our data shall have been extended to younger

and perhaps older subjects.

The above prediction standard was used to calculate the me¬

tabolic “rates” of the individuals in the original series, both

first and later tests ; and the rates for the same tests were also

318 Wisconsin Academy of Sciences , Arts , and Letters .

calculated by the Aub-DuBois prediction, the Harris-Benedict

prediction for adult women, extrapolated for these ages, and

Benedict’s special prediction for girls from 12 to 20. The same

calculations were carried out for a comparable test-series of

presumably normal girls of the same ages from Wisconsin

(other than standard series), Florida, South Carolina and Bos¬

ton, whose metabolic measurements have been reported by vari¬

ous recent observers.

Distribution charts of the rates in the same tests as indicat¬

ed by the 4 different predictions showed very similar pictures

for the two large series, and demonstrated in a striking way

that the Aub-DuBois and the Harris-Benedict predictions give

figures which are markedly too high, and the Benedict predic¬

tion figures which are too low, to represent with any degree of

satisfaction the “basal” or standard metabolism of large groups

of normal girls of these ages. The Benedict prediction showed

itself inferior to the other three by excessive scatter of rates

about the average expectation. It proved to indicate “normal”

rates for overweght girls, but abnormally high ones for the un¬

derweight.

The older predictions were then centered by applying the con¬

stant percentage correction to each which would make its zero-

point correspond practically to that of the Wisconsin standard

series by their own prediction. This was done, not as a perma¬

nent suggestion, but as the only basis possible to permit fair

comparisons in respects more fundamental than mere success in

predicting the average absolute level of heat-production for

groups of subjects. Various criteria were then applied to deter¬

mine which of the standards, with predictions now in the same

range, would afford the best all-around “fit” for these two con¬

siderable groups of normal subjects.

The criteria of suitability that were applied included : aver¬

age rates, standard error of predicting the rates, percentiles

within plus or minus 15% of the average expectation, and ex¬

treme ranges of rates by the different predictions.

Conclusions

By all the above criteria the Wisconsin standard proved to

furnish a somewhat better fit for both Wisconsin and other

normals than did either of the two most satisfactory of the old-

Stark— -Standards for Predicting Basal Metabolism. 319

er standards, that of Harris and Benedict, or that of Aub and

DuBois, even after centering to make their zero-points coincide.

However, of the two latter, the Harris-Benedict, when extrap¬

olated for the ages in question and lowered by 7%, showed itself

nearly as satisfactory in the various respects as did the Wiscon¬

sin Standard which was established by similar methods of an¬

alysis for a comparatively large number of subjects within a

narrow range of ages.

A special series of comparisons which determined average

and extreme ranges of rates for the 38 individuals from the

total series of controls who were from 12 to 40 pounds over- or

underweight for their heights according to the standard Bar¬

deen height-weight tables, showed practically the same degree

of satisfaction from any of the three standards after the two

older ones were centered to fit the average Wisconsin data. The

Wisconsin standard showed a slight advantage in smaller total

spread of individual rates. General comparisons indicated that

these subjects could be expected to show essentially “normal”

rates.

The various studies of the Wisconsin group and of the meas¬

urements made upon it, in comparison with those of a large

series of normal girls of the same ages from widely different

localities, suggest that the Wisconsin prediction may safely be

used as a standard of reference for similar classes of girls who

are studied with comparable technique anywhere in America.

References

(1) BENEDICT, F. G., Amer. J. Physiol ., 1928, lxxxv, 607. Basal Me¬

tabolism Data on Normal Men and Women (Series II) with Some

Considerations on the Use of Prediction Standards.

(2) DuBOIS, E. F., J. Nutr., 1930, iii, 217. Recent Advances in the

Study of Basal Metabolism. Part II: Basal Metabolism and Sur¬

face Area, ibid, p. 331.

(3) AUB, J. C., and DuBOIS, E. F., Arch . Int. Med., 1917, xix, 823.

Note at end of Clinical Calorimetry, 19th Paper: The Basal Me¬

tabolism of Old Men.

(4) BENEDICT, F. G., Proc. Amer. Phil. Soc., 1924, Ixiii, No. 1, 25.

Physical Factors in Predicting Basal Metabolism of Girls.

(5) BENEDICT, F. G., and HENDRY, Mary F., Bost. Med. and Surg.

J., 1921, clxxxiv, 217, 257, 282, 297 and 329. The Energy Require¬

ments of Girls from 12 to 17 Years of Age.

320 Wisconsin Academy of Sciences , Arts, and Letters.

(6) McCLENDON, F. J., Physiol. Rev., 1927, vii, 189. Distribution of

Iodine with Special Reference to Goiter.

(7) DuBOIS, E. F., Lea and Febiger, 2d ed., 1927. Basal Metabolism

in Health and Disease.

(8) GRIFFITH, Fred R., Jr., PUCHER, Geo. W., BROWNELL, Kath¬

erine, KLEIN, Jennie D., and CARMER, Mabel E., Amer. J.

Physiol., 1928-9, Ixxxvii, 602. Studies in Human Physiology. I.

The Metabolism and Body Temperature (Oral) under Basal Con¬

ditions.

(9) MacLEOD, Grace, CROFTS, Eliz. E., and BENEDICT, F. G., Amer.

J. Physiol., 1925, lxxiii, 449. The Basal Metabolism of Some

Orientals.

(10) HARRIS, J. A., and BENEDICT, F. G., Carnegie Inst, of Wash.

Publ. #279, 1919. A Biometric Study of Basal Metabolism in

Man.

(11) DuBOIS, E. F., Arch. Int. Med., 1916, xvii, 877. Clinical Calorimet¬

ry, 12th Paper: The Metabolism of Boys 12 and 13 Years Old

Compared with the Metabolism at Other Ages.

(12) DuBOIS, E. F., Amer. J. Med. Sci., 1916, cli, 781. The Respiration

Calorimeter in Clinical Medicine.

(13) Krogh’s Recording Respiration Apparatus, Tables of Normal Me¬

tabolic Rates after DuBois and Benedict-Harris, J. H. Schultz,

Copehagen, 1925. (Quoted from DuBois, Ref. 7 above, p. 174-5.)

(14) BOOTHBY, W. M., and SANDIFORD, Irene., Amer. J. Physiol,

1929, xc, 290. Normal Values of Basal or Standard Metabolism.

A Modification of the DuBois Standards.

(15) Sanborn, F. B. (Editor) : Basal Metabolism. Its Determination

and Application. Boston, 1922, p. 238. (Quoted from DuBois,

Ref. 7 above, p. 179.)

(16) STONER, Boston Med. and Surg. J., 1924, cxci, 1026, 1030.

(17) KESTNER, Otto, and KNIPPING, H. W., Julius Springer, Berlin,

2d ed., 1926. Die Ernahrung des Menschen.

(18) TILT, Jennie, J. Biol. Chem., 1930, lxxxvi, 635. The Basal Me¬

tabolism of Young College Women in Florida.

(19) REMINGTON, R. E., and CULP, F. B., Arch Int. Med., 1931, xlvii,

366. Basal Metabolic Rate of Medical Students and Nurses in

Training at Charleston, S. C.

(20) KROGH, August, Bost. Med. and Surg. J., 1923, clxxxix, 313. De¬

termination of Standard (Basal) Metabolism of Patients by a Re¬

cording Apparatus.

(21) BARDEEN, C. R., Carnegie Inst. Wash. Publ. #272, 1920.

THE DECAPOD CRUSTACEANS OF WISCONSIN

Edwin P. Creaser

Previous lists of the decapod crustacean fauna of Wisconsin

have been published by Bundy (1882), Faxon (1885), Harris

(1908), and Graenicher (1913). The first list compiled by

Bundy contained eleven species. Many of these are synonyms.

Graenicher’s list contains seven species of crayfish, including

C. rusticus of which no specimens from Wisconsin were exam¬

ined. Six species of crayfish are now definitely known to oc¬

cur within the state, and in addition the shrimp, Palaemonetes

exilipes.

The writer has examined the extensive collections obtained

through the efforts of the Wisconsin Geological and Natural

History Survey. For two summers the writer collected for the

Survey in Wisconsin. In addition to these collections, those

belonging to the Zoology Department of the University of Wis¬

consin have been examined, and also a few of the specimens

in the collections of the Public Museum of Milwaukee. Dr.

Waldo L. Schmitt of the United States National Museum has

furnished several records of Wisconsin crayfish. Miss Myrtle

Creaser of Kenosha has very kindly furnished many specimens

from that region. Decapod crustaceans have been examined

from nearly every section of Wisconsin. The following coun¬

ties are not represented: Door, Kewaunee, Brown, Oconto,

Calumet, Manitowoc and Sheboygan. Most of these counties

are bordered by either Lake Michigan or Green Bay.

Most of the collecting has been done with seines. This meth¬

od does not allow a fair sampling for distributional studies of

the borrowing species, Cambarus diogenes and Cambarus gra¬

cilis. Additional collections are especially needed of these two

species.

Family Astacidae

Two families of decapods occur in Wisconsin. Most of these,

the crayfishes, are classed in the Astacidae, a family of fresh¬

water custaceans found in Asia, Europe, North and Central

America. Two genera, as now defined, occur in the United

322 Wisconsin Academy of Sciences , Arts , and Letters.

Fig. 1. Diagrams of Cambarus virilis. ABD., abdomen; AN., anten-

nule; ANT., antennae; ARE., areola; A. S., antennal scale; B., basipo-

dite; C., carpopodite (carpus) ; CAR., carapace; C. G., cephalic groove;

CO., coxopodite; D., dactylopodite ; EYE , eye; HO., hook; I., ischiopo-

dite (ischium); I. UR., inner ramus of uropod; L. S., lateral spine; M.,

meropodite (merus) ; MAX., third maxilliped; O. UR., outer ramus of

uropod; P., propodite (propodus) ; P. R., postorbital ridge; RO., rostrum;

S. O., sexual organ; SJF., swimmeret; TEL., telson; 1, 2, 3, U, 5, walking

legs.

States; the genus Astacus in California, Washington, Oregon,

Idaho and Wyoming; and the genus Cambarus in the states

which drain into the Atlantic Ocean. In Mexico the crayfishes

of the genus Cambarus are found in the Pacific as well as in the

Atlantic drainage areas. A species of Cambarus , (C. clarhii)

has apparently been introduced in California within recent

years.

The word crayfish is possibly derived from the French

ecrevisse, or the Low Dutch crevik. Names commonly applied

Greaser — Decapod Crustaceans of Wisconsin . 323

to these animals are: crayfish, crawfish, crawdads and inap¬

propriately, crabs*

The crayfish frequently serve as an intermediate host of par¬

asitic worms, of which Pamgonimus sp. is a typical example.

An ostracod, Entocythere cambaria , originally described from

Wisconsin, is parasitic on the gills of crayfish. Aquatic oli-

gochaetes (earthworms) of the family Discodrilidae are often

found as parasites or symbionts on crayfish.

Identification of the species of crayfish is based largely upon

the structure of the first abdominal appendages (sexual or¬

gans) of the male. Two forms of this appendage are known in

each species of the genus Cambarus . These are usually desig¬

nated as Form I and Form II. Males of the first form can be

immediately distinguished by the horny color of the tips of the

outer part of the sexual organ. Female specimens have an or¬

gan between the last pair of walking feet called either an annu¬

lus ventralis or receptaculum seminalis. Its shape is charac¬

teristic for any given species.

The key which follows is based largely on males of the first

form. The reader is cautioned at the outset that some varia¬

bility exists in the form of the male sexual organ. All of the

drawings, except those of C. gracilis and C. immunis, were

made from Wisconsin specimens. Figure 1 and the glossary at

the end of this paper explain the terminology used in the key.

Key to the Wisconsin Crayfishes

Genus Cambarus

la. Sexual organ of male terminating in two, slender, elongated, taper¬

ing tips, either straight or gently curved. Third walking legs of

male with hooks. Wisconsin species of subgenus Faxonius.

2a. Both tips of male sexual organ curved posteriorly.

3a. Antennal scale uniformly rounded. Chelae moderately broad,

moveable finger with straight cutting edge. Length of posterior

section of carapace contained less than twice in the length of the

anterior section. Male sexual organ gently curved. Female an¬

nulus ventralis with a median transverse fossa. (Fig. 2.)

Cambarus virilis Hagen 1870.

3b. Antennal scale irregularly rounded. Chelae narrow, moveable

finger with incision at base of cutting edge. Length of posterior

section of carapace contained twice in the length of the anterior

324 Wisconsin Academy of Sciences, Arts, and Letters .

I , antennal scale; 2 , annulus

ventralis ; 3 , sexual organ,

form I; sexual organ, form

II.

nis. 1 , antennal scale; 2, an¬

nulus ventralis; 3, sexual or¬

gan, form I; 4, sexual organ,

form II.

section. Male sexual organ abruptly curved. Female annulus ven¬

tralis with a deep fossa to the left.1 (Fig. 3.)

Cambarus immunis Hagen 1870.

2b. Tips of male sexual organ straight.

4a. Rostrum with a well developed median carina or elevation above.

Sexual organ stout with short tips; outer tip about the same length

as the inner. Annulus ventralis flat, fossa very small or absent,

sinus nearly straight. (Fig. 4.)

Cambarus propinquus Girard 1852.

lb. Sexual organ of male terminating with two short tips curved at about

right angles with the main shaft, the outer of the two tips (the top

one) flattened laterally, the inner one rounded. Hooks on the third

walking legs of the male. (See also item 1c. below). Wisconsin

species of the subgenus Cambarus .

5a. Areola obliterated. Carapace higher than wide. Anterior margin

of carapace with a triangular protuberance behind the antennal

scale. Rostrum without lateral spines.

Cephalothorax without lateral spines. Annulus ventralis with

sinus curved to the right2; fossa short, median and transverse.

(Fig. 5.)

Cambarus diogenes Girard 1852.

1 That is, to the observer’s left when the animal is seen in ventral view with

the anterior end up.

2 That Is, to the observer’s right when the animal is seen in ventral view with

the anterior end up.

Creaser— Decapod Crustaceans of Wisconsin .

325

Fig. 4. Cambarus propin-

quus. 1, antennal scale; 2,

annulus ventralis; 3 , sexual

organ, form I; 4, sexual or¬

gan, form II.

Fig. 6. Cambarus gracilis.

I , antennal scale; 2, annulus

ventralis; 3, sexual organ,

form I ; 4, sexual organ, form

II.

*

Fig. 5. Cambarus dio ge¬

nes. 1, antennal scale; 2,

annulus ventralis; 3, sexual

organ, form I; 4, sexual or¬

gan, form II.

Fig. 7. Cambarus bland-

ingii acutus. 1, antennal

scale; 2, annulus ventralis;

3, sexual organ, form I; U,

sexual organ, form II.

326 Wisconsin Academy of Sciences , Arts , and Letters .

lc. Sexual organ of male truncate or blunt at the tip; outer part ending

in three rather short teeth; inner part terminating in an acute spine.

Hooks on the third or on the third and fourth walking legs of the

male. Wisconsin species of the subgenus Ortmannicus.

6a. Areola obliterated. Margin of antennal scale evenly rounded. Hooks

on third walking legs of male. Chelae broad with moderate fing¬

ers. Inner tip of sexual organ exceeding teeth of outer part. Outer

part without row of setae at apex. Female annulus ventralis

nearly round, sinus irregular, fossa shallow. (Fig. 6.)

Cambarus gracilis Bundy 1876.

6b. Areola narrow but distinct. Margin of antennal scale irregular.

Hooks on third and fourth walking legs of male. Chelae long,

fingers slender, palm oval. Inner tip of sexual organ curved

obliquely outward, not exceeding teeth of outer part. Apex of

outer part with row of setae. Female annulus ventralis wide,

with two longitudinal tubercles. (Fig. 7.)

Cambarus blandingii acutus Girard 1852.

Distribution and Habits of Wisconsin Crayfish

1. Cambarus virilis Hagen

Range: Chihuahua, Texas, Colorado, Oklahoma, Kansas, Ar¬

kansas, Alabama (?), Missouri, Iowa, Minnesota, North Da¬

kota, South Dakota, Wisconsin, Michigan, Illinois, Indiana,

Ohio, Ontario, Saskatchewan and Manitoba.

Distribution in Wisconsin (Fig. 8) : Without question this

species is the commonest in the state. It occurs in all the major

drainage areas and probably will be found in every county.

Ecology: Cambarus virilis inhabits lakes, streams and riv¬

ers. It is found typically under stones, although I have often

taken specimens from muddy creeks and in aquatic vegetation.

Females with eggs are found during the early spring, usually

before the last of April. This species occurs even in cold trout

streams. In the region of Green Bay it is of considerable eco¬

nomic importance. At certain seasons great numbers are

caught and shipped to the Chicago market. This species is the

largest in the state and frequently attains a size of over eight

inches.

2. Cambarus immunis Hagen

Range: Colorado, Oklahoma, Kansas, Missouri, Nebraska,

Iowa, Wyoming, North Dakota, Minnesota, Wisconsin, Illinois,

Indiana, Ohio, Michigan, Ontario, New York and Massachu¬

setts.

Creaser— -Decapod Crustaceans of Wisconsin.

827

Distribution in Wisconsin (Fig. 9) : This crayfish is appar¬

ently very local, being confined, as far as we now know, to

the southeastern corner of the state and in the region of Lake

Pepin. It is not common in Wisconsin, but probably is more

abundant than indicated on the map, as it frequently occurs in

temporary ponds which have not been adequately examined for

crayfishes.

Ecology : This species is encountered in various types of

ecological situations. In rivers and streams it avoids strong

currents. It is frequently found in small lakes, especially those

with muddy bottoms. When disturbed, it often darts back-

328 Wisconsin Academy of Sciences , Arts, and Letters .

wards and downwards, striking the bottom so forcibly that the

specimen cannot be seen through the murky mud. Cambarus

immunis occurs sometimes in temporary ponds. As the season

. advances and the ponds become dry, it resorts to shallow bur¬

rows. Females with eggs are found from November to Febru¬

ary. During the winter months this crayfish remains in a dor¬

mant state, buried in the mud. Graenicher (1913, p. 122) gives

the following note regarding this species in Wisconsin : “Along

the Wisconsin side of Lake Pepin north of Maiden Rock in

Pierce Co., the water is extremely shallow, and in many places

the bottom is covered with a dark, sticky mud. In the summer

Creaser — Decapod Crustaceans of Wisconsin .

329

Fig. 10. Distribution of Cambarus propinquus in Wisconsin.

of 1910 the water in the Mississippi river was extremely low,

and males and females of C. immunis were found on August 3,

and 9, in burrows along the wet shore, quite a distance from

the lake.” In life this crayfish is greenish brown on the cara¬

pace and abdomen. On the dorsal surface of the abdomen a

design is formed by a contrasting lighter color.

3. Cambarus propinquus Girard

Range: Quebec, Ontario, Michigan, Wisconsin, Iowa, New

York, Pennsylvania, West Virginia, Ohio, Indiana, Illinois.

Distribution in Wisconsin (Fig. 10) : This species presents

a most unique distribution in the state. It occurs throughout

330 Wisconsin Academy of Sciences , Arts, and Letters .

the Lake Michigan drainage area to the east, and in the Mis¬

sissippi drainage northward as far as the Wisconsin river and

tributaries. It is not found over the western half of the state

above the Wisconsin river drainage area. To the writer it

seems quite apparent that this indicates a relatively recent in¬

vasion from the south and from the east.

Ecology : This species is typically a stream and lake form.

It has a habitat preference similar to C. virilis, being found

most frequently in clear streams and lakes with stony bottoms.

It occurs, however, in other habitats and is frequently taken in

dense mats of aquatic vegetation. This species never attains

Greaser— Decapod Crustaceans of Wisconsin.

331

the size of C. virilis and accordingly has an economic use only

as bait and as fish food. Females with eggs attached occur in

Michigan from April to June (Pearse, 1910, p. 16).

4. Comb ar us diogenes Girard

Range: Wyoming, Colorado, Kansas, Arkansas, Missouri,

Iowa, Minnesota, Wisconsin, Illinois, Indiana, Ohio, Michigan,

Ontario, New York, Pennsylvania, New Jersey, Maryland, West

Virginia, Virginia, North Carolina. The subspecies ludovicianus

occurs in Mississippi and Louisiana.

Distribution in Wisconsin (Fig. 11) : This burrowing cray¬

fish doubtless occurs throughout the entire state. The gaps in

the map are explainable as resulting from the fact that most

of the crayfish collecting in the state has been done with seines.

Ecology: Cambarus diogenes is a burrowing species which

resorts to streams, rivers and lakes only during the breeding

season in the early spring. It is a common sight to see hun¬

dreds of the burrows of this species along a water course. Fre¬

quently the tops are covered with mud “chimneys”. These bur¬

rowing crayfish are often forced to go down several feet before

they reach the water level. I have dug this species from bur¬

rows which extended more than three feet beneath the surface.

Sometimes many small crustaceans, amphipods, ostracods, and

copepods, are found in the water pocket of these burrows

(Creaser, 1931, pp. 243-244). Female specimens with eggs

were found in Indiana on April 17, 1930. Pearse (1910, p. 20)

records a female with young taken in Michigan in June.

5. Cambarus gracilis Bundy

Range: Oklahoma, Kansas, Missouri, Iowa, Illinois, Wiscon¬

sin, Indiana (?), Ohio (?).

Distribution in Wisconsin (Fig. 12) : As yet this species is

known only from Racine and Milwaukee Counties. It may oc¬

cur elsewhere in the prairie regions of southern Wisconsin,

where it may have escaped notice on account of its burrowing

habits.

Ecology: Cambarus gracilis is another burrowing crayfish.

Its burrows, unlike those of Cambarus diogenes are seldom

found along streams, rivers, or lakes but are most frequently

332 Wisconsin Academy of Sciences, Arts, and Letters,

Fig. 12. Distribution of Cambarus gracilis , Cambarus blandingii acutus

and Palaemonetes exilipes in Wisconsin.

found in the vicinity of small ponds. In Missouri I dug a speci¬

men from a burrow which extended more than six feet be¬

neath the surface before the water level was reached. Females

with young attached are taken in Missouri as late as October.

This crayfish can often be obtained after a rain storm, when in¬

dividuals leave their burrows.

6. Cambarus blandingii acutus Girard

Range: Michigan, Ohio, Indiana, Illinois, Wisconsin, Iowa,

Missouri, Arkansas, Kansas, Oklahoma, Tennessee, Mississippi

(?), Texas, Vera Cruz, N. Carolina (?), S. Carolina (?).

Greaser —Decapod Crustaceans of Wisconsin.

333

Distribution in Wisconsin (Fig. 12) : In the Mississippi

river drainage this species extends northward slightly further

than La Crosse, and in the Lake Michigan drainage as far

northward as Milwaukee.

Ecology : This crayfish frequently occurs in temporary ponds

where it builds shallow burrows, probably for mating. It is

also found along rivers, especially those with marshy banks.

It never occurs in rapidly flowing streams. Turner (1926, p.

154) records this species as carrying eggs or young during the

months of March, July, and September. This species probably

does not have a restricted breeding season.

Family Palaemonidae

First two pairs of legs chelate. Pleura of first abdominal

segment overlapped by those of the second. Abdomen with a

sharp bend. Rostrum armed with teeth, immoveable, long and

compressed. Mandibles deeply cleft. Gills developed as phyllo-

branchiae.

The crustacean family Palaemonidae, essentially a marine

group, is represented in the fresh-water fauna of the United

States by five species: (1) Macrobrachium jamaicensis from

Florida and Texas, (2) M. acanthurus from Florida, (3) M.

ohionis from the Mississippi and Ohio rivers, (4) Palaemonetes

antrorum , a blind species, from an artesian well at San Marcos,

Texas, (5) P . exilipes from the eastern half of the United

States.

Genus Palaemonetes

Both pairs of legs approximately the same size. Rostrum

with teeth above and usually below. Mandibles without a palp.

7. Palaemonetes exilipes Stimpson

Range: Iowa, Wisconsin, Michigan, Ontario, New York,

Pennsylvania, Ohio, Indiana, Illinois, Kentucky, Tennessee, Ar¬

kansas, Oklahoma, Texas, Nuevo Leon, Louisiana, Mississippi,

Alabama, Florida, Georgia, South Carolina, North Carolina,

Virginia, District of Columbia, Maryland.

Distribution in Wisconsin (Fig. 12) : This species is known

from four localities along the Mississippi river in Wisconsin.

334 Wisconsin Academy of Sciences , Arts , and Letters .

It is doubtless of general occurrence in and near the Missis¬

sippi river.

Diagnostic characters : Fifth walking leg exceeding rostrum.

Antennae longer than body. Antennules triflagellate, shortest

flagellum closely attached to longest and free but for a short

distance. Antennal scale reaching to a point even with apex

of rostrum. Rostrum long, vertical, armed above with 6-8

teeth, and below with 1-3. Second dorsal tooth of rostrum usu¬

ally above base of eye stalk. Carapace rounded above, branch-

iostegal and hepatic spine present. Abdomen abruptly curved

at end of third segment.

Ecology: Palaemonetes exilipes prefers slowly moving or

stagnant waters. It is frequently abundant in overflow ponds

where its enemies are few. These small crustaceans are eaten

by many species of fishes. They could be very easily raised in

large numbers and could be used as food for trout in hatch¬

eries. Palaemonetes exilipes can be kept for long periods in the

laboratory and they make admirable aquarium animals. Eggs

are found attached to the females in the spring and fall. This

indicates two breeding seasons a year, but adequate studies of

the life history of this interesting shrimp have not been made.

Greaser — Decapod Crustaceans of Wisconsin .

335

Species of Doubtful Occurrence or Validity

In this study no specimens of C. rusticus have been found.

This species has previously been recorded in Wisconsin from

Racine Co., Ironton, Sauk Co., and Beloit, Rock Co. It is my

belief that C. rusticus does not occur in Wisconsin, and that the

specimens previously referred to this species are, in reality,

aberrant forms of C. propinquus.

Bundy (Forbes, 1876, pp. 3-4) has described a new species of

crayfish (C. stygius) from Lake Michigan, near Racine. This

species has persisted in the literature as one of doubtful va¬

lidity. From Bundy’s later account (1882, p. 180) it is quite

apparent that he described immature, newly moulted speci¬

mens. The differences noted between this species and C. blan-

dingii acutus are so slight that it seems almost certain that C.

stygius is a synonym of C. b. acutus.

The following list gives the synonymy of the Wisconsin spe¬

cies mentioned by Bundy (1882, p. 180-183).

Species

1. C, blandingii acutus,

2. C. virilis,

3. C. propinquus,

4. C, gracilis,

5. C, dio genes,

6. C . immunis,

7.

Synonymy of Bundy

la. C, acutus,

lb. C, stygius,

2a. C, virilis,

2b. C. wisconsinensis.

2c. C. debilis,

3a. C, propinquus,

3b. C, placidus, (?)

3c. C. rusticus. (?)

4a. C. gracilis.

5a. C. obesus.

6a. _ _ _

7a. C. bartonii ,3

The Wisconsin species with the exception of C. gracilis, are

of general occurrence in adjacent states as shown in Table I.

C. gracilis is a southwestern form which has probably become

established in Wisconsin by an immigration up the Mississippi

drainage. The decapod fauna of Wisconsin and adjacent states

may be conveniently grouped as follows :

8 Cambarus bartonii has never been taken in Wisconsin.

336 Wisconsin Academy of Sciences, Arts, and Letters .

1. Species of general occurrence: C. diogenes, C. b. acutus,

C. immunis, C. propinquus, P. exilipes.

2. Southwestern prairie species: C. gracilis .

3. Species of northeastern drainage areas: C. robustus, C.

virilis.

4. Species of the Ohio valley: C. rusticus (A confusing ar¬

ray of subspecies of this crayfish exist elsewhere.)

5. Species confined to the Mississippi and Ohio rivers: M .

ohionis .

6. Blind cave species: C. pellucidus and C. pellucidus testii .

7. Local species: C . bartonii laevis, C . ortmanni, C. sloanii

and C. indianensis .

8. Species of southeastern occurrence : C. juvenilis .

Table I. Decapod occurrence in Wisconsin and adjacent states.

C. fodiens (C. argillicola) has been reported from Michigan,

Lower Ontario, Ohio, Indiana, Illinois, Mississippi, Louisiana,

Texas and North Carolina. The records for the four states last

named are surely doubtful. Consequently C. fodiens may be a

species of rather wide spread occurrence in the north central

states. This species is a burrower and is found in the spring¬

time in temporary woodland ponds. A search should be made

for this crayfish in southern Wisconsin. (It is a species of the

subgenus Cambarus, and is characterized by having the areola

Creaser — Decapod Crustaceans of Wisconsin .

387

obliterated and the moveable finger of the chela with a deep

incision at the base of the inner margin.)

Literature

Bundy, W. F. 1882. A List of the Crustacea of Wisconsin with Notes

on Some New or Little Known Species. Trans . Wis. Acad. Sci. 5 :

177-184.

Creaser, E. P. 1931a. Some Cohabitants of Burrowing Crayfish. Ecol¬

ogy 12 (1) : 243-244.

1931b. The Michigan Decapod Crustaceans. Pap. Mich. Acad. Sci. 13 :

257-276, 9 figs., 6 maps.

Faxon, Walter. 1884. Descriptions of New Species of Cambarus. Proc.

Am. Acad. Arts and Sci. 20 : 107-158.

1885. A Revision of the Astacidae. Mem. Mus. Comp. Zool. 10 (Part

4) : 1-186.

1890. Notes on North American Crayfishes. Proc. U. S. Nat. Mus.

12 : 619-634.

1898. Observations on the Astacidae in the United States National

Museum and in the Museum of Comparative Zoology .... Proc. U. S.

Nat. Mus. 20 : 643-694, 9 pis.

1914. Notes on the Crayfishes in the United States National Museum

and in the Museum of Comparative Zoology .... Mem. Mus. Comp.

Zool. 40 (8) : 351-427, 13 pis.

Forbes, S. A. 1876. List of Illinois Crustacea with Descriptions of New

Species. Bull. III. Mus. Nat. Hist. 1 : 3-25, 1 pi.

Graenicher, S. 1913. Some Notes on the Habits and Distribution of

Wisconsin Crawfishes. Bull. Wis. Nat. Hist. Soc. 10 (3-4) : 118-123.

Hagen, H. A. 1870. Monograph of the North American Astacidae. III.

Cat. Mus. Comp. Zool., Harvard College 3 : 1-110, 11 pis.

Harris, J. A. 1903. An Ecological Catalogue of the Crayfishes Belong¬

ing to the Genus Cambarus. Kan. TJniv. Sci. Bull. 2 (3) : 51-187.

Hay, W. P. 1896. The Crayfishes of the State of Indiana. Ann. Rept.

Ind. Geol. Survey , 20 : 475-507, 15 figs.

Huntsman, A. G. 1915. The Fresh-Water Malacostraca of Ontario.

Contr. Canad. Biol. 1911-1914 (Sessional Paper No. 39b) : 145-163.

Ortmann, A. E. 1905. The Mutual Affinities of the Species of the Genus

Cambarus, and Their Dispersal over the United States. Proc. Am.

Philosoph. Soc. 44 : 91-136, 1 map.

1931. IV. Crawfishes of the Southern Appalachians and the Cumber¬

land Plateau. Ann. Carnegie Museum 20 (2) : 61-160.

Pearse, A. S. 1910. The Crayfishes of Michigan. Mich. State Biol. Surv.,

Bull. 1 : 9-22, 8 pis.

Smith, S. I. 1874. The Crustacea of the Fresh Waters of the United

States. Rep. U. S. Comm. Fish. 1872-1873, 2 : 637-665.

Steele, Mary. 1902. The Crayfish of Missouri. Bull. No. 10 Univ. Cin¬

cinnati 2 (Ser. 2) : 1-54, 4 pis.

Turner, C. L. 1926. The Crayfishes of Ohio. Ohio State Univ. Bull. 30

(11) : 145-195, 2 pis.

338 Wisconsin Academy of Sciences , Arts, and Letters.

Glossary

Branchiostegal spine, a spine situated near the anterior margin of the

carapace, ventrad to the hepatic spine (in shrimp).

Cephalo thorax, the solid front part of the body covered by a continuous

chitinous shield. The cephalic groove divides this into an anterior

and a posterior section. Also called the carapace.

Fossa, a cavity, pit or depression.

Hepatic spine, a spine laterally situated on the carapace, behind and be¬

low the eye.

Obliterated areola, areola limited to posterior and anterior triangular

fields by the fusion of the limiting (branchio-cardiac) grooves.

Palp, the endopodite or inner ramus of a mouth part.

Phyllobranchiae, gills with two rows of broad flat lamellae.

Pleura, the side of an abdominal segment.

Sinus, an elongated groove.

Truncate, having the terminal edge square or even.

PRELIMINARY LIST OF THE HYDRACARINA OF

WISCONSIN

Part II

Ruth Marshall

Part I of the Preliminary List of the Hydracarina of Wis¬

consin (Marshall, 1931) recorded fourteen species belonging to

seven genera of the red water mites, the super-family Lim-

nocharae. The present paper treats in the same way a portion

of the much larger super-family Hygrobatae, namely, seventeen

species of nine genera belonging to four large families found

in the state. Of these, one species is new. Notes on distribu¬

tion and one or more drawings are given for each species. For

complete characterizations of the species the student is referred

to titles in the bibliography. A few outstanding features of

each are noted, however, which, together with the drawings

will, it is hoped, be sufficient in most cases to identify the spe¬

cies.

The water mites of this group are of medium size, variously

and often brightly colored, the skin usually thin but sometimes

heavily chitinized; the paired eyes of each side do not lie on

capsules; the palpi are not chelate; the epimera are extensive

and varied and more or less united; the legs usually end in

claws which are double toothed ; the genital plates are well de¬

veloped and often placed apart from the epimera and sexual

dimorphism is usually well marked.

For most of the extensive collections from Green Lake and

the Trout Lake, Vilas County, region and for all of the collec¬

tions from given depths the author is indebted to the courtesy

of the Wisconsin Natural History Survey.

Lebertia porosa Thor

PI. VII, fig. 1.

The epimera in the Lebertia are united into a shield which

partly encloses the genital area. This cosmopolitan species is

recognized by details of the palpi and epimeral plates and their

relation to the genital plates.

340 Wisconsin Academy of Sciences , Arts , and Letters .

The species is found throughout Europe and in northern

Asia; in Canada, Alaska, Colorado, Montana, Wyoming, Illi¬

nois, Iowa, Michigan, Indiana and New York. In Wisconsin

it has been found in Lake Winnebago, the Madison lakes, Drake

Lake (Waupaca), Goose Pond (Adams Co.), in four lakes of

Vilas County and in several collections from Green Lake at

depths from the surface to a few meters.

Lebertia quinquemaculosa Mar.

PI. VII, fig. 2-4.

Adults, which may attain a length of 2 mm., are usually rec¬

ognized when alive by the presence of five red spots on the

dorsal surface. The first epimera are narrow and the fourth

forms a wide bay for the genital plates. The nymphs have ven¬

tral plates characteristic for the genus.

Specimens have been found in British Columbia and Indiana ;

in Wisconsin they have been taken in Mirror Lake (Delton),

Deep and Parker lakes (Adams Co.) and in Green, Powers and

Twin Lakes, all from shallow water.

Oxus connatus Mar.

PI. VIII, fig. 9, 10.

In the genus Oxus the epimera are fused into a shield. In 0.

connatus this shield is not extensive and it only partly encloses

the genital area and the body is relatively low.

Specimens have been found in Ontario. In Wisconsin they

have been taken in Mirror Lake, Goose Pond (Adams Co.),

Lake Mendota, Green Lake and several of the lakes of Vilas

County.

Oxus elongatus Mar.

PI. VIII, fig. 13, 14.

The epimeral shield is somewhat more extensive than in the

last species and the body is higher.

The species has been found in Ontario and in Wisconsin in

several of the lakes of Vilas County, near the surface and at

depths of a few meters.

Oxus intermedins Mar.

PI. VIII, fig. 11, 12.

The epimeral shield is extensive, extending to the dorsal side,

and the body is elevated.

Marshall— Hydracarina of Wisconsin.

341

It has been taken in Minnesota, in the Madison lakes and

in several of the lakes of Vilas County together with specimens

of the last two species.

Frontipoda americana Mar.

PI. VIII, fig. 15-17.

This species, the only one of the genus so far reported in

North America, is common in shallow waters. It is green (oc¬

casionally red), with brown markings, and is readily recognized

by the laterally compressed body and the great development of

the epimeral shield which entirely encloses the genital area and

extends over most of the dorsal surface, crowding the leg at¬

tachments well to the front.

The nymph is now known. It shows the epimeral shield in

two parts and the typical genital area for the genus.

Specimens have been found in Maine, New York, Michigan,

Indiana, Iowa, Minnesota and Louisiana. In Wisconsin they

have been found in lakes and ponds near Cable, Madison and

Eagle River ; in Green, Fox, Mirror and Storr lakes ; in Adams

County and in thirteen lakes in Vilas County.

Atractides jordanensis Mar.

PI. X, fig. 46, 47.

Water mites of the genus Atractides (formerly Torrenticola)

have hard and porous integuments and an unusual development

of plates so that the body appears to lie in layers. This species

shows a typical arrangement of the dorsal plates, one being

very large with four small ones on its anterior border. The

rostrum of the mouth region is short. Colors are bright and

variegated.

Individuals are usually found in silt. They have been taken

in small numbers in Jordan Lake and in the neighboring Goose

Pond; in Razor Back Lake, Vilas Co., and in Green Lake at a

depth of fifteen meters.

Atractides indistinctus Mar.

PI. X, fig. 42-45.

This species is distinguished by the incomplete development

of the four anterior dorsal plates and by the large rostrum,

which with the ends of the first epimeral pair extend well for¬

ward beyond the body margin.

342 Wisconsin Academy of Sciences, Arts, and Letters .

The male is now known. It is smaller than the female (0.575

mm. to the end of the rostrum) ; as usual in this sex, the united

first pair of epimera do not reach to the genital area, as they

do in the female, due to the greater development of the second

and third pairs.

The nymph has likewise been found. The surface is finely

ridged and the dorsal side shows the typical four conspicuous

plates, the posterior one large and shield shaped.

Specimens have been found at moderate depths in Indiana

and in Green and Winnebago lakes in Wisconsin.

Tyrrellia ovalis nov. spec.

PI. VIII, fig. 18-21.

The body is oval ; the largest individual found measures 0.95

mm. The surface is thickly beset with tiny thorns directed

backward; the epimera and legs are porous. Colors are not

known. The anterior dorsal surface shows two pairs of small

irregularly oblong chitinized plates and a number of small hair

plates. The epimera are heavy; in form they resemble those

of Limnesia and all bear a few hairs; the fourth is broad and

bears the articulation for the last leg well toward its inner

posterior margin. The two elongated genital plates lying close

to the epimera bear each three conspicuous acetabula, the two

posterior ones close together; in the specimens found these re¬

semble the plates of female Limnesia. The palpi resemble those

of T . circularis and are likewise similar to those of certain spe¬

cies of the related genus ; a small peg on a conspicuous papilla

shows on the second segment, while the fourth bears a cluster

of hairs on the inner side near the distal end. The legs are

heavy and short, none of them being as long as the body; all

are provided with many small bristles but no swimming hairs

and they end in large weak claws. The third leg is slightly

shorter than the others. The fifth segment of the fourth leg

is bent, as in the related species.

The genus Tyrrellia was erected by Dr. Koenike to contain

the one species, T . circularis, found near Ottowa. The author

has examined this material which is deposited in the Depart¬

ment of Agriculture in Ottawa. Unfortunately the two slides

containing the specimens are not in good condition, but Dr.

Koenike’s descriptions and figures are clear and detailed. The

new species falls well within his characterization of the genus

Marshall — Hydracarina of Wisconsin .

343

except that in place of a single dorsal anterior plate there is

present a pair of small plates. It is distinguished from the

type species not only by this difference in the dorsal surface but

also by its more elongated form and by the shape of the genital

plates of the female. In the few recorded measurements in¬

dividuals of the new species are smaller.

Three specimens of T. ovalis were found in Mendota Bay at

Madison ; it is inferred that they are females. Dr. R. W. Wol¬

cott, in Ward & Whipple's Fresh Water Biology (p. 869) states

that he found specimens of the genus in Michigan lakes; two

species were present, one of which was apparently T . circularis ,

but further data are not given.

Limnesia cornuta Wol.

PI. IX, fig. 22-24.

This rare species of the large genus Limnesia is recognized

by the unusual chitinous meshwork of the body surface, the

presence of a small posterior dorsal plate and the large, finely

serrated antenniform bristles.

It is known for Ontario, Michigan and Tennessee. In Wis¬

consin it has been taken in Clear-Crooked Lake (Vilas Co.) and

in Goose Pond (Adams Co.) at depths from the surface to six

meters.

Limnesia maculata (Mull.)

PL VII, fig. 5.

Dr. R. Piersig (1905), commenting on Dr. Wolcott’s detailed

account of this species in North America (1903), designated it

as a new variety, L. americana, basing his opinion on certain

details of palpi, epimera and genital plates as shown in Wol¬

cott’s figures of a young female. The author, after examining

a large number of specimens and comparing them with identi¬

fied European material, does not find any constant or important

differences existing between them which justifies the formation

of a variety and consequently the name has been dropped. In

respect only to color does there appear to be any appreciable

difference. In European literature the species is described as

almost always red; observations on living material here indi¬

cate that the entire body is only seldom entirely red, though

red spots are common, and that green, yellow and blue predomi-

344 Wisconsin Academy of Sciences , Arts, and Letters .

nate. These color varieties, however, have been reported for

various parts of Europe.

Re-examination of material shows that L. elliptica Mar.

(1924), described from one young female taken in Alaska, is

a synonym for L. maculata and that the name must be dropped.

The species L. maculata is one of the largest, individuals

sometimes measuring 2 mm. It is recognized by the small size

of the palpi and correlated with this the small size of the maxil¬

lary shield which bears them, as well as by the form of the first

epimera, the inner borders of which tend to approach each other

for a greater part of their length.

The species has been found all over Europe, in Turkestan,

northern Asia and northern Africa. In North America it is

known for British Columbia, Ontario, Alaska, New York, Mich¬

igan, Iowa and Montana. In Wisconsin it has been found in

Spooner and Green lakes, Goose Pond (Adams Co.), pools near

East Winona and in Trout Lake and twenty smaller lakes in its

vicinity. It appears to be commoner in northern waters and at

some distance below the surface, since the greater number of

individuals have been found at depths from two to twenty-nine

meters.

Limnesia paucispina Wol.

PI. IX, fig. 25-27.

Mites of this fairly common species are small, measuring less

than 1 mm. Colors are pale browns, sometimes with reds or

blue and orange. Spines and swimming hairs on the legs are

scarce; the maxillary shield has straight sides; and the palpi

are distinctive, being large and stout with a long spine on the

second segment resting on a very short papilla.

Dr. Wolcott erected the species from the examination of a

single preserved female. The male is now known. The relative

size and position of the three acetabula of the genital plates in

both sexes have been found to be variable, as in other species

of the genus.

This species is known for Michigan and Ohio; in Wisconsin

it has been found in small numbers in Green, Lauderdale and

Buffalo Lakes; in pools near Wisconsin Dells and Green Bay;

in the Yahara River and pools near Madison; and in several

lakes in Vilas County at depths from the surface to 7.5 meters.

(One record for 29 meters may be accidental.)

Marshall — Hydracarina of Wisconsin .

345

Limnesia fulgida Koch

PL IX, fig. 31-33.

American forms of this species have been referred to as

variety wolcotti, a name given by Dr. Piersig (1905) in review¬

ing Dr. Wolcott's description of the species in North America

(1903), since he found certain small differences in details of

the genital plates and palpi. The author is now of the opinion

that the creation of a new variety was not justified.

This species and the following, L. undulata, both common in

the Old World, are very closely related; their separation is

difficult and has given rise to much confusion in the literature.

The author, after the examination of hundreds of North Ameri¬

can Limnesia and the study of identified European specimens

has come to the conclusion that the two species intergrade and

are not clearly separated. The same condition has been found

true for two other common American species, Arrhenurus meg-

alurus and A . marshallae, as already reported. There appears

to be, in these Limnesia, individual, sex and age variations;

perhaps the particular environment affects the individual in

some cases, and crossing may occur. Results of this study in¬

dicate that, in general, the two species are separated chiefly on

the basis of certain small, more or less constant difference in

color, palpi and genital plates, especially those of the male ; that

the decision as to which of the two specfes a given individual

is assigned to rests largely on the judgment of the investigator

in balancing these small and sometimes variable characters.

Both species are common, large, and frequently found together.

As interpreted by the author, L. fulgida (formally L. his-

trionica) is brightly colored, sometimes entirely red (as usu¬

ally given in European literature) ; the palpi are large and the

stout second segment has two rows of bristles and a moderately

large, sometimes conical process provided with a peg, often

set obliquely; and on the male genital plates the small hair

papillae are numerous and conspicuous and closely follow the

outer margins of the large acetabula.

This cosmopolitan species is found in nearly all shallow wa¬

ters in the state and probably throughout eastern United States

and Canada.

346 Wisconsin Academy of Sciences , Arts , and Letters .

Limnesia undulata (Mull.)

PL IX, fig. 28-30.

The species is closely related to L. fulgida, as stated under

the description of the latter species, and is distinguished from

it with difficulty. In general it is duller in color and never en¬

tirely red ; the palpi, relative to the legs, are larger and stouter,

although the fifth segment is slimmer, its second segment shows

fewer spines and its process with a peg is longer and slimmer ;

and on the male genital plates the fine hair papillae are not usu¬

ally so numerous nor conspicuous nor placed as irregularly as

in L. fulgida.

This large species appear to be somewhat more widely dis¬

tributed in the United States and Canada than the related spe¬

cies. It is found, usually abundantly, in all shallow waters of

Wisconsin.

Limnesiopsis anomala (Koen.)

PI. VII, fig. 6-8.

This species, the only one known for the genus, closely re¬

sembles the large Limnesia but is never so abundant. It is

readily recognized by the presence of the numerous small ace-

tabula on the genital plates.

The nymph is now known ; its genital plate closely resembles

that found in Limnesia larvae.

The species was first described by Dr. Koenike who took it

for an unusual Limnesia ; it is still regarded by some authors

as forming only a subgenus. The original material came from

Ontario ; this has been examined by the author as well as ma¬

terial since taken from Lake Simcoe, near Toronto. It has also

been found in New York and Michigan. In Wisconsin it has

been found in Green, Winnebago, Pewaukee, Waukesha and the

Madison lakes and in thirteen bodies of water in Vilas County,

usually near the surface.

Hygrobates longipalpis (Herm.)

PL X, fig. 38-41.

Details of the anterior epimeral group and the structure of

the palpi distinguish this genus from Megapus which it closely

resembles ; all of these mites are of moderate size, usually one

millimeter or less in length. Characteristic of this species is

the well developed process on the second palpal segment, the

shape of the fourth epimera and the position of the acetabula

Marshall — Hydracarina of Wisconsin .

847

on the genital plates of the female. Adults when alive are usu¬

ally recognized by their bright brown color on which are con¬

spicuous irregular white branched streaks and sometimes red

spots.

The species H. ruber (Marshall, 1926) has been found on re¬

examination of the material to be a young H . longipalpis; hence

the name, being a synonym, must be dropped.

The species is common in Europe and has also been found in

Asia Minor and northern Africa. It was first reported for the

New World by Dr. Koenike who recognized it in material from

British Columbia which the author has also examined. It has

since been found in Ontario, Wyoming, Montana, Iowa, Illinois,

Michigan, Indiana, Ohio, Tennessee and New York. In Wis¬

consin it has been taken from Winnebago, Green, Spooner, Pe-

waukee, Lauderdale, Nashota and Twin Lakes (Kenosha Co.),

the Madison lakes and from ten lakes in Vilas County, at depths

from the surface to ten meters.

Megapus parvis cutus (Mar.)

PI. X, fig. 34-87.

Members of the genus Megapus (formerly Atractides) are

distinguished from Hygrobates by the more complete separa¬

tion of the first pair of epimera from the capitulum and by

differences in the palpi and first pair of legs. This species

shows less modification of the ends of the last two segments of

the first pair of legs than do most species of the genus.

Re-examination of material now shows that M. (Atractides)

orthopes (Marshall, 1915) is the male of M. parviscutus and

the former name becomes invalid in consequence. The status

of the species M. phenopleces described by the author in the

same paper from one female found in Lake Spooner is in doubt

and will await the study of more material.

The species M. parviscutus has been found in Indiana, Illi¬

nois and Michigan. In Wisconsin it has been found in shallow

water, usually in southern counties, the localities as follows:

lakes Lauderdale, Como, Delavan, Twin, Green, Mirror, Nasho¬

ta, Nagowicka, Buffalo and Spooner, Goose Pond (Adams Co.)

and pools near Minocqua.

Rockford College,

September 1, 1931.

348 Wisconsin Academy of Sciences, Arts, and Letters .

Bibliography

Titles are confined to papers having authors’ descriptions of the

species cited and to general papers describing cosmopolitan species.

Koenike, F. 1895. Nordamerikanische Hydrachniden. Abh. Naturwiss.

Ver. Bremen 13 (2) : 167-226, pi. 1-3.

Marshall, R. 1914. Some New American Water Mites. Trans. Wis.

Acad. 47 (2) : 1300-1304, pi. 92-93.

1915. American Species of the Genus Atractides (Megapus). Trans.

Am. Micr . Soc. 44 (3) : 185-188, pi. 6.

1926. Water Mites of the Okoboji Region. Univ. Iowa Studies Nat.

Hist. 11 (3) : 28-35, pi. 1-4.

1929. The Water Mites of Lake Wawasee. Proc. Indiana Acad. Sci.

38 : 315-320, fig. 1-18.

1929. Canadian Hydracarina. Univ. Toronto Studies, Biol. Series 33

: 57-93, pi. 1-7.

1930. The Water Mites of the Jordan Lake Region. Trans. Wis. Acad.

25 : 245-253, pi. 5-6.

1931. Preliminary List of the Hydracarina of Wisconsin. Trans. Wis.

Acad. 26 : 311-319, pi. 7-8.

Piersig, R. 1905. (Review). Zool. Centralbl. 12 : 197-198.

Soar and Williamson. 1927. British Hydracarina, Vol. II. The Ray

Society, No. 112. London.

Thor, S. 1906. Lebertia-Studien. Zool. Anz. 29 : 761-780, fig. 32-46, 49.

Wolcott, R. W. 1903. The North American Species of Limnesia. Trans.

Am. Micr. Soc. 24 : 85-107, pi. 12-13.

1905. A Review of the Genera of the Water Mites. Trans. Am. Micr.

Soc. 26 : 161-242, pi. 18-27.

350 Wisconsin Academy of Sciences , Arts, and Letters .

Plate VII.

1. Lebertia porosa, ventral plates.

2. Lebertia quinquemaculosa , palpus.

3. Lebertia quinquemaculosa, ventral surface.

4. Lebertia quinquemaculosa, nymph, ventral plates.

5. Limnesia maculata, ventral plates, female.

6. Limnesiopsis anomala, ventral plates, female.

7. Limnesiopsis anomala, genital plates, male.

8. Limnesiopsis anomala, genital plates, nymph.

TRANS. WIS. ACAD., VOL. 27

PLATE VII

352

Wisconsin Academy of Sciences , Arts , and Letters.

Plate VIII.

9. Oxus connatus, ventral view.

10. Oxus connatus, lateral view.

11. Oxus intermedius, ventral view.

12. Oxus intermedius, dorsal view.

13. Oxus elongatus, ventral view.

14. Oxus elongatus, palpus.

15. Frontipoda americana, dorsal view.

16. Frontipoda americana, nymph, ventral view.

17. Frontipoda americana, lateral view.

18. Tyrrellia ovalis, ventral view.

19. Tyrrellia ovalis, right palpus.

20. Tyrrellia ovalis, dorsal view.

21. Tyrrellia ovalis, leg I, left.

TRANS. WIS. ACAD., VOL. 27

354 Wisconsin Academy of Sciences , Arts , and Letters .

Plate IX.

22. Limnesia cornuta, dorsal view.

23. Limnesia cornuta , ventral plates, male.

24. Limnesia cornuta, hair plate and chitinous mesh work, (after Wol¬

cott) .

25. Limnesia paucispina, end of leg IV.

26. Limnesia paucispina, ventral view, male.

27. Limnesia paucispina, palpus.

28. Limnesia undulata, left palpus.

29. Limnesia undulata, genital plates, male.

30. Limnesia undulata, genital plates, nymph.

31. Limnesia fulgida, genital plates, male.

32. Limnesia fulgida, left palpus.

33. Limnesia fulgida, ventral plates, female.

TRANS. WIS. ACAD., VOL. 21

356 Wisconsin Academy of Sciences, Arts, and Letters .

Plate X.

34. Megapus parviscutus , ventral view, male.

35. Megapus parviscutus , palpus.

36. Megapus parviscutus , end of leg I.

37. Megapus parviscutus , genital plates, female.

38. Hygrobates longipalpis, palpus.

39. Hygrobates longipalpis, dorsal view.

40. Hygrobates longipalpis, genital plates, female.

41. Hygrobates longipalpis, genital plates, male.

42. Atractides indistinctus, dorsal view.

43. Atractides indistinctus, nymph, dorsal view.

44. Atractides indistinctus, ventral view, male.

45. Atractides indistinctus, palpus.

46. Atractides jordanensis, lateral view.

47. Atractides jordanensis, anterior dorsal region.

TRANS. WIS. ACAD., VOL. 27

PLATE X

A REPORT ON THE MOLLUSCA OF THE NORTHEAST¬

ERN WISCONSIN LAKE DISTRICT

J. P. E. Morrison

Notes from the Limnological Laboratory of the Wisconsin Geological

and Natural History Survey

No. XLVII.

Introduction

The following is a report on the present knowledge of the

molluscan life of the Highland Lake District of Wisconsin. This

region of the state has been practically unexplored as far as

this group of animals is concerned.

The work of Chadwick was limited, including a few records

from the Wisconsin drainage, near the town of Eagle River.

The work of Winslow was largely limited to the vicinity of the

Arbor Vitae lakes, in the Tomahawk drainage. The records

of Cahn come principally from the vicinity of Sayner and in¬

clude some from Muskellunge Lake.

Baker's work on Tomahawk Lake was largely descriptive, be¬

ing a fairly complete account of one of the large lakes in the

district. He described the habitats of all the species collected,

and classified them on the basis of ecological succession. Strik¬

ingly brought out in his report is the great diversity of eco¬

logical conditions within such a small area.

There are many lakes without inlets or outlets in the area

studied, since the region is topographically young, so little re¬

moved from the effects of the last glacial era, and the streams

have not had time enough to cut back to drain all the lakes that

were left. A brief study of the drainage lines, together with

the great abundance of the lakes, of all sizes, will indicate the

youthfulness of the region. This region is one of the headwa¬

ters of several drainage areas. It includes the headwaters of

the Flambeau River (Chippewa drainage), of the Tomahawk

and Wisconsin Rivers (Wisconsin drainage), of the Montreal

River, West Branch of the Ontonagon River, and the South

Branch of the Presque Isle River (Lake Superior drainage),

and of the Pine River (Lake Michigan drainage).

360 Wisconsin Academy of Sciences , Arts, and Letters.

Examination of the district was undertaken at the suggestion

of Prof. C. Juday, under whom the writer had the pleasure of

working during the summers of 1929 and 1930. The work on

the Mollusca was done in whatever time was to spare from the

quantitative work on the bottom fauna of these lakes.

In the preparation of the lists, records have been secured

from the following sources: (1) Collections made during the

summers of 1929 and 1930. (2) Previous collections, hitherto

unrecorded, including those from the Wis. Geol. & Nat. Hist.

Survey (1928), and some made by Dr. 0. Park, near Sayner,

during September 1927. (3) Previous records, included in Bak¬

er’s Monograph of Wisconsin Fresh Water Mollusca.

Acknowledgments are due the following people who have

aided the work: Prof. Juday under whose supervision the work

was done; Dr. Wm. J. Clench for determination of the Physi-

dae; Dr. Victor Sterki for determination of the Sphaeriidae;

Dr. Bryant Walker for determination of the Ancylidae; to Ed¬

ward Schneberger, Mrs. J. P. E. Morrison, and others whose

assistance in the field has been invaluable.

The lakes in the region examined for Mollusca show a wide

range in degree of softness, with a corresponding range in

acidity. The amount of fixed carbon dioxide present in the open

water of the lakes varies from 1.0 to 30.5 parts per million.

The pH range is from 5.1 to 8.3. In the softest lakes the cal¬

cium content of the water is as low as 0.1 part per million.

It would seem at first sight that molluscs would be unable to

exist in such soft waters as is indicated by a fixed carbon di¬

oxide content of from 1.0 to 5.0 parts per million. However,

careful search has shown their presence in even the softest and

most acid of the lakes. There are two general types of the ex¬

tremely soft lakes : ( 1 ) the type with clear water and usually a

sandy or rocky gravel margin, more or less devoid of plants;

(2) the type with highly colored water, surrounded usually in

part by bog.

Two of the characteristic forms of molluscs found in the soft,

clear lakes are Pisidium and Campeloma. It is a puzzle as to

how Pisidium can draw enough substance for a shell (thin, to

be sure) from water with a pH of 6.0 and a fixed carbon dioxide

content of 1.0 part per million. The snail Campeloma builds

a much larger and thicker shell under the same conditions.

M orrison — M ollusca of Northeastern Wisconsin . 861

In lakes of the soft, bog-surrounded type, there is usually a

little more dissolved carbonate (3.0-5.0 p.p.m.), with a pH of

5.1 to 6.1. Here are to be found in certain lakes, some of the

largest and finest specimens of Pisidium (sp. undescr.), nearly

reaching the dimensions of the largest found in the state. Are

these small bivalves able to hoard enough of the shell building

materials from the water, or is there a better supply in the

particular place in the bottom they inhabit?

None of the family Valvatidae are found at a pH lower than

7.1 and in water softer than that containing 8 parts per mil¬

lion of fixed carbon dioxide.

The Campelomas, the only representatives of the family Vi-

viparidae in the region are able to withstand the more extreme

conditions of a pH of 5.7 or 5.8 and a fixed C02 content of 1.0

part per million. The range of the two species is almost iden¬

tical, showing both of them to be equally generalized in their

habitats.

Among the Amnicolidae, only two species are widespread,

and of these only one is found at any great range below neu¬

trality. The commonest species ( Amnicola limosa porata) is

found in situations ranging from pH 5.7 to 8.3, and from 1 to

30 parts per million of fixed C02 ; all the other species are found

above pH 6.8 and 8 parts per million of fixed C02.

Among the gill-breathing snails, only three species are able

to tolerate the conditions of the extremely soft waters of the

clear lake type. There is a probability that the data used for

these snails may be in error in certain cases. It is only reason¬

able to suppose that Campeloma , where found in abundance in

the (glacial till) clay bottom of an extremely soft water lake,

is getting its necessary supply of carbonates from the clay bot¬

tom directly, and not from the open water of the lake. In the

case of Amnicola, an extra source of shell-building materials

must be sought in the plant food.

Among the Lymnaeidae, three forms are especially tolerant

of acid water (pH to 6.0), while the majority of the species

are found only in water having an alkaline reaction (pH 7.0

to 8.0). Specialization of habitat seems to be rather well

marked in this group, as indicated by the attendant chemical

data. The genus Lymnaea is restricted to waters of pH 7.2 or

more, and a fixed carbon dioxide content of 15 or more parts

362 Wisconsin Academy of Sciences , Arts , and Letters .

per million. In the genus Stagnicola, we find the common form

of southern Wisconsin ( S . palustris elodes) largely replaced in

the northern lakes and ponds by S. exilis and S . lanceata.

These two last named species are found in more acid and softer

lakes than is palustris elodes . On the other hand, the species

of the emarginata group seem to be confined to lakes of pH 7

to 8. In the genus Fossaria, the common species obrussa is

found from pH 5.9 to 8.3, while the supposed ancestral form

F. o. decampi is found under much more restricted conditions,

chiefly in waters having a H-ion concentration of pH 7.5. This

immediately raises the question as to which is the ancestral

form, and which the special form found under a peculiar set of

conditions attendant upon recently formed glacial lakes.

Examination of the family as a whole shows that only Stag¬

nicola and Fossaria are generally distributed under variable

conditions while Lymnaea , Acella, Pseudosuccinea, and Bu-

limnaea are restricted to greater or less degree. Must not the

four last-named genera be considered as more highly specialized

or “senescent” groups as compared with Stagnicola and Fos-

saria ?

Examination of the several described varieties of Helisoma

antrosa brings out some interesting conclusions. The thin-

shelled form, H . a. unicarinata, seems to be restricted to the

softer, more acid waters of the region, while H . a. sayi, which

has a noticeably thicker shell, is not found in lakes that are acid

(pH below 7.0). On the other hand, the two other varieties, H .

a. antrosa and H . a . cahni, are found under variable conditions

(pH 6.0 to 8.0).

The range of the varieties of H. trivolvis and H. campanulata

show simply that the varieties are more restricted in habitat

than is the typical form of each species; for example, H. t .

pilsbryi is found within narrower pH limits than is the typical

H. trivolvis .

In the case of campanulata, the varieties listed in order of

increasing restriction are H . c. campanulata, c . wisconsinensis ,

c. minor, and c. ferrissii.

Different sets of chemical conditions in these lakes seem to

produce specific varieties in a few cases. Also, it would seem

that the variation in chemical nature of the habitat may be the

stimulus for production of non-specific variation in form of

the animal or of the shell it builds.

Morrison — Mollusca of Northeastern Wisconsin . 363

Among the small Planorbids, the forms of the Genus Gyrau-

luSy when regarded in the subgeneric groups, show a tendency

toward serial arrangement of the different forms across the

different conditions of the lakes. In the subgenus Gyranlus

sensu stricto listed from more acid to more alkaline limits of

range are: G. deflectus, G. d. obliquus, G. hirsutus. Of these

three species, that found in the more acid conditions is most

carinate, and the one found under most alkaline conditions is

the least carinate on the periphery of the whorl. In the sub¬

genus TorquiSy a much more marked series is indicated, consist¬

ing of : G. circumstriatus, G. parvus, G. arcticus.

In spite of the paucity of records in the Ancylidae, one dif¬

ference is indicated. Ferrissia parallela is the only species in

the region found in neutral or acid waters. The other three

species are bunched (with one record each) at about pH 7.6.

Parallela is to be found from pH 6.0 to the most alkaline of the

lakes examined for mollusks (pH 8.4).

Two species of Physa show up in a wide range of conditions.

These two are large, thin-shelled, and apparently annuan in

these lake habitats. P. sayii ranges as far as pH 5.7 on the

acid side, while P. laphami is found down to pH 6.4. P. gyrina,

which is more common in southern Wisconsin than in these

northern lakes, is not in acid waters in the lakes. Likewise,

the four other forms recorded were restricted to alkaline water

(pH 7.6-8.0) .

All the species of the Unioninae in the region are restricted

to streams of slightly alkaline reaction (pH 7.0-8.0). The low¬

er limit of fixed carbon dioxide observed was 12.07 parts per

million. The only one of the forms of this subfamily found in

lakes in Vilas Co., is recorded from a lake in the same range of

acidity and hardness of water.

In the subfamily Anodontinae , all except species of Anodonta

are similarly restricted in the chemical nature of the habitat.

Thinnest-shelled of the genus, among the species to be found

in northern Wisconsin, Anodonta marginata is found in many

of these northern lakes, in water varying from pH 6.0 to 8.4

and in fixed carbon dioxide content from 2.6 to 30.5 parts per

million. Under the extremely soft and acid water conditions,

the shell developed by this form is so thin, that it may be twist¬

ed (when fresh and still wet) through about 20 degrees, with-

364 Wisconsin Academy of Sciences, Arts, and Letters .

out even cracking. It is impossible to twist the thicker shells

developed when the animals have grown under slightly alkaline

conditions.

All of the species of the subfamily Lampsilinae, like the ma¬

jority of species of the fresh water mussels, are limited to

slightly alkaline waters. Detailed examination of the range of

the two species of Lampsilis shows that the lake and the stream

variety of each have approximately the same limits. The de¬

velopment of the lake form is not due to differences of H-ion

concentration or of the amount of fixed carbon dioxide present,

as far as the writer’s studies are concerned.

In the Sphaeriidae, some striking differences of chemical na¬

ture of the habitat are seen. In general the distribution of spe¬

cies of the “Finger-nail” and “Pill” Clams shows the condition

expected of a diversified group, some widespread, some inter¬

mediate, and some species confined to narrow limits of H-ion

concentration and of amount of fixed carbon dioxide present in

the water.

On examination of the groups within the family, or within

genera, we get more precise information. For example : Pisidi-

um surpasses the other two genera in tolerance for acidity and

ability to thrive in the softest waters. It is found in water

with pH 5.7 and a fixed carbon dioxide content of 1.5 parts per

million. Musculium, which has a proportionately thinner shell,

is found only as low as pH 5.9 and with a fixed carbon dioxide

content of 2.6 parts per million. Sphaerium, as a unit, is found

in habitats approximately neutral, or alkaline in reaction (pH

6.8-8.4) and with a fixed carbon dioxide content of 9.3 or more

parts per million. But there is one straggler. S. occidentale is

restricted to the acid side of the scale, having been taken in the

region only from temporary ponds, with pH 5. 8-5. 9 and a fixed

carbon dioxide content of 5.5 to 7.5 parts per million. Is this

physiological difference not marked enough to indicate that S .

occidentale may be less closely related to the other Sphaeria

than usually regarded ? Another good example of physiological

isolation of species is seen in the group of Pisidium rotunda -

turn. In this group P. ferrugineum and P. vesicular e are both

found between pH 7.2 and 8.2 and a fixed carbon dioxide con¬

tent between 11 and 22.5 parts per million. In direct contrast,

P. rotundatum is found between pH 5.8 and 6.2 and from a

fixed carbon dioxide content of 2.0 to 9.0 parts per million.

Morrison — Mollusca of Northeastern Wisconsin . 365

The lakes that are intermediate in hardness (10.0-20.0 p.p.m.

fixed carbon dioxide, and a pH of 7.0-7.6) harbor the greatest

number of species. As would be expected, the hardest lakes

examined contain the greatest abundance of individuals.

Stream conditions are chemically rather uniform in the dis¬

trict, paralleling the intermediate lakes in character (Fig. 127).

Chemical factors are thus not a limiting factor for molluscs in

the streams. Geographic distribution and size and flow of the

streams do seem to be important.

The number of species of Unionids in the small headwater

streams of the Lake Superior and Green Bay (Lake Michigan)

drainages is about one-half that found in similar streams, un¬

der comparable conditions, in the headwaters of the Flambeau,

Tomahawk, and Wisconsin drainages.

The Wisconsin River, examined at various places from its

source to a point in northern Oneida County, shows remarkably

well the variation and increase of the molluscan fauna in co¬

ordination with the increase in size of the stream, as noted by

Adams, Ortmann, Grier, and Baker.

In this northern lake region, where some streams are ponded

for mile after mile, with swampy or bog margins, and others

are rapid, with sand or gravel beds, the molluscan fauna of the

streams shows a corresponding difference. For example,

Sphaerium fallax and S. rhomb oideum are found in the swampy

margins of ponded streams, while S. stamineum and S. emargi-

natum are characteristic of streams with a good current over

sandy bottom.

In all, some ninety-six lakes and thirty-eight stream localities

have been examined, included in Vilas County and the adjoin¬

ing portions of Iron, Price, Oneida, and Forest Counties. A

total of one hundred twenty-nine forms of molluscan life are

here recorded from the area. These are distributed as follows

in the major groups:

Gill-breathing univalves _ _ _ 11

Lung-breathing univalves _ _ 51

Unionidae (bivalves) _ 26

Sphaeriidae (bivalves) _ _ _ 41

Total _ _ 129

366 Wisconsin Academy of Sciences , Arts , and Letters .

Three forms are added to those known to occur in the state,

namely: Pseudosuccinea columella chalybea (Gould), Pisidium

fallax septentrionale Sterki, Pisidium punctatum Sterki.

The system of classification followed in this report is that of

Baker’s Monograph. For further references, the reader is re¬

ferred to that publication.

Systematic Catalogue of Species

In the following list the name of each species or variety is

followed by a record of the localities where it is known to oc¬

cur, listed according to drainage areas. Except where the au¬

thority for the record is otherwise stated, the records are those

of the 1929-1930 collections of the Wisconsin Geological and

Natural History Survey.

The area included in this brief report has not been ex¬

haustively explored: there are about a thousand lakes in the

entire district! Any additions and corrections will be grate¬

fully received by the author.

Class GASTROPODA.

Subclass Streptoneura Spengel.

Order Ctenorranchiata Schweigger.

Suborder Platypoda Lamarck.

Superfamily Taeniglossa Bouvier.

Family Valvatidae Gray.

Genus Valvata Muller.

Valvata tricarinata (Say).

pH— 7.16-8.37; fixed carbon dioxide=8.16-30.56 p.p.m. (Fig. 1.).

Lake Superior Drainage: Palmer Lake.

Flambeau Drainage: Allequash L.; Lake Laura; Mann L.; Silver

L.; Trout L.; White Sand L.; Wildcat Lake.

Tomahawk Drainage: Kawaguesaga L.; Little Arbor Vitae Lake

(Winslow, Baker).

Wisconsin Drainage: Plum L.; Razorback L.; Star Lake.

Valvata sincera nylanderi Dali.

pH—7.6; fixed carbon dioxide— 22.5 p.p.m. (Fig. 2.).

Tomahawk Drainage: Little Arbor Vitae Lake (Winslow, Baker).

Valvata lewisii (Currier).

pH — 7.35-7.7 ; fixed carbon dioxide™ 10.65-22.1 p.p.m. (Fig. 3.).

Morrison ■ — Mollusca of Northeastern Wisconsin. 867

Lake Superior Drainage: Palmer Lake.

Flambeau Drainage: Papoose L.; Trout L.; Upper Gresham L.;

Whitefish L.; White Sand Lake.

Tomahawk Drainage: Brandy Lake.

Wisconsin Drainage: Plum Lake.

Family Viviparidae (Gray) Gill.

Subfamily Lioplacinae (Gill) Baker.

Genus Campeloma Rafinesque.

Campeloma decisum (Say).

pH = 5.68-8.37; fixed carbon dioxide = 1.2-25.75 p.p.m. (Fig. 4).

Lake Superior Drainage: South Branch, Presque Isle River, at

Winegar.

Green Bay Drainage: Butternut Lake.

Flambeau Drainage: Big L.; Diamond L.; Fishtrap L.; Helen L.;

High L.; Inlet of Trout L.; Little Long L.; Manitowish River, at

Boulder Junction and 4 mi. southwest; Mann L. Outlet; Marion

L.; Rest L.; South Fork, Flambeau River, at Fifield; Trout L.;

Trout River, at Trout L.; Turtle River, below Lake of the Falls;

White Sand Lake inlet.

Tomahawk Drainage: Little Star Lake.

Wisconsin Drainage: Gilmore Creek and Wisconsin River, north¬

east of Lake Tomahawk (Baker) ; Deerskin River, 6 mi. south of

Phelps; Finley L. ; Plum L. ; Wisconsin River, at Lac Vieux Des¬

ert, at Otter Rapids, 5 mi. west of Eagle River, and at Rainbow

Rapids, southeast of Lake Tomahawk.

Campeloma milesii (Lea).

pH=5. 86-8.0 ; fixed carbon dioxide=l.l-24.73 p.p.m. (Fig. 5).

Lake Superior Drainage: Anna L.; Carlin L. ; Palmer L.; Katinka

L.; Presque Isle Lake.

Flambeau Drainage: Big Muskellunge and White Sand Lakes (Cahn,

Baker) ; Lower Gresham Lake (Juday, Baker) ; Big Muskellunge

L.; Boulder L.; Crooked L.; Ike Walton L.; Inlet of White Sand

L.; Irving L. Outlet; L. Constance; Little White Birch L.; Lost

Canoe L.; Mary L.; Trout L.; Turtle River, at Winchester; White-

fish L.; White Sand Lake.

Tomahawk Drainage: Tomahawk Lake (Baker); Brandy L. ; John¬

son L. ; Skunk L.; Tomahawk River, 4 mi. west of Minocqua; Web¬

er Lake.

Wisconsin Drainage: Plum Lake (Cahn, Baker); Crescent L.; Plum

L.; Razorback L.; Star L.; Sterrett L.; Wisconsin River, 5 mi. be¬

low Lac Vieux Desert.

368 Wisconsin Academy of Sciences, Arts , and Letters .

Family Amnicolidae (Tryon) Gill.

Subfamily Amnicolinae Gill.

Genus Amnicola Gould and Haldeman.

Amnicola limosa (Say) .

pH=7.95; fixed carbon dioxide=30.56 p.p.m. (Fig. 6).

Flambeau Drainage: Wildcat Lake.

Amnicola limosa porata (Say).

pH=5.68-8.37; fixed carbon dioxide=1.2-30.56 p.p.m. (Fig. 7).

Lake Superior Drainage : Harris L.; Montreal River, at Pine L.;

Palmer L.; Presque Isle L.; South Branch, Presque Isle River, at

Winegar.

Flambeau Drainage : Allequash L. ; Big Lake Outlet ; Big Muskel-

lunge L. ; Boulder L.; Catfish L. ; Clear Crooked L. ; Dead Pike L.;

Diamond L.; Fishtrap L.; Harvey L.; Helen L.; High L.; Ike Wal¬

ton L. ; Inlet of White Sand L. ; Inlet of Trout L.; L. Laura ; Little

Crooked L. ; Little White Birch L.; Lost Canoe L. ; Mann L.; Mann

Lake Outlet ; Nebish L. ; Nixon Lake Outlet; Papoose L.; Par¬

tridge L.; Trout L.; Whitefish L.; White Sand L.; Whitney L.;

Wildcat L.; Wolf Lake.

Tomahawk Drainage : Little Arbor Vitae Lake (Winslow, Baker) ;

Tomahawk Lake (Baker) ; Blue L.; Brandy L.; Carroll L.; Clear

L. ; Kawaguesaga Lake.

Wisconsin Drainage : Bragonier L.; Crescent L.; Plum L.; Razor-

back L.; Star L. ; Wisconsin River, at Rainbow Rapids, southeast

of Lake Tomahawk.

Amnicola limosa parva (Lea).

pH=7.64; fixed carbon dioxide=18.87 p.p.m. (Fig. 8).

Flambeau Drainage : Trout Lake.

Amnicola lustrica decepta Baker.

pH=6.85-8.37; fixed carbon dioxide=9.3-30.56 p.p.m. (Fig. 9).

Lake Superior Drainage : Ontonagon River, Mich., 3 mi. north of

Tenderfoot L.; Palmer L.; Presque Isle Lake.

Flambeau Drainage : Big Muskellunge L.; Boulder L. ; High L.;

Lake Laura ; Little Crooked L.; Little Rice L.; Little White Birch

L.; Mann L.; Trout L.; Upper Gresham L.; Whitefish L.; White

Sand L.; Whitney L.; Wildcat L.; Wolf Lake.

Tomahawk Drainage : Little Arbor Vitae Lake (Winslow, Baker).

Wisconsin Drainage : Plum Lake ( Cahn, Baker) ; Crescent L.; Plum

L.; Star Lake.

Amnicola walkeri Pilsbry.

pH—7.16-7.64; fixed carbon dioxide=8.16-22.5 p.p.m. (Fig. 10).

Flambeau Drainage : Big Muskellunge Lake (Cahn, Baker) ; Fish-

trap L.; Trout River, at Trout Lake.

Tomahawk Drainage : Little Arbor Vitae Lake (Winslow, Baker).

Wisconsin Drainage : Razorback Lake.

Morrison — Mollusca of Northeastern Wisconsin. 369

Subfamily Lithoglyphinae Fisher.

Genus Somato gyrus Gill.

Somatogyrus tryoni Pilsbry and Baker.

pH=7.0 ; fixed carbon dioxide=13.0 p.p.m. (Fig. 11).

Wisconsin Drainage: Wisconsin River, at Otter Rapids, 5 mi. west

of Eagle River, and at Rainbow Rapids, southeast of Lake Toma¬

hawk.

Subclass Euthyneura Spengel.

Order Pulmonata Cuvier.

Suborder Basommatophora A. Schmidt.

Superfamily Limnophila.

Family Lymnaeidae (Broderip) Baker.

Genus Lymnaea Lamarck.

Lymnaea stagnalis jugularis Say.

pH=7.6~8.16; fixed carbon dioxide=15. 8-23.0 p.p.m. (Fig. 12).

Flambeau Drainage: Inlet stream, Trout Lake; Outlet of Big Lake.

Tomahawk Drainage: Tomahawk Lake (Baker); Brandy L.; Car-

roll L.; Johnson Lake.

Wisconsin Drainage: Plum Lake; Plum Creek.

Lymnaea , stagnalis lillianae F. C. Baker.

pH = 7.2-8.02; fixed carbon dioxide = 14.9-30.56 p.p.m. (Fig. 13).

Lake Superior Drainage: Ontonagon River, Mich., 3 mi. north of

Tenderfoot Lake.

Flambeau Drainage: Big L. ; Fishtrap L. ; High L.; Trout L. ; Trout

River at Trout Lake; Wildcat Lake.

Tomahawk Drainage: Tomahawk Lake (Baker).

Wisconsin Drainage: Star Lake.

Lymnaea stagnalis sanctamariae Walker.

pH=7.35-8.0; fixed carbon dioxide=16.45-24.73 p.p.m. (Fig. 14).

Lake Superior Drainage: Presque Isle Lake.

Green Bay Drainage: Butternut Lake.

Tomahawk Drainage: Little Arbor Vitae Lake (Juday, Winslow,

Baker) ; Ponds and Stream at State Fish Hatchery, Woodruff.

Genus Stagnicola (Leach) Jeffreys.

Stagnicola palustris elodes (Say).

pH=7.4; fixed carbon dioxide=21.0 p.p.m. (Fig. 15).

Lake Superior Drainage: Pond near South Branch, Presque Isle

River, at Winegar.

Flambeau Drainage: Stream at Fish Hatchery, Woodruff.

870 Wisconsin Academy of Sciences , Arts , and Letters .

Stagnicola exilis (Lea).

pH=5.9-7.74; fixed carbon dioxide=7.5-22.56 p.p.m. (Fig. 16).

Flambeau Drainage: Fishtrap L.; Forest Ponds, 10 mi. northeast

of Boulder Junction; High L.; Turtle River, below Lake of the

Falls.

Tomahawk Drainage: Little Star Lake.

Stagnicola lanceata (Gould).

pH=6.95-7.7; fixed carbon dioxide=7.5-22.56 p.p.m. (Fig. 17).

Lake Superior Drainage: Armour Lake.

Flambeau Drainage: High Lake.

Tomahawk Drainage: Tomahawk Lake (Baker); Little Rice River.

Wisconsin Drainage: Plum Lake (Cahn, Baker).

Stagnicola emarginata (Say).

pH=7. 5-8.0 ; fixed carbon dioxide=14.3-24.73 p.p.m. (Fig. 18).

Lake Superior Drainage: Presque Isle Lake.

Flambeau Drainage: Rest Lake.

Tomahawk Drainage: Kawaguesaga Lake.

Wisconsin Drainage: Plum Lake (Cahn, Baker); Plum Creek and

Lake.

Stagnicola emarginata vilas ensis F. C. Baker.

pH=7.21; fixed carbon dioxide=9.59 p.p.m. (Fig. 19).

Flambeau Drainage: Big Muskellunge Lake (Cahn, Baker).

Stagnicola emarginata wisconsinensis F. C. Baker.

pH = 7.21; fixed carbon dioxide = 16.7-22.5 p.p.m. (Fig. 20).

Tomahawk Drainage: Little Arbor Vitae Lake (Winslow, Baker);

Tomahawk Lake (Baker).

Stagnicola catascopium (Say).

pH=7.64; fixed carbon dioxide=18.87 p.p.m. (Fig. 21).

Flambeau Drainage: Trout Lake.

Genus Acella Haldeman.

Acella haldemani (“Desh.” Binney).

pH = 7.36-7.7; fixed carbon dioxide= 17.0-22.56 p.p.m. (Fig. 22).

Lake Superior Drainage: Harris Lake.

Flambeau Drainage: Fishtrap Lake; Channel between Fishtrap and

High Lakes; High Lake.

Genus Pseudosuccinea Baker.

Pseudosuccinea columella (Say).

pH=6.13-7.6; fixed carbon dioxide=2.75-16.7 p.p.m. (Fig. 23).

Lake Superior Drainage: Anna Lake.

Flambeau Drainage: Channel between Fishtrap and High Lakes.

Tomahawk Drainage: Tomahawk Lake (Baker); Clear Lake.

Morrison — Mollusca of Northeastern Wisconsin. 371

Pseudosuccinea columella chalybea (Gould).

pH=6.06-7.8; fixed carbon dioxide=3.06-18.36 p.p.m. (Fig. 24).

Flambeau Drainage: Catfish L.; Fishtrap L.; Helen Lake.

Genus Bulimnaea Haldeman.

Bulimnaea megasoma (Say).

pH=6.6-8.37; fixed carbon dioxide— 9.3-25.75 p.p.m. (Fig. 25).

Lake Superior Drainage: Pond near South Branch, Presque Isle

River, at Winegar.

Flambeau Drainage: Duck L.; Fishtrap L.; Channel between Fish-

trap and High Lakes; High L.; Little Rice L.; Mann L. Outlet;

Pike L. inlet; Trout L.; Turtle River, below Lake of the Falls;

White Sand Lake.

Tomahawk Drainage: Tomahawk Lake (Baker).

Wisconsin Drainage: Plum Lake (Cahn, Baker); Slough along Wis¬

consin River, northeast of Lake Tomahawk (Baker).

Genus Fossaria Westerlund.

Fossaria modicella (Say).

pH=7.0; fixed carbon dioxide— 13.0 p.p.m. (Fig. 26).

Wisconsin Drainage: Wisconsin River, northeast of Lake Toma¬

hawk (Baker).

Fossaria obrussa (Say).

pH = 5.86-8.37; fixed carbon dioxide = 1.26-25.75 p.p.m. (Fig. 27).

Flambeau Drainage: Ike Walton L. ; Little Rice L. ; Mann L. Out¬

let; Pond along Mann L. Outlet; Trout Lake.

Tomahawk Drainage: Tomahawk Lake (Baker).

Wisconsin Drainage: Found Lake (Cahn, Baker); Star Lake.

Fossaria obrussa decampi (Streng).

pH=7.42-7.7; fixed carbon dioxide=10.65-18.87 p.p.m. (Fig. 28).

Flambeau Drainage: Upper Gresham Lake (Juday, Baker); Little

White Birch L.; Trout L.; Whitefish Lake.

Wisconsin Drainage: Plum Lake.

Fossaria exigua (Lea).

pH=7.7-8.37; fixed carbon dioxide=13.0-25.75 p.p.m. (Fig. 29).

Lake Superior Drainage: Montreal River, at Pine Lake.

Flambeau Drainage: Mann Lake.

Family Planorbidae H. & A. Adams.

Genus Helisoma Swainson.

Helisoma antrosa (Conrad)

pH=6.03-8.02; fixed carbon dioxide=2.66-30.56 p.p.m. (Fig. 30).

Lake Superior Drainage: Montreal River, at Pine Lake; Palmer L.;

Presque Isle Lake.

372 Wisconsin Academy of Sciences , Arts , and Letters.

Green Bay Drainage: Butternut Lake.

Flambeau Drainage: Big L.; Big Muskellunge L. ; Boulder L.;

Helen L. ; High L.; L. George; Lost Canoe L.; Manitowish River,

4 mi. southwest of Boulder Junction; Outlet of Big L.; Rest L.;

Trout L. ; Trout River, at Trout L.; Whitefish L.; Wildcat Lake.

Tomahawk Drainage: Brandy L.; Little Star L.; Skunk L.; Stream,

10 mi. southwest of Hazelhurst; Willow River Flowage, 14 mi.

southwest of Hazelhurst.

Wisconsin Drainage: Crescent L.; Deerskin River, 6 mi. south of

Phelps; Plum L.; St. Germaine River; Star L.; Wisconsin River,

at Rainbow Rapids, southeast of Lake Tomahawk.

H elisoma antrosa unicarinata (Haldeman).

pH=6.05-7.85; fixed carbon dioxide=l.l-18.36 p.p.m. (Fig. 31).

Green Bay Drainage: Kentuck Lake.

Flambeau Drainage: Big Muskellunge L. ; Fishtrap L. ; Channel be¬

tween Fishtrap and High Lakes; Mary L.; Nixon L. Outlet; White

Sand L. Inlet.

Tomahawk Drainage: Tomahawk Lake (Baker); Little Rice River;

Pond near State Fish Hatchery Ponds, at Woodruff; Weber Lake.

Wisconsin Drainage: Razorback L. ; Star Lake.

H elisoma antrosa sayi F. C. Baker.

pH—7.13-8.37; fixed carbon dioxide=9.59-25.75 p.p.m. (Fig. 32).

Flambeau Drainage: Big Muskellunge L.; Nixon Lake (Cahn, Bak¬

er) ; Fishtrap L. ; Mann L. ; Outlet of Mann L. ; White Sand Lake.

Tomahawk Drainage: Little Arbor Vitae Lake (Winslow, Baker);

Tomahawk Lake (Baker).

Wisconsin Drainage: Plum L. ; Found Lake (Cahn, Baker).

H elisoma antrosa cahni F. C. Baker

pH=6.13-8.0; fixed carbon dioxide=2.75-24.73 p.p.m. (Fig. 33).

Lake Superior Drainage: Anna L.; Armour L.; Presque Isle Lake.

Flambeau Drainage: Big Muskellunge Lake (Baker); Silver Lake.

H elisoma trivolvis (Say).

pH=6.6-8.37; fixed carbon dioxide=7.5-30.56 p.p.m. (Fig. 34).

Lake Superior Drainage: Black Oak L.; Palmer Lake.

Flambeau Drainage: Allequash L.; Duck L.; Fishtrap L.; High

L.; Inlet of Trout L.; Inlet of White Sand L.; Irving L. Outlet;

Little Rice L. ; Mann L. ; Outlet of Mann L.; Outlet of Nixon L.;

Pike L.; Trout L.; Trout River, at Trout L.; Turtle River, below

Lake of the Falls; White Sand L.; Wildcat Lake.

Tomahawk Drainage: Tomahawk Lake (Baker) ; Willow River Flow-

age, 14 mi. southwest of Hazelhurst.

Wisconsin Drainage: Crescent L. ; Deerskin River, 6 mi. south of

Phelps; Plum L. ; Rice Creek, near Plum Lake.

H elisoma trivolvis pilsbryi (F. C. Baker).

pH=7.2-8.37; fixed carbon dioxide=13.3-25.75 p.p.m. (Fig. 35).

Morrison — Mollusca of Northeastern Wisconsin. 37B

Lake Superior Drainage: Ontonagon River, Mich., 3 mi. north of

Tenderfoot Lake.

Flambeau Drainage: Boulder L. ; Fishtrap L.; High L. ; Mann Lake.

Tomahawk Drainage: Tomahawk Lake (Baker); Brandy Lake.

Helisoma trivolvis winslowi (F. C. Baker).

ptL~7.6-7.65 ; fixed carbon dioxide=22. 5-22.6 p.p.m. (Fig. 36).

Flambeau Drainage: Manitowish River (Winslow, Baker).

Tomahawk Drainage: Big and Little Arbor Vitae Lakes (Winslow,

Baker) .

Helisoma pseudo trivolvis (F. C. Baker).

PH=7 .23; fixed carbon dioxide=10.8 p.p.m. (Fig. 37).

Flambeau Drainage: Lake Laura.

Helisoma campanulata (Say).

pH—6.6-8.16 ; fixed carbon dioxide=7.5-30.56 p.p.m. (Fig. 38).

Lake Superior Drainage: Palmer Lake.

Green Bay Drainage: Butternut L.; Kentuck Lake.

Flambeau Drainage: Allequash L.; Big L.; Big Muskellunge L.;

Boulder L. ; Fishtrap L.; High L. ; Papoose L.; Trout L. ; White-

fish L.; Wildcat L.; Wolf Lake.

Tomahawk Drainage: Brandy L.; Carroll L.; Johnson L.; Kawa-

guesaga L. ; Little Star L. ; Willow River Flowage, 14 mi. south¬

west of Hazelhurst.

Wisconsin Drainage: Crescent L. ; Plum L. ; Razorback L.; Star

Lake.

Helisoma campanulata minor (Bunker).

pH—6.6-7.85 ; fixed carbon dioxide=9. 59-18. 87 p.p.m. (Fig. 39).

Lake Superior Drainage: Ontonagon River, Mich., 3 mi. north of

Tenderfoot Lake.

Flambeau Drainage: Big Muskellunge L.; Catfish L.; Inlet of White

Sand L. ; Outlet of Nixon L. ; Trout Lake.

Wisconsin Drainage: Plum L.; Star Lake.

Helisoma campanulata ferrissii (F. C. Baker).

pH=7.05; fixed carbon dioxide=13.7 p.p.m. (Fig. 40).

Flambeau Drainage: Island Lake.

Helisoma campanulata wisconsinensis (Winslow) .

pH=6.95-8.37 ; fixed carbon dioxide=7.5-25.75 p.p.m. (Fig. 41).

Lake Superior Drainage: Armour L.; Harris L.; Presque Isle Lake.

Flambeau Drainage: Big Muskellunge L. ; Nixon L. ; White Sand

Lake (Cahn, Baker); Allequash L.; Big Muskellunge L.; High L.;

Lost Canoe L. ; Mann L. ; Turtle River, below Lake of the Falls;

White Sand Lake.

Tomahawk Drainage: Big Arbor Vitae L.; Little Arbor Vitae L.;

Tomahawk L.; Madeline Creek, near Woodruff (Winslow, Baker);

Little Arbor Vitae Lake (Cahn, Baker); Tomahawk Lake (Baker).

Wisconsin Drainage: Found L.; Plum Lake (Cahn, Baker); St.

Germaine Lakes (Winslow, Baker).

374 Wisconsin Academy of Sciences, Arts, and Letters .

Genus Planorbula Haldeman.

Planorbula armigera (Say).

pH=6.6-7.6; fixed carbon dioxide=7.5-16.7 p.p.m. (Fig. 42).

Tomahawk Drainage: Tomahawk Lake and swamp ponds in vicinity

(Baker); Willow River Flowage, 14 mi. southwest of Hazelhurst.

Wisconsin Drainage: Ponds in swamp along Wisconsin River, 4 mi.

northeast of Tomahawk Lake (Baker).

Genus Menetus H. & A. Adams.

Menetus exacuous (Say).

pH=7.0-7.64; fixed carbon dioxide=9.3-22.5 p.p.m. (Fig. 43).

Lake Superior Drainage: Palmer Lake.

Flambeau Drainage: Fishtrap L.; Little Rice L. ; Manitowish River,

4 mi. southwest of Boulder Junction; Pond along outlet of Mann

L.; Trout Lake.

Tomahawk Drainage: Little Arbor Vitae Lake (Winslow, Baker).

Wisconsin Drainage: Crescent Lake.

Menetus exacuous meg as (Dali).

pH=7.1-8.37; fixed carbon dioxide=9.59-25.75 p.p.m. (Fig. 44).

Flambeau Drainage: Big Muskellunge Lake (Cahn, Baker); Big

Muskellunge L.; Mann L.; Outlet of Nixon L.; Trout Lake.

Tomahawk Drainage: Kawaguesaga Lake.

Genus Gyraulus Charpentier.

Gyraulus hirsutus (Gould).

pH=7.1-7.95; fixed carbon dioxide=9.5-30.56 p.p.m. (Fig. 45).

Flambeau Drainage: Boulder L. ; Little White Birch L. ; Nelson L.;

Partridge L.; Trout L.; Wildcat Lake.

Tomahawk Drainage: Little Arbor Vitae Lake (Winslow, Baker);

Tomahawk Lake (Baker).

Wisconsin Drainage : Found L. ; Plum Lake (C'ahn, Baker) ; Plum

L.; Star Lake.

Gyraulus deflectus (Say).

pH=6.2-8.37; fixed carbon dioxide=2.1-30.56 p.p.m. (Fig. 46).

Lake Superior Drainage: Armour Lake.

Flambeau Drainage: Allequash L.; Dead Pike L.; Fishtrap L.; High

L.; Inlet of Trout L.; Outlet of Mann L. ; Pond along Mann L.

Outlet; Whitefish L.; Wildcat Lake.

Tomahawk Drainage: Clear L. ; Little Rice River; Willow River

Flowage, 14 mi. southwest of Hazelhurst.

Wisconsin Drainage: Bragonier Lake.

Gyraulus deflectus obliquus (DeKay).

pH=6.4-8.37 ; fixed carbon dioxide— 8.16-30.56 p.p.m. (Fig. 47).

Lake Superior Drainage: Montreal River, at Pine L.; Palmer L.;

Presque Isle Lake.

M orris on — M ollusca of Northeastern Wisconsin . 375

Flambeau Drainage: Fishtrap L.; Inlet of Trout Lake; Mann L.;

Papoose L. ; Trout L. ; Whitefish L.; White Sand L.; Wolf Lake.

Tomahawk Drainage: Little Arbor Vitae Lake (Winslow, Baker);

Brandy L. ; Carroll L. ; Johnson L. ; Pond near State Fish Hatch¬

ery, at Woodruff.

Wisconsin Drainage: Plum Lake (Cahn, Baker); Shore pools, Wis¬

consin River, 4 mi. northeast of Tomahawk Lake (Baker) ; Crescent

L.; Razorback Lake.

Gyraulus parvus (Say).

pH = 7.0-8.16; fixed carbon dioxide = 8.16-30.56 p.p.m. (Fig. 48).

Lake Superior Drainage: Montreal River, at Pine L.; Ontonagon

River, Mich., 3 mi. north of Tenderfoot L. ; Pond, near South

Branch, Presque Isle River, Winegar; Presque Isle Lake.

Flambeau Drainage: Big Muskellunge Lake (Cahn, Baker); Big

Muskellunge L.; Boulder L.; Inlet of Trout L.; Lake Laura; Little

Rice L. ; Little White Birch L. ; Outlet of Big L.; Outlet of Nixon

L.; Silver L.; Trout L.; Upper Gresham L.; Whitefish L.; White

Sand L. ; Wildcat Lake.

Tomahawk Drainage: Tomahawk Lake and kettle hole ponds in vi¬

cinity (Baker); Carroll L.; Stream, 10 mi. southwest of Hazel-

hurst.

Wisconsin Drainage: Plum Lake (Cahn, Baker); Razorback Lake.

Gyraulus circumstriatus (Try on).

pH=5.9-7.7; fixed carbon dioxide— 2.9-18.87 p.p.m. (Fig. 49).

Flambeau Drainage: Forest Ponds, 10 mi. northeast of Boulder

Junction; Trout L.; Whitefish Lake.

Tomahawk Drainage: Clear Lake.

Wisconsin Drainage: Plum Lake.

Gyraulus arcticus (“Beck” Moller).

pH—8.37; fixed carbon dioxide=25.75 p.p.m. (Fig. 50).

Flambeau Drainage: Mann Lake.

Family Ancylidae Menke.

Subfamily Ferrissiinae Walker.

Genus Ferrissia Walker.

Ferrissia parallela (Haldeman).

pH—6.05-8.37 ; fixed carbon dioxide— 2.75-25.75 p.p.m. (Fig. 51).

Lake Superior Drainage: Ontonagon River, Mich., 3 mi. north of

Tenderfoot L.; Palmer Lake.

Flambeau Drainage: Boulder L.; Fishtrap L.; High L.; Mary L.;

Mud L. ; Outlet of Mann L. ; Turtle River, below Lake of the Falls.

Tomahawk Drainage: Tomahawk Lake, and kettle hole ponds in

the vicinity (Baker) ; Stream, 10 mi. southwest of Hazelhurst.

Wisconsin Drainage: Deerskin River, 6 mi. south of Phelps; Plum

L.; Razorback Lake.

376 Wisconsin Academy of Sciences , Arts , and Letters .

Ferrissia tarda (Say).

pH=7.63; fixed carbon dioxide~2Q.l p.p.m. (Fig. 52).

Tomahawk Drainage: Tomahawk River, 4 mi. west of Minocqua.

Ferrissia fusca (C. B. Adams).

pH=7.58; fixed carbon dioxide=15.2 p.p.m. (Fig. 53).

Flambeau Drainage: White Sand Lake.

Ferrissia kirklandi (Walker).

pH=7.6; fixed carbon dioxide=22.5 p.p.m. (Fig. 54).

Tomahawk Drainage: Little Arbor Vitae Lake (Winslow, Baker).

Family Physidae Dali.

Genus Physa Draparnaud.

Physa laphami (Baker).

pH=6.4-8.02; fixed carbon dioxide=2.9-24.73 p.p.m. (Fig. 55).

Lake Superior Drainage: Armour L.; Harris L.; Montreal River,

at Pine L. ; Ontonagon River, Mich., 3 mi. north of Tenderfoot L.;

Presque Isle Lake.

Flambeau Drainage: Big L.; High L.; Lost Canoe L.; Whitney

Lake.

Tomahawk Drainage : Clear L. ; Little Star L. ; Pond near State Fish

Hatchery ponds, at Woodruff.

Wisconsin Drainage: Crescent L. ; Wisconsin River, at Rainbow

Rapids, southeast of Lake Tomahawk.

Physa sayii Tappan.

pH=5.68-7.96; fixed carbon dioxide=1.2-22.5 p.p.m. (Fig. 56).

Lake Superior Drainage: Palmer Lake.

Flambeau Drainage: Big Muskellunge L.; Nixon Lake (Cahn, Bak¬

er) ; Allequash L. ; Ballard L. ; Big Muskellunge L. ; Catfish L.;

Crystal L.; Dead Pike L.; Diamond L.; Fishtrap L.; Harvey L.;

Island L. ; Little Rice L. ; Manitowish River, 4 mi. southwest of

Boulder Junction; Marion L.; Silver Lake.

Tomahawk Drainage: Little Arbor Vitae Lake (Cahn, Winslow,

Baker); Tomahawk Lake (Baker); Brandy L.; Johnson Lake.

Wisconsin Drainage: Plum Lake (Cahn, Baker); Deerskin River, 6

mi. south of Phelps ; Plum L. ; Razorback L. ; Star Lake.

Physa obrussoides (F. C. Baker).

pH=7.64; fixed carbon dixoide=18.87 p.p.m. (Fig. 57).

Flambeau Drainage: Roadside spring, 3 mi. northwest of Winches¬

ter; Trout River, at Trout Lake.

Physa gyrina Say.

pH=7.1-8.37 ; fixed carbon dioxide=9.5-25.75 p.p.m. (Fig. 58).

Flambeau Drainage: Mann L.; Nelson Lake.

Tomahawk Drainage: Stream, 10 mi. southwest of Hazelhurst.

Wisconsin Drainage: Pools along Wisconsin River, 4 mi. northeast

of Tomahawk Lake (Baker); Rice Creek, near Plum Lake.

Morrison — Mollusca of Northeastern Wisconsin . 377

Physa gyrina elliptica Lea.

pH—7.64; fixed carbon dioxide=18.87 p.p.m. (Fig. 59).

Flambeau Drainage: Trout Lake.

Physa integra Haldeman.

pH=8.0; fixed carbon dioxide— 24.73 p.p.m. (Fig. 60).

Lake Superior Drainage: South Branch, Presque Isle River, at Wine-

gar.

Physa michiganensis Clench.

pH—8.02; fixed carbon dioxide=23.0 p.p.m. (Fig. 61).

Flambeau Drainage: Outlet of Big Lake.

Genus Aplexa Fleming.

Aplexa hypnorum (L.).

No chemical data.

Wisconsin Drainage: Pools in swamp along Wisconsin River, 4 mi.

northeast of Tomahawk Lake (Baker).

Class PELECYPODA Goldfuss.

Order Prionodesmacea Dali.

Superfamily Naiadacea Menke.

Family Unionidae (d’Orbigny) Ortmann.

Subfamily UNIONINAE (Swainson) Ortmann.

Genus Fusconaia Simpson.

Fusconaia flava (Rafinesque) .

pH— 7.1-8.02; fixed carbon dioxide=12.07-23.0 p.p.m. (Fig. 62).

Flambeau Drainage: Inlet of White Sand L. ; Manitowish River, at

Boulder Junction, and 4 mi. southwest; Outlet of Big L.; South

Fork, Flambeau River, at Fifield, and 2 mi. east; Turtle L. ; Turtle

River, below Lake of the Falls.

Tomahawk Drainage: Tomahawk River, 4 mi. west of Minocqua.

Wisconsin Drainage: Clear Water Lake (Chadwick, Baker); St.

Germaine River; Wisconsin River, at Lac Vieux Desert, and 5 mi.

below.

Genus Amblema Rafinesque.

Amblema costata Rafinesque.

pH=7.1-7.7; fixed carbon dioxide=12.07-18.87 p.p.m. (Fig. 63).

Flambeau Drainage: Manitowish River, at Boulder Junction; Trout

River, at Trout L.; Turtle River, at Winchester, and below Lake

of the Falls.

Wisconsin Drainage: Clear Water Lake (Chadwick, Baker); Wis¬

consin River, 4 mi. northeast of Tomahawk Lake (Baker) ; Wis¬

consin River, 5 mi. below Lac Vieux Desert, and at Otter Rapids,

5 mi. west of Eagle River.

378 Wisconsin Academy of Sciences , Arts , and Letters.

Genus Pleurobema (Rafinesque) Agassiz.

Fleur obema coccineum (Conrad).

pH=7.15-7.63; fixed carbon dioxide=12.07-20.1 p.p.m. (Fig. 64).

Flambeau Drainage: Manitowish River, at Boulder Junction; Turtle

River, at Winchester.

Tomahawk Drainage: Tomahawk River, 4 mi. west of Minocqua.

Wisconsin Drainage: Wisconsin River, 5 mi. below Lac Yieux Des¬

ert.

Genus Elliptic Rafinesque.

Elliptio dilatatus (Rafinesque).

pH=7.3-7.5; fixed carbon dioxide=13. 3-14.0 p.p.m. (Fig. 65).

Flambeau Drainage: Manitowish River, at Boulder Junction.

Wisconsin Drainage: Wisconsin River, 5 mi. below Lac Vieux Des¬

ert.

Elliptio dilatatus delicatus (Simpson).

pH=7.1-8.02; fixed carbon dioxide=12. 07-23.0 p.p.m. (Fig. 66).

Flambeau Drainage: Manitowish River, 4 mi. southwest of Boulder

Junction; Outlet of Big L.; South Fork, Flambeau River, at Fifield,

and 2 mi. east; Turtle River, at Winchester, and below Lake of

the Falls.

Tomahawk Drainage: Tomahawk River, 4 mi. west of Minocqua.

Elliptio dilatatus sterkii Grier.

pH=7.15; fixed carbon dioxide=12.07 p.p.m. (Fig. 67).

Flambeau Drainage: Turtle Lake.

Subfamily Anodontinae Ortmann.

Genus Lasmigona Rafinesque.

Lasmigona compressa (Lea).

pH=7.1-8.02; fixed carbon dioxide— 12.07-24.73 p.p.m. (Fig. 68).

Lake Superior Drainage: Montreal River, at Pine L.; Ontonagon

River, Mich., 3 mi. north of Tenderfoot L.; South Branch, Presque

Isle River, at Winegar.

Flambeau Drainage: Inlet of White Sand L.; Manitowish River, 4

mi. southwest of Boulder Junction; Outlet of Big L.; South Fork,

Flambeau River, at Fifield; Trout River, at Trout L.; Turtle L.;

Turtle River, at Winchester.

Tomahawk Drainage: Tomahawk River, 4 mi. west of Minocqua.

Wisconsin Drainage: Gilmore Creek (Baker); Wisconsin River, at

Lac Vieux Desert, and 5 mi. below.

Lasmigona costata (Rafinesque).

pH=7.1-8.14; fixed carbon dioxide— 12.07-23.0 p.p.m. (Fig. 69).

Flambeau Drainage: Inlet of Trout L.; Inlet of White Sand L.;

Outlet of Big L.; Manitowish River, at Boulder Junction, and 4

Morrison — Mollusca of Northeastern Wisconsin . 379

mi. southwest; South Fork, Flambeau River, at Fifield, and 2 mi.

east; Trout River, at Trout L.; Turtle River, at Winchester, and

below Lake of the Falls.

Tomahawk Drainage: Tomahawk River, 4 mi. west of Minocqua.

Wisconsin Drainage: Gilmore Creek (Baker); Little St. Germaine

River; Plum Creek; St. Germaine River; Wisconsin River, 5 mi.

below Lac Yieux Desert, and at Otter Rapids, 5 mi. west of Eagle

River.

Lasmigona complanata (Barnes).

pH=7.3-8.14; fixed carbon dioxide=13.4-16.95 p.p.m. (Fig. 70).

Wisconsin Drainage: Little St. Germaine River; Plum L.; St. Ger¬

maine River; Wisconsin River, at Lac Vieux Desert, 5 mi. below

Lac Vieux Desert, and at Otter Rapids, 5 mi. west of Eagle River.

Genus Anodonta Lamarck.

Anodonta grandis plana Lea.

pH = 6.9-8.37; fixed carbon dioxide = 9.3-25.75 p.p.m. (Fig. 71).

Lake Superior Drainage: Montreal River, at Pine L.; South Branch,

Presque Isle River, at Winegar.

Flambeau Drainage: Inlet and outlet of Big L.; Inlet of White

Sand L.; Little Rice L.; Manitowish River, at Boulder Junction;

Outlet of Irving L. ; Outlet of Mann L. ; Outlet of Tamarac L. ;

Trout River, at Trout Lake; Turtle River, at Winchester, and be¬

low Lake of the Falls.

Tomahawk Drainage: Stream at State Fish Hatchery, near Wood¬

ruff.

Wisconsin Drainage: Gilmore Creek (Baker) ; Deerskin River, 6 mi.

south of Phelps; Plum Creek; St. Germaine River; Wisconsin Riv¬

er, at Lac Vieux Desert, 5 mi. below Lac Vieux Desert, and at Ot¬

ter Rapids, 5 mi. west of Eagle River.

Anodonta grandis footiana Lea.

pH=6.7-8.02; fixed carbon dioxide=3.2-30,56 p.p.m. (Fig. 72).

Lake Superior Drainage: Presque Isle Lake.

Flambeau Drainage: Adelaide L.; Big L.; Fishtrap L.; Little Long

L.; Lost Canoe L.; Trout L.; Turtle L.; Whitefish L.; Wildcat

Lake.

Tomahawk Drainage: Tomahawk Lake (Baker); Brandy L.; John¬

son L.; Little Star Lake.

Wisconsin Drainage: Found Lake (Cahn, Baker); Plum Lake.

Anodonta kennicottii Lea.

pH—7.85-8.0; fixed carbon dioxide=15.46-24.73 p.p.m. (Fig. 73).

Lake Superior Drainage: Palmer L:; Presque Isle Lake.

Flambeau Drainage: High L.; Silver L.; Trout Lake.

Anodonta marginata Say.

pH=6.03-8.37; fixed carbon dioxide— 2.6-30.56 p.p.m. (Fig. 74).

Lake Superior Drainage: Anna L.; Armour L.; Horsehead L.; Mon-

380 Wisconsin Academy of Sciences , Arts , and Letters .

treal River, at Pine L.; Ontonagon River, 3 mi. north of Tender¬

foot L.; Presque Isle Lake.

Green Bay Drainage: Butternut L.; Kentuck Lake.

Flambeau Drainage: Adelaide L.; Allequash L.; Big Muskellunge

L.; Big L. Outlet; Big L.; Cranberry L.; Favil L.; Fishtrap L.;

High L.; Inlet of Trout L. ; Inlet of White Sand L. ; Irving L. Out¬

let; L. Constance; L. George; L. Laura; Little Long L.; Little

Rice L.; Little White Birch L.; Lost Canoe L.; Manitowish River,

at Boulder Junction, and 4 mi. southwest; Mann L.; Marion L.;

Outlet of Mann L.; Outlet of Nixon L.; Outlet of Tamarac L.;

Silver L.; Trout L.; Trout River, at Trout L.; Turtle L.; Turtle

River, at Winchester; Wildcat Lake.

Tomahawk Drainage: Tomahawk Lake (Baker); Brandy L.; Clear

L.; Johnson L.; Stream at State Fish Hatchery, near Woodruff.

Wisconsin Drainage: Gilmore Creek (Baker); Crescent L.; Deer¬

skin River, 6 mi. south of Phelps; Little St. Germaine River; Plum

L. ; St. Germaine River; Razorback L.; Star L.; Wisconsin River,

at Lac Vieux Desert.

Genus Utterbackia F. C. Baker.

Utterbackia imbecillis (Say).

pH=7.1; fixed carbon dioxide=17.3 p.p.m. (Fig. 75).

Flambeau Drainage: Inlet of White Sand L.; Manitowish River, at

Boulder Junction; Turtle River, below Lake of the Falls.

Genus Anodontoides Simpson.

Anodontoides ferussacianus (Lea).

pH=7.0; fixed carbon dioxide=9.3 p.p.m. (Fig. 76).

Flambeau Drainage: Little Rice Lake.

Anodontoides ferussacianus subcylindraceus (Lea).

pH=6.9-8.37; fixed carbon dioxide=10. 65-30. 56 p.p.m. (Fig. 77).

Lake Superior Drainage: Montreal River, at Pine Lake.

Flambeau Drainage: Fishtrap L.; High L.; Inlet of White Sand

L.; Irving L. Outlet; Manitowish River, 4 mi. southwest of Boulder

Junction; Mann L.; Silver L.; Trout River, at Trout L.; Turtle

River, at Winchester; Whitefish L.; Wildcat Lake.

Tomahawk Drainage: Brandy L.; Tomahawk River, 4 mi. west of

Minocqua.

Wisconsin Drainage: Deerskin River, 6 mi. south of Phelps; Plum

L. ; Wisconsin River, at Lac Vieux Desert, and 5 mi. below.

Anodontoides birgei F. C. Baker.

pH=8.0; fixed carbon dioxide=24.73 p.p.m. (Fig. 78).

Lake Superior Drainage: South Branch, Presque Isle River, at

Winegar.

Morrison — Mollusca of Northeastern Wisconsin . 881

Genus Alasmidonta Say.

Alasmidonta marginata variahilis F. C. Baker.

pH=7.1-8.14; fixed carbon dioxide=13.3-20.1 p.p.m. (Fig. 79).

Flambeau Drainage: Manitowish River, 4 mi. southwest of Boulder

Junction; South Fork, Flambeau River, at Fifield, and 2 mi. east.

Tomahawk Drainage: Tomahawk River, 4 mi. west of Minocqua.

Wisconsin Drainage: Little St. Germaine River; Wisconsin River,

at Otter Rapids, 5 mi. west of Eagle River.

Genus Strophitus Rafinesque.

Strophitus rugosus pavonius (Lea).

pH=7.1-8.14; fixed carbon dioxide— 12.07-23.0 p.p.m. (Fig. 80).

Flambeau Drainage: Big L. Outlet; Inlet of Trout Lake; Inlet of

White Sand L.; Manitowish River, 4 mi. southwest of Boulder

Junction; South Fork, Flambeau River, at Fifield; Trout River, at

Trout L.; Turtle River, at Winchester, and Below Lake of the

Falls.

Tomahawk Drainage: Tomahawk River, 4 mi. west of Minocqua.

Wisconsin Drainage: Gilmore Creek (Baker); Little St. Germaine

River; Plum Creek; St. Germaine River; Wisconsin River, at Lac

Vieux Desert.

Subfamily Lampsilinae Ortmann.

Genus Actinonaias Fischer & Crosse.

Actinonaias carinata (Barnes).

pH—7.0-8.14; fixed carbon dioxide=12.07-23.0 p.p.m. (Fig. 81).

Flambeau Drainage: Inlet of Trout L.; Outlet of Big L.; South

Fork, Flambeau River, at Fifield; Turtle River, at Winchester, and

below Lake of the Falls.

Tomahawk Drainage: Tomahawk River, 4 mi. west of Minocqua.

Wisconsin Drainage: Clear Water Lake Creek (Chadwick, Baker);

Gilmore Creek, and Wisconsin River, 4 mi. northeast of Tomahawk

Lake (Baker); Little St. Germaine River; St. Germaine River;

Wisconsin River, at Lac Vieux Desert, 5 mi. below Lac Vieux Des¬

ert, at Otter Rapids, 5 mi. west of Eagle River, and at Rainbow

Rapids, southeast of Lake Tomahawk.

Genus Ligumia Swainson.

Ligumia recta (Lamarck).

pH~7.15; fixed carbon dioxide=12.07 p.p.m. (Fig. 82).

Flambeau Drainage: Turtle Lake.

Ligumia recta latissima (Rafinesque).

pH=7.1-8.14; fixed carbon dioxide=12.07-20.1 p.p.m. (Fig. 83).

Flambeau Drainage: Manitowish River, at Boulder Junction, and 4

mi. southwest; South Fork, Flambeau River, at Fifield, and 2 mi.

382 Wisconsin Academy of Sciences, Arts, and Letters .

east; Turtle River, at Winchester, and below Lake of the Falls.

Tomahawk Drainage: Tomahawk River, 4 mi. west of Minocqua.

Wisconsin Drainage: Wisconsin River, 4 mi. northeast of Toma¬

hawk Lake (Baker); Little St. Germaine River; Wisconsin River,

at Otter Rapids, 5 mi. west of Eagle River.

Genus Lampsilis Rafinesque.

Lampsilis siliquoidea (Barnes).

pH=6.9-8.14; fixed carbon dioxide— 9.3-24.73 p.p.m. (Fig. 84).

Lake Superior Drainage: Ontonagon River, Mich., 3 mi. north of

Tenderfoot L. ; South Branch, Presque Isle River, at Winegar.

Flambeau Drainage: Inlet and Outlet of Big L.; Inlet of Trout L.;

Little Rice L. ; Manitowish River, at Boulder Junction, and 4 mi.

southwest; Outlet of Tamarac L.; South Fork, Flambeau River,

at Fifield, and 2 mi. east; Trout River, at Trout L.; Turtle River,

at Winchester, and below Lake of the Falls.

Tomahawk Drainage: Stream at State Fish Hatchery, near Wood¬

ruff.

Wisconsin Drainage: Clear Water Creek (Chadwick, Baker); Gil¬

more Creek, and Wisconsin River, 4 mi. northeast of Tomahawk

Lake (Baker); Deerskin River, 6 mi. south of Phelps; Little St.

Germaine River; St. Germaine River; Wisconsin River, at Lac

Vieux Desert, 5 mi. below Lac Vieux Desert, at Otter Rapids, 5 mi.

west of Eagle River, and at Rainbow Rapids, southeast of Lake

Tomahawk.

Lampsilis siliquoidea rosacea (DeKay).

pH=6.95-8.37; fixed carbon dioxide=7.5-30.56 p.p.m. (Fig. 85).

Lake Superior Drainage: Armour L.; Horsehead L.; Presque Isle

Lake.

Flambeau Drainage : Allequash L. ; Big L. ; Boulder L. ; Fishtrap L. ;

High L.; Mann L.; Trout L.; Turtle L.; Whitefish L.; White Sand

L.; Wildcat Lake.

Tomahawk Drainage: Tomahawk Lake (Baker); Brandy Lake.

Wisconsin Drainage: Plum Lake; Plum Creek.

Lampsilis ventricosa occidens (Lea).

pH—7.0-8.14; fixed carbon dioxide=12. 07-23,0 p.p.m. (Fig. 86).

Flambeau Drainage: Inlet and Outlet of Big L.; Manitowish Riv¬

er, at Boulder Junction, and 4 mi. southwest; South Fork, Flam¬

beau River, at Fifield, and 2 mi. east; Trout River, at Trout L.;

Turtle River, at Winchester, and below Lake of the Falls.

Tomahawk Drainage: Tomahawk River, 4 mi. west of Minocqua.

Wisconsin Drainage: Clear Water Creek (Chadwick, Baker); Gil¬

more Creek, and Wisconsin River, 4 mi. northeast of Lake Toma¬

hawk (Baker) ; Little St. Germaine River; St. Germaine River;

Wisconsin River, at Lac Vieux Desert, 5 mi. below Lac Vieux Des¬

ert, at Otter Rapids, 5 mi. west of Eagle River, and at Rainbow

Rapids, southeast of Lake Tomahawk.

M orris on— M ollusca of Northeastern Wisconsin . 383

Lampsilis ventricosa lurida Simpson*

pH=7.15-8.02; fixed carbon dioxide— 12.07-23.0 p.p.m. (Fig. 87).

Flambeau Drainage: Big L.; Fishtrap L.; High L.; Inlet of Trout

L.; Trout Lake.

Order Teleodesmacea Dali.

Superfamily Cyrenacea Tryon.

Family Sphaeriidae Dali.

Subfamily Sphaeriinae F. C. Baker.

Genus Sphaerium Scopoli.

Sphaerium sulcatum (Lamarck).

pH=6.9~8.37; fixed carbon dioxide=9.3-25.75 p.p.m. (Fig. 88).

Flambeau Drainage: Big L.; Big Muskellunge L.; Fishtrap L.; Irv¬

ing L. Outlet; Little Rice L.; Outlet of Mann L.; Outlet of Nixon

L. ; Trout L. Inlet.

Tomahawk Drainage: Tomahawk Lake (Baker).

Wisconsin Drainage: Deerskin River, 6 mi. south of Phelps; Plum

L. ; Rice Creek, near Plum Lake.

Sphaerium eras sum Sterki.

pH—7.1; fixed carbon dioxide=17.3 (Fig. 89).

Flambeau Drainage: Turtle River, below Lake of the Falls.

Wisconsin Drainage: Wisconsin River, at Otter Rapids, 5 mi. west

of Eagle River.

Sphaerium fallax Sterki.

pH=6.85-8.37; fixed carbon dioxide— 11.75-30.56 p.p.m. (Fig. 90).

Lake Superior Drainage: Ontonagon River, Mich., 3 mi. north of

Tenderfoot L.; Palmer L.; Presque Isle Lake.

Flambeau Drainage : High L. ; Island L. ; Marion L. ; Outlet of Mann

L.; Outlet of Tamarac L.; Turtle River, at Winchester; Wildcat

Lake.

Wisconsin Drainage: Wisconsin River, at Lac Vieux Desert.

Sphaerium solidulum (Prime).

pH—7.7; fixed carbon dioxide— -16.95 p.p.m. (Fig. 91).

Wisconsin Drainage: Plum Creek.

Sphaerium stamineum (Conrad)

pH—6.9-8.37; fixed carbon dioxide— 13.0-25.75 p.p.m. (Fig. 92).

Lake Superior Drainage: Ontonagon River, Mich., 3 mi. north of

Tenderfoot L.; South Branch, Presque Isle River, at Winegar.

Flambeau Drainage: Inlet of Trout L. ; Manitowish River, at Boul¬

der Junction; Mann L. Outlet; Outlet of Big L.; Trout River, at

Trout Lake.

Tomahawk Drainage: Tomahawk River, 4 mi. west of Minocqua.

Wisconsin Drainage: Deerskin River, 6 mi. south of Phelps; Wiscon-

384 Wisconsin Academy of Sciences, Arts, and Letters.

sin River, 5 mi. below Lac Vieux Desert, at Otter Rapids, 5 mi.

west of Eagle River, and at Rainbow Rapids, southeast of Lake

Tomahawk.

Sphaerium emarginatum (Prime).

pH=7.1-7.95; fixed carbon dioxide=15.5-17.3 p.p.m. (Fig. 93).

Flambeau Drainage: Inlet of Trout L.; Inlet of White Sand L.;

Manitowish River, at Boulder Junction; South Fork, Flambeau

River, at Fifield; Turtle River, below Lake of the Falls.

Sphaerium bakeri Sterki.

pH=7.7; fixed carbon dioxide— 16.95 p.p.m. (Fig. 94).

Wisconsin Drainage: Plum Creek.

Sphaerium striatinum (Lamarck).

pH=7.1; fixed carbon dioxide=17.3 p.p.m. (Fig. 95).

Flambeau Drainage: Turtle River, below Lake of the Falls.

Wisconsin Drainage: Wisconsin River, 4 mi. northeast of Toma¬

hawk Lake (Baker).

Sphaerium rhomb oideum (Say).

pH=7.1-7.36; fixed carbon dioxide=14.0-18.5 p.p.m. (Fig. 96).

Lake Superior Drainage: Ontonagon River, Mich., 3 mi. north of

Tenderfoot Lake.

Flambeau Drainage: Fishtrap L.; Outlet of Nixon Lake.

Sphaerium occidentale Prime.

pH=5.8-5.9; fixed carbon dioxide=5.5-7.5 p.p.m. (Fig. 97).

Green Bay Drainage: Pools in lumber slashings, 4 mi. east of But¬

ternut Lake.

Flambeau Drainage: Forest Ponds, 10 mi. northeast of Boulder

Junction.

Wisconsin Drainage: Swamp along Wisconsin River, 4 mi. north¬

east of Tomahawk Lake (Baker).

Genus Musculium Link.

Musculium jayense (Prime).

pH=7. 1-7.23 ; fixed carbon dioxide=10.8-13.0 p.p.m. (Fig. 98).

Flambeau Drainage: L. Laura; Outlet of Tamarac Lake.

Musculium par tumeium (Say).

No chemical data.

Wisconsin Drainage: Small Ponds in swamp along Wisconsin River,

4 mi. northeast of Tomahawk Lake (Baker).

Musculium truncatum (Linsley).

pH=6.05-8.37; fixed carbon dioxide— 2.75-25.75 p.p.m. (Fig. 99).

Flambeau Drainage: Catfish L.; Fishtrap L.; Harvey L.; L. Laura;

Mary L.; Outlet of Mann Lake.

Tomahawk Drainage: Little Rice River.

Morrison — Mollusca of Northeastern Wisconsin. 885

Musculium rosaceum (Prime).

pH—6.4-7.64; fixed carbon dioxide=9.3-18.87 p.p.m. (Fig. 100).

Flambeau Drainage: Big Muskellunge L.; Little Rice L.; Outlet of

Nixon L.; Trout Lake.

Tomahawk Drainage: Pond, near State Fish Hatchery ponds, near

Woodruff.

Musculium ryckholti (Normand).

No chemical data.

Tomahawk Drainage: Small Kettle-hole Pools near Tomahawk Lake

(Baker).

Musculium securis (Prime).

pH=5.9-8.37; fixed carbon dioxide=2.75-25.75 p.p.m. (Fig. 101).

Lake Superior Drainage: Black Oak Lake.

Flambeau Drainage: Allequash L.; Forest ponds, 10 mi. northeast

of Boulder Junction; Helen L.; Little Long L.; Mary L.; Outlet

of Mann L.; Pond along Mann L. Outlet.

Tomahawk Drainage: Pond near Tomahawk Lake, and Tomahawk

Lake (Baker); Pond, near State Fish Hatchery ponds, at Wood¬

ruff.

Wisconsin Drainage: Wisconsin River, at Rainbow Rapids, southeast

of Lake Tomahawk.

Musculium steinii (A. Schmidt).

pH=6.6; fixed carbon dioxide— 12.9 p.p.m. (Fig. 102).

Flambeau Drainage: Inlet of White Sand Lake.

Subfamily Pisidiinae F. C. Baker.

Genus Pisidium C. Pfeiffer.

Pisidium virginicum (Gmelin).

pH=7.0~7.7; fixed carbon dioxide=13.0-16.95 p.p.m. (Fig. 103).

Wisconsin Drainage: Wisconsin River, 4 mi. northeast of Tomahawk

Lake (Baker); Plum L.; Wisconsin River, at Rainbow Rapids,

southeast of Lake Tomahawk, and Vz mi. below.

Pisidium idahoense Roper.

pH — 5.8; fixed carbon dioxide = 1.5 p.p.m. (Fig. 104).

Tomahawk Drainage: Walker Lake.

Pisidium compressum Prime.

pH=7.0-8.37; fixed carbon dioxide— 9.3-30.56 p.p.m. (Fig. 105).

Lake Superior Drainage: Palmer L.; Presque Isle L.; South Branch,

Presque Isle River, at Winegar.

Flambeau Drainage: Big L. Outlet; Big Muskellunge L.; Boulder

L. ; Inlet of Trout L. ; Irving L. Outlet ; Little Rice L. ; Little White

Birch L. ; Lost Canoe L. ; Mann L. Outlet; Trout L. ; Upper

Gresham L.; Whitefish L.; White Sand L.; Wildcat Lake.

Tomahawk Drainage: Brandy L.; Kawaguesaga Lake.

386 Wisconsin Academy of Sciences, Arts, and Letters .

Wisconsin Drainage: Little St. Germaine River; Plum L. ; Star L.;

Wisconsin River, at Lac Vieux Desert, at Rainbow Rapids, south¬

east of Lake Tomahawk, and V2 mi. below.

Pisidium fallax septentrionale Sterki.

pH=7.95; fixed carbon dioxide=16.6 p.p.m. (Fig. 106).

Flambeau Drainage: Inlet of Trout L.; Inlet of White Sand Lake.

Pisidium punctatum Sterki.

pH=7.0; fixed carbon dioxide— 13.0 p.p.m. (Fig. 107).

Wisconsin Drainage: Wisconsin River, at Rainbow Rapids, south¬

east of Lake Tomahawk.

Pisidium variabile Prime.

pH=5.72-8.37; fixed carbon dioxide=l. 72-30.56 p.p.m. (Fig. 108).

Lake Superior Drainage: Ontonagon River, Mich., 3 mi. north of

Tenderfoot L.; Palmer L.; Presque Isle Lake.

Flambeau Drainage: Big Muskellunge L.; Boulder L.; Clear Crooked

L.; Dead Pike L.; Fishtrap L.; Little Rice L.; Mann L.; Outlet of

Mann L.; Outlet of Nixon L. ; Outlet of Tamarac L. ; Pauto L.;

Trout L.; Upper Gresham L.; Whitefish L.; White Sand L.; Wild¬

cat Lake.

Tomahawk Drainage: Brandy L. ; Kawaguesaga L.; Trilby Lake.

Wisconsin Drainage: Crescent L.; Plum L.; Razorback L.; Star L.;

Wisconsin River, at Rainbow Rapids, southeast of Lake Tomahawk.

Pisidium minus culum Sterki.

pH=7.48-7.64; fixed carbon dioxide=12.96-18.87 p.p.m. (Fig. 109).

Flambeau Drainage: Little White Birch L.; Trout Lake.

Pisidium adamsi Prime.

pH=6.05-7.7; fixed carbon dioxide=2.75-18.36 p.p.m. (Fig. 110).

Flambeau Drainage: Fishtrap L.; Irving L. Outlet; Mary Lake.

Wisconsin Drainage: Plum Lake.

Pisidium sargenti Sterki.

pH—6.05-8.14; fixed carbon dioxide=2.75-23.0 p.p.m. (Fig. 111).

Lake Superior Drainage: Palmer Lake.

Flambeau Drainage: Big L. Outlet; Little White Birch L.; Mani-

towish River, at Boulder Junction; Mary L.; Trout L.; Trout River,

at Trout L.; Whitefish Lake.

Tomahawk Drainage: Clear Lake.

Wisconsin Drainage: Crescent L.; Little St. Germaine River; Plum

L.; Star L. ; Wisconsin River at Lac Vieux Desert, and at Rain¬

bow Rapids, southeast of Lake Tomahawk.

Pisidium neglectum Sterki.

pH=6.66-7.1; fixed carbon dioxide—2. 9-14.0 p.p.m. (Fig. 112).

Flambeau Drainage: Outlet of Nixon Lake.

Tomahawk Drainage: Clear Lake.

Morrison — Mollusca of Northeastern Wisconsin. 387

Pisidium lilljehorgi Clessin. ( —scutellatum Sterki.)

pH=6.16-8.02; fixed carbon dioxide— 1.97-23.0 p.p.m. (Fig. 113).

Lake Superior Drainage: Katinka Lake.

Flambeau Drainage: Big L. Outlet; Boulder L.; Little White Birch

L.; Trout L.; Whitefish Lake.

Tomahawk Drainage: Little Arbor Vitae Lake (Winslow, Baker).

Wisconsin Drainage: Crescent L.; Plum L.; Star Lake.

Pisidium lilljehorgi cristatum Sterki.

pH—7.35-7.64; fixed carbon dioxide— 18.87-19.5 p.p.m. (Fig. 114).

Lake Superior Drainage: Palmer Lake.

Flambeau Drainage: Trout Lake.

Pisidium roperi Sterki.

pH=5.8-6.4; fixed carbon dioxide— 5.S-9.5 p.p.m. (Fig. 115).

Green Bay Drainage: Pools in lumber slashings, 4 mi. east of But¬

ternut Lake.

Flambeau Drainage : Forest ponds, 10 mi. northeast of Boulder Junc¬

tion.

Tomahawk Drainage : Kettle-hole pools near Tomahawk Lake

(Baker) ; Pond, near State Fish Hatchery ponds, near Woodruff.

Pisidium strengi Sterki.

pH=5. 84-7.95; fixed carbon dioxide— 2.13-30.56 p.p.m. (Fig. 116).

Flambeau Drainage: Wildcat Lake.

Tomahawk Drainage: Trilby Lake.

Wisconsin Drainage: Finley Lake.

Pisidium abditum Haldeman.

pH=7.6; fixed carbon dioxide=16.7 p.p.m. (Fig. 117).

Tomahawk Drainage: Tomahawk Lake and kettle-hole pools in vi¬

cinity (Baker).

Pisidium subrotundatum Sterki.

No chemical data.

Wisconsin Drainage: Wisconsin River, swampy places, 4 mi. north¬

east of Tomahawk Lake (Baker).

Pisidium splendidulum Sterki.

pH=6.32; fixed carbon dioxide=1.98 p.p.m. (Fig. 118).

Wisconsin Drainage: Sterrett Lake.

Pisidium levissimum Sterki.

pH—7.64 ; fixed carbon dioxide— 18.87 p.p.m. (Fig. 119).

Flambeau Drainage: Trout Lake.

Pisidium pauper culum Sterki.

pH=7. 0-8.0 ; fixed carbon dioxide— 9.3-24.73 p.p.m. (Fig. 120).

Lake Superior Drainage: Palmer L.; Presque Isle Lake.

Flambeau Drainage: Big Muskellunge L.; Boulder L.; L. Laura;

Little Rice L.; Trout Lake.

388 Wisconsin Academy of Sciences , Arts and Letters .

Tomahawk Drainage: Brandy Lake.

Wisconsin Drainage: Crescent L.; Plum L.; Star L.; Wisconsin

River, at Rainbow Rapids, southeast of Lake Tomahawk.

Pisidium rotundatum Prime.

p 11=5.8-6.2; fixed carbon dioxide=l. 97-9.0 p.p.m. (Fig. 121).

Lake Superior Drainage: Katinka Lake.

Green Bay Drainage: Pools in lumber slashings, 4 mi. east of But¬

ternut Lake.

Flambeau Drainage: Forest pond, 10 mi. northeast of Boulder Junc¬

tion.

Pisidium vesiculare Sterki.

pH=7.64; fixed carbon dioxide=18.87 p.p.m. (Fig. 122).

Flambeau Drainage: Trout Lake.

Pisidium ferrugineum Prime.

pH=7.23-8.14; fixed carbon dioxide~10.8-22.5 p.p.m. (Fig. 123).

Flambeau Drainage: L. Laura; Trout Lake.

Tomahawk Drainage: Little Arbor Vitae Lake (Winslow, Baker).

Wisconsin Drainage: Crescent L.; Little St. Germaine River; Star

Lake.

Pisidium concinnulum Sterki.

pH—5.72-7.48; fixed carbon dioxide— 1.72-15-46 p.p.m. (Fig. 124).

Green Bay Drainage: Pools in lumber slashings, 4 mi. east of But¬

ternut Lake.

Flambeau Drainage : Forest ponds, 10 mi. northeast of Boulder Junc¬

tion ; Pauto L. ; Silver L. ; Springs in Tamarack bog north of Trout

Lake.

Pisidium pusillum (Gmelin) Jenyus.

No chemical data.

Flambeau Drainage: Pool along Mann Lake Outlet.

Bibliography

Baker, F. C. 1911. The molluscan fauna of Tomahawk Lake, Wis.

Trans. Wis. Acad. Sci., Arts, Lett., XVII, pp. 200-246, pi. xi-xvii.

1928a. The Fresh Water Mollusca of Wisconsin. Part 1, Gastropoda.

(Bull. 70, part 1. Wis. Geol. & Nat. Hist. Sur.) Mon. Aquatic Gastro¬

poda of Wisconsin. Wis. Acad. Sci. Arts and Lett.

1928b. The Fresh Water Mollusca of Wisconsin. Part 2, Pelecypoda.

(Bull. 70, part 2, Wis. Geol. & Nat. Hist. Sur.) Bull. U. of Wis. Serial

No. 1527, general series, No. 1301.

Chadwick, G. H. 1905. List of Wisconsin shells. Nautilus, XIX, pp. 57-60.

Winslow, M. L. 1926. The varieties of Planorbis campanulatus Say. Occ.

Papers, Mus. Zool. Univ. Mich., No. 180, pp. 1-9, pi. i, ii.

Morrison — Mollusca of Northeastern Wisconsin. 389

Figs. 1-16. The pH and fixed carbon dioxide ranges of various mol-

lusks. Ordinates indicate fixed carbon dioxide in parts per million; ab¬

scissae indicate pH. The circumscribed area presents for comparison the

total range of these factors in all the lakes of the Highland Lake District

for which data are available.

390 Wisconsin Academy of Sciences , Arts , and Letters .

Fig. 17-32. The pH and fixed carbon dioxide ranges of various mol-

lusks. Ordinates indicate fixed carbon dioxide in parts per million; ab¬

scissae indicate pH. The circumscribed area presents for comparison the

total range of these factors in all the lakes of the Highland Lake Dis¬

trict for which data are available.

Morrison — Mollusca of Northeastern Wisconsin. 391

Figs. 33-48. The pH and fixed carbon dioxide ranges of various mol-

/usks. Ordinates indicate fixed carbon dioxide in parts per million; ab¬

scissae indicate pH. The circumscribed area presents for camparison the

total range of these factors in all the lakes of the Highland Lake Dis¬

trict for which data are available.

392 Wisconsin Academy of Sciences , Arts , and Letters .

Figs. 49-64. The pH and fixed carbon dioxide ranges of various mol-

lusks. Ordinates indicate fixed carbon dioxide in parts per million; ab¬

scissae indicate pH. The circumscribed area presents for comparison the

total range of these factors in all the lakes of the Highland Lake Dis¬

trict for which data are available.

Morrison — Mollusca of Northeastern Wisconsin . 393

Pigs. 65-80. The pH and fixed carbon dioxide ranges of various mol-

lusks. Ordinates indicate fixed carbon dioxide in parts per million; ab¬

scissae indicate pH. The circumscribed area presents for comparison the

total range of these factors in all the lakes of the Highland Lake Dis¬

trict for which data are available.

394 Wisconsin Academy of Sciences , Arts, and Letters.

Figs. 81-96. The pH and fixed carbon dioxide ranges of various mol-

lusks. Ordinates indicate fixed carbon dioxide in parts per million; ab¬

scissae indicate pH. The circumscribed area presents for comparison the

total range of these factors in all the lakes of the Highland Lake Dis¬

trict for which data are available.

Morrison— Mollusca of Northeastern Wisconsin . 395

r.o 6.® TO' 8.0 5.0 6.o 7.0 8.0 £0 fe.e 7.0 fi.© £0 tJO 7.0 8j©

Figs. 97-112. The pH and fixed carbon dioxide ranges of various mol-

lusks. Ordinates indicate fixed carbon dioxide in parts per million; ab¬

scissae indicate pH. The circumscribed area presents for comparison the

total range of these factors in all the lakes of the Highland Lake Dis¬

trict for which data are available.

396 Wisconsin Academy of Sciences , Arts , and Letters .

f.o 6.6 l.o 8.0 5.0 6.0 7.6 fl.o 5.0 4o 7.0 8.0 5.0 6.0 7.0 8.6

Figs. 113-127. pH and fixed carbon dioxide ranges: (113-124) for

various mollusks, (125) for all the lakes of the Highland Lake District

for which records are available, (126) for all lakes in which mollusks

were found, and (127) for stream localities found to harbor mollusks.

Ordinates represent fixed carbon dioxide; abscissae indicate pH.

STUDIES ON THE LIFE HISTORY OF

ACELLA HALDEMANI (“DESH.” BINNEY.)

J. P. E. Morrison

Notes from the Limnological Laboratory of the Wisconsin Geological

and Natural History Survey

No. XLVIII.

Introduction

There are several problems that are outstanding in regard

to the snail Acella. Among these are the breeding season,

length of life, and whereabouts during most of the year. This

paper is written in answer to these questions.

This species, the slenderest of American Lymnaeas, has a lim¬

ited range, in the Great Lakes Drainage, from Vermont to

northern Minnesota, and from lower Canada to northern Illi¬

nois and northern Ohio. Described by Jay, from Lake Cham¬

plain in 1839, little was known of the ecology for many years.

Kirkland has contributed many interesting observations. Bak¬

er, in monographing the American Lymnaeas, has collected all

the available notes on Acella. All of the earlier collectors found

the species only in the fall of the year, and only in the adult

condition. One important reference in the literature has been

overlooked. This is the note of DeCamp’s collecton of Acella

(6) “in one year in May. He once told me he collected eighty-

five on the rushes, ‘where they had come to spawn’.” Just why

Kirkland should have forgotten this lead, is hard to under¬

stand. In later notes (1) he says, “This is a deep water spe¬

cies, which migrates shoreward in the fall, doubtless for spawn¬

ing purposes, as adults only have been captured, — ”.

Baker (3) after finding young individuals in July 1916, says,

“It may be that the animal descends to the Pond-weed Zones

in the winter and lays its eggs on the Potamogeton and that

they subsequently hatch out in the spring.”

In the course of studies on the mollusks of the Northeastern

Wisconsin Lakes, Acella was found. Additional collecting in

these lakes at various times of the year has resulted in tracing

the complete life history.

398 Wisconsin Academy of Sciences, Arts, and Letters .

The writer is indebted to the following people: Dr. F. C.

Baker, Univ. of Ill. Museum; Dr. W. J. Clench, M.C.Z. at Har¬

vard; Dr. Paul Bartsch, U. S. Nat. Museum; and Dr. W. G.

Van Name, Am. Mus. Nat. Hist.; for information regarding

Acella in the collections of these museums ; to W. A. Dence, N.

Y. State College of Forestry for loan of specimens; to Paul

Armand, Fishtrap Lake, Vilas Co. Wis. ; to G. E. Burdick, for

assistance in Photography and to Prof. Chancey Juday for

chemical data on the lakes and many helpful suggestions and

criticisms.

The ecological conditions under which Acella is found in Vilas

Co., Wisconsin, are restricted. The observed average pH of

the lakes it inhabits in this region is between 7.36 and 7.7.

The amount of fixed carbon dioxide present in these same lakes

is also within relatively narrow limits of variation.

Fishtrap Lake —pH 7.36 fixed C02 18.36 p.p.m.

Harris Lake —pH 7.7 fixed C02 17.0 p.p.m.

High Lake —pH 7.7 fixed C02 22.56 p.p.m.

The narrowness of the limits under which Acella has been found

in these lakes is made significant by the fact that the variation

in all the lakes of the district for which there are data is so

much greater. There are lakes here that have an average pH

of 4.4 and others as alkaline as 8.9. The range in hardness of

water of these lakes is indicated by the fixed carbon dioxide

content, which is as low as 0.2 in some lakes, and as high as

31.5 p.p.m. in others (Fig. 1).

minimum —pH 4.4 fixed C02 0.2 p.p.m.

maximum —pH 8.9 fixed C02 31.5 p.p.m.

It was at first thought that the species would be restricted to

a muddy type of bottom (silted), but further observations have

shown that the species is found in situations having both sand

and silt bottoms, though perhaps more often on the silted type.

This is in direct support of the observed preference for a more

or less protected situation.

As regards its niche in the habitat, it is primarily an inhabi¬

tant of vegetation. But what becomes of the snail, when the

winter conditions kill down the annual growths of such plants

as the reeds, pond lilies, etc.? Assuredly it must have some

other places for protection from being silted under on the bot-

Morrison — Life History of Acella,

399

Fig. 1. Distribution of lakes of the Northeastern Wisconsin District

in regard to hydrogen-ion concentration (pH) and hardness of water

(represented by fixed carbon dioxide in parts per million), with the re¬

stricted distribution of Acella indicated on the same scales.

400 Wisconsin Academy of Sciences, Arts, and Letters.

tom and for the procuring of food. It is pretty well established

that this species feeds on the small filamentous algae growing

on the plants on which it has been collected. Why should not

this snail be found on any other object in the habitat upon

which these plant foods are found? It stands as unquestioned

that the snail should be able to move on the bottom, in order

to travel from one plant to another, and the observations of the

species both in the juvenile and the adult stages, show that it

frequently moves about over the bottom.

The fallacy that has done most to create the mystery in re¬

gard to Acella, is the inference that it must migrate, since it

had been found in shallow water only in the adult stage. As

will be evident from observations to be mentioned later, this

erroneous idea must be given up ; there are, in fact, only occa¬

sional wanderings of individuals away from the places where

they feed to another plant or log.

As previously noted by Kirkland and Baker, this species is

more sluggish in its movements than others of the family. This

slowness of motion will account in part for the colonial habits,

since none of the individuals ever go very far from the spot

where they hatched from the egg, and near the same place

they will lay their eggs, thus keeping the species in the same

location from generation to generation.

The habitat of Acella has always been listed by previous

writers as vegetation. Decamp, Kirkland and others have re¬

corded it from rushes (Scirpus) ; Sargent (5) mentions having

collected it from the under side of pond lily leaves; Baker

(S) has recorded it from Oneida Lake, N. Y., from the follow¬

ing plants: “Smith’s bullrush (Scirpus smithii) on the stem.

Floating pond-weed (Potamogeton natans) on leaves and stem.

Pond-weed (P. interruptus) on leaves and stem. White water-

lily (Castalia odorata) on leaves and stem. Yellow water-lily

(Nymphaea advena) on leaves and stem.”

The writer has found the habitat of Acella even more vari¬

able, including objects other than vegetation. The eggs have

been noted on rushes, on the dead and decaying portions of

Potamogeton and burreed plants, and on the small sticks and

logs on the bottom, in the same zone of vegetation.

Juvenile individuals have been collected from Potamogeton,

from Yellow water-lily leaves, from burreed leaves and stems,

and from rushes ( Scirpus , 2 species), and from stumps and

Morrison — Life History of Acella .

401

snags in the same marginal zone. One stray individual was

taken from silt bottom in deeper water and another from gravel

bottom (at mouth of Dead River, Illinois). Adults have been

collected from rushes (both living and dead stems), from Po-

tamogeton, burreed, pickerel weed, yellow pond-lily stems, and

from snags and logs. This last mentioned habitat of snags,

etc., has been observed only in the spring season.

As to depth of water in which Acella is to be found, Kirk¬

land says, (1) “in water from one to three feet deep ; and in¬

variably from six to eight inches from the bottom, the apex

of the shell pointing downwards, — though in a few instances

the apex has been upwards, as if in the act of descending/’

Baker says it is found in water of about the same depth limits

(.3 to over 1 m. deep.). The writer has noted the following

depths: Dead River: most of the Acella were found near the

surface, in from 4 to 24 inches of water; Fishtrap Lake: the

snails were generally distributed, from near the bottom to near

the top, on all parts of the plants, and with the apex of the

shell pointing in any direction, in water 10 to 30 inches deep;

in the channel between Fishtrap and High Lakes: in 15-36

inches of water, just on the edge of the current, the snails

always within 18 inches of the surface; High Lake: near the

surface in water 10 to 18 inches deep; Harris Lake: in water

16-20 inches deep.

Three of the colony locations in Vilas County were examined

on February 1st and 2nd, 1931, to find out to what winter con¬

ditions these snails are subjected. At the site of the colony on

Fishtrap Lake, the following conditions were found: ice 9

inches thick, 2 inches slushy-snow ice, and about 10 inches

snow over all; the temperature of the water was 1.5 degrees

Centigrade; the depth of water was 24 inches. No specimens

were found at this time. The channel between Fishtrap and

High Lakes showed the following: ice to 3 inches thick;

snow covering the ice 8 to 10 inches thick; water about 10

inches deep; water temperature 1.5 degrees Centigrade. Two

adults were found on decaying burreed stems on this date. Har¬

ris Lake showed: 1 foot of ice, 2 inches soft snow-ice, and 2

to 4 inches of snow; water 2 feet 5 inches deep; water tem¬

perature near 0 degrees Centigrade, since it froze rapidly over

the holes cut in the ice, even at 1 p. m. No Acella found here,

402 Wisconsin Academy of Sciences , Arts, and Letters .

because the exact location of the colony was not found. These

measurements indicate conditions for the snails in a mild win¬

ter. In the case of the channel habitat, the extremely thin

ice indicates either that there are springs in this swamp along

the channel, or that the current is spread out more than in the

summer when the water is open.

Egg masses of these animals were seen on May 16, 1925

from Dead River, Illinois, on the stems of rushes. At this

time, the eggs had not hatched, but the embryonic shell was of

such size as to almost fill the egg capsule. These egg masses

were identified as belonging to Acella by the shape of the em¬

bryonic shell, which, as Baker (1) has said, is peculiar in that

it alone of all the slender Lymnaerds shows elongation of the

nuclear whorls. Again May 10, 1930, egg masses were found

sparingly on the stems, leaves, of Potamogeton and burreed

plants, in the colony previously discovered on the south side

of Fishtrap Lake, Vilas Co., Wisconsin. Further collections

have brought out the fact that in this northern Wisconsin re¬

gion, the period of egg laying extends from some time previ¬

ous to May 10 to about the middle of June. One egg mass was

collected on June 21, 1930. When examined, it was in the early

developmental stages.

The process of egg laying was observed on March 5, 1931.

One of the adult individuals in an aquarium in the laboratory

was seen laying eggs on the leaf of eel-grass (V allisneria) .

The process is as follows: The animal remains almost sta¬

tionary, with the long axis of the shell parallel to the leaf, and

pointing upward; first, the gelatinous covering of the eggmass

is laid down in part and then the long, oval egg capsules are

deposited inside this envelope; when all the eggs are laid, the

covering of the egg mass is completed and the animal moves

slowly away. As the eggs are being deposited, the snail moves

forward very slowly, just enough to make room for them. The

whole process of laying this egg mass containing eight eggs,

took approximately fifteen minutes. For the first few hours

after deposition, the egg mass is less transparent than it be¬

comes later. The gelatinous material of the sheath is appar¬

ently more dense, and somewhat milky in appearance. The

milkiness disappears as time passes, apparently as the sheath

absorbs more water, and swells to full thickness.

Morrison — Life History of Acella.

403

The egg masses are elongate, semi-cylindrical, with rounded

ends. The width is approximately two millimeters ; the length

varies from 3.5 to 6 mm. The individual egg capsules are 1

mm. long and 0.6 mm. wide. The gelatinous outer covering of

the egg mass is about 0.3 mm. thick. The number of eggs

varies from 3 to 12 (Plate XI, fig. 1-4).

Eggs from the individuals collected in February were first

seen one month later (March second). In the case of the in¬

dividuals collected in May, eggs were seen one week after the

date of collection (Plate XI, fig. 1). This indicates that egg

laying is controlled by some sort of rhythm which is attendant

on the arrival of more favorable conditions of growth in the

spring. Temperature seems to be the controlling factor, or the

immediate one at any rate.

The problem of studying the growth of the earliest juveniles

of Acella is much more easily accomplished when the eggs are

brought into the laboratory and hatched out in small aquaria.

In this way the rate of development may be more accurately

noted and the obvious difficulty of searching over all surfaces

of the water plants in a colony location for minute individuals

that are less than 2 mm. in length is overcome. The speci¬

mens upon which the early growth studies were made, were all

raised in battery- jar aquaria, kept under more or less constant

temperature conditions, though somewhat warmer than those

prevailing in the shallow water habitats.

After three days, the embryos are in the trochophore stage,

slowly revolving in the uppermost end of the elongate egg cap¬

sule. About five days after the eggs are laid, the embryos have

developed a shell and are seen to be continually crawling (or

moving by ciliary action?) in all directions over the inner sur¬

face of the capsule (Plate XI, fig. 2-3). At the end of ten

days, hatching of the young has started and continues over four

or five days. (Plate XI, fig. 4.) There is apparently some

variation in the hatching of the snails as in the growth of the

young afterwards.

The shell is 1.3 to 1.5 mm. in length when the young are

hatched; its width is about 0.5 mm. Sixteen days after the

eggs are laid and about three after the young individuals are

hatched, the shell averages 1.8 mm. in length and 0.6 mm. in

width. In twenty days (one week after hatching) the young

404 Wisconsin Academy of Sciences , Arts , and Letters ,

Fig. 2. The growth of Acella, including experimentally raised individuals. The vertical lines represent variation in

size from the average.

Morrison — Life History of Acella.

405

individuals vary in size from 1.6 to 3.4 mm. with an average

shell length of 2.6 mm. The nuclear whorls of these individuals

shows as a portion visibly distinct from the growth that has

occurred since hatching. (Plate XI, fig. 5-6). After a month

(two weeks after hatching) the shell of the juvenile individuals

varies in length from 2.4 to 3.5 mm., with an average size of

3.1 mm.

One group of juvenile Acella was raised in the laboratory

for three and one half months. Measured shortly after hatch¬

ing (June 22, 1930) they averaged 1.8 mm. in shell length, vary¬

ing from 1.4 to 2.0 mm. One month later (July 23, 1930) they

measured between 3.5 and 7.0 mm. with an average length of

5.5 mm. At the end of six weeks (August 6, 1930) they were

between 5.7 and 11.7 mm. long. The average was 8.5 mm. One

individual 9.0 mm. long was found dead August 22, 1930.

(Plate XII, fig. 34). After nine weeks (Sept. 1, 1930) they

averaged 10.8 mm. with a variation of 7.6 to 13.9 mm. One

individual found dead on Sept. 30, 1930, had grown to a length

of 14.0 mm. (Plate XII, fig. 35). The shell length of the re¬

mainder (dead Oct. 5, 1930) varied from 8.7 to 15.0 mm., with

an average of 11.6 mm. (Plate XII, fig. 36-41).

Juvenile individuals have been collected in the summer

months of the year only. Baker, in the course of ecological

studies on Oneida Lake, New York, found immature individu¬

als on July 17 and 24, 1916. He says, “The specimens collect¬

ed were all young, none exceeding 11 mm. in length, the greater

number being 3 to 5 mm. long. . . . Five specimens gave the

following measurements :

Whorls 2 ; length 3.0; breadth .6; aperture length 1.5;

breadth .5 mm.

Whorls 2%; length 4.0; breadth 1.0; aperture length 2.0;

breadth .75 mm.

Whorls 2V2 ; length 5.5; breadth 1.4; aperture length 2.0;

breadth 1.0 mm.

Whorls 3 ; length 8.0; breadth 1.7; aperture length 3.5;

breadth 1.0 mm.

Whorls 3% ; length 10.5; breadth 2.5; aperture length 5.0;

breadth 1.5 mm.

The whorls are usually flatsided as in the adult shell, but in

two specimens, they were somewhat rounded.”

Of the few specimens of these juvenile individuals of Acella

loaned by the New York State College of Forestry, two were

406 Wisconsin Academy of Sciences , Arts, and Letters .

dated July 17, 1916. These were 4.0 and 5.4 mm. long and

averaged 4.7 mm. in shell length (Plate XI, fig. 7-8). Another

specimen, collected July 24, 1916, measured 10.0 mm. (Plate

XI, fig. 9).

Juvenile Acella were collected on July 81, 1930, from a small

bit of marshy shore in a small bay at the southeast corner of

High Lake, Vilas Co., Wisconsin. Measurements of these varied

from 4.6 to 14.9 mm., with the average of 9.7 mm. shell length.

(Plate XII, fig. 1-11). Again on August 2, 1930, juveniles

were found in the small bay immediately to the west of that

collected in a few days previously. The shell length of these

individuals varied from 6.7 to 16.5 mm. with an average of

11.4 mm. (Plate XII, fig. 12-15). The Fishtrap Lake colony of

Acella was searched for young on August 15, 1930. This search

revealed one empty juvenile shell and seven live ones. These

had a shell length of 8.0 to 18.4 mm. with an average of 14.3

mm. (Plate XII, fig. 16-20). The Harris Lake colony was lo¬

cated by the finding of juveniles on August 16, 1930. Here, the

individuals varied in size between 8.6 and 20.0 mm. with an

average shell length of 15.0 mm. (Plate XII, fig. 21-25). One

individual was collected from Dead River, Illinois on August

22, 1925. This specimen, almost full grown, was taken by a

member of Dr. W. C. Allee’s Field Zoology Course of the Uni¬

versity of Chicago, in the course of ecological studies on Dead

River. This individual had a shell length of 19.2 mm.

Specimens collected in September and October in Oneida

Lake, N. Y., by Baker, were loaned by the New York State Col¬

lege of Forestry. Thirty-seven specimens collected Sept. 10,

1916 vary in size from 12.3 to 21.8 mm., with an average shell

length of 16.9 mm. Eight collected Sept. 14, 1916 vary from

18.8 to 22.0 mm., with an average shell length of 20.3 mm.

Five collected Sept. 18, 1916 vary from 18.5 to 25.2 mm., with

an average length of 21.6 mm. Five individuals collected Octo¬

ber 12, 1915 vary from 18.0 to 25.0 mm., with an average of

21.9 mm. shell length.

Of the specimens collected in Dead River, Illinois on Octo¬

ber 13, 1929, twenty-two individuals were measured. They

vary in size between 18.1 and 26.5 mm., with an average shell

length of 22.8 mm. (Plate XII, fig. 26-29, 42). Of the Acella

collected in Fishtrap Lake, Vilas Co., Wisconsin on November

16, 1929, sixty-one were measured. They vary in size between

M orris oiv—Life History of Acella.

407

18.2 and 25.2 mm., with an average length of 21.6 mm. (Plate

XII, fig. 30-33, 43).

Two individuals were collected from the colony in the chan¬

nel between Fishtrap and High Lakes on Feb. 1, 1931. These

measured 15.4 and 16.1 mm. when collected. The average shell

length was 15.7 mm. That these individuals are not just

juveniles that have not completed their growth is indicated by

the amount of erosion evident at the apex of the shell. These

had just 3 and 31/2 whorls remaining when collected. Obser¬

vations on the growth of these two specimens while they were

in the laboratory are of interest. Measured before they started

laying eggs, on February 24th, they were 16.1 and 17.3 mm.

long. Measured at the end of the egg laying period, they were

seen to be 16.0 and 17.2 mm. long, respectively. Later, on

March 19, they showed a total length of 17.7 and 18.7 mm.

From these observations it is evident that shell growth stops

during the egg laying period. The decrease in length at this

time, may be explained by continued erosion of the shell.

There were thirty-seven adult individuals collected from the

Fishtrap Lake colony on May 10, 1930. The erosion of the

spire of these shells is in direct contrast to the perfect speci¬

mens collected the preceding fall. Instead of 5 full whorls,

there were only 2 to 3 whorls to the shell. The tip of the ani¬

mal was about one quarter turn behind the eroding tip of the

shell. These varied in length from 16.8 to 24.1 mm. with an

average shell length of 20.4 mm. These shells also show some

additional shell growth, after a winter ring on the shell. When

kept in the laboratory, these same individuals show more shell

growth after the date of collection. Examination of the indi¬

viduals that lived longest, under laboratory conditions, shows

as much as two or three millimeters growth of shell beyond the

line of the aperture on the date of collection. Again on June

17, 1930, six adults were collected from the Fishtrap Lake Col¬

ony. These individuals also show a winter ring with addi¬

tional growth beyond. The smaller shells, probably those that

were under slightly less favorable growth conditions the pre¬

ceding summer, have more nearly cylindrical shells, while the

larger ones, with a shell of greater diameter, have the aperture

flared out distinctly, more especially on the second season's

growth. These varied in length from 15.6 to 21.7 mm., with

an average shell length of 19.3 mm. Thirty-three individuals

408 Wisconsin Academy of Sciences, Arts, and Letters,

were collected from the colony in the channel between Fishtrap

and High Lakes on June 21 , 1930. They have an average shell

length of 16.5 mm,, varying between 14.5 and 19.8 mm. These

Acella show the same erosion of the apex of the shell as was

noticed on the individuals collected in May and on June 17.

There were about three whorls remaining. The shell shows a

crowded group of growth lines, representing the winter and

there is a narrow band of new shell growth, lighter in color

from one to two millimeters in width. This evidently repre¬

sents the growth up to the date of collection, this season. (Plate

XII, fig. 45-50). One adult individual was found in the sec¬

ond High Lake colony on July 31, 1930. While there is not

much erosion of the apex of this shell, the surface is pitted

somewhat (Plate XII, fig. 44). This individual measured 24.0

mm. when collected. The writer was unable to keep any of the

Acella alive in the laboratory beyond August 11, 1930. That

these snails died of “old age” is the most probable explanation.

This opinion is supported by observations on the amount of

general activity and of the heart beat rate, contrasting them

with the juvenile individuals being reared in the laboratory.

The heart beat rate of the individuals in their second season

(after laying eggs) is approximately one-half that of the juve¬

niles two weeks after they are hatched.

The Dead River habitat was examined for Acella in October

1924 and again in November 1925. These searches resulted

only in the finding of empty shells, probably those of the sea¬

son preceding. These shells are all full grown, with the flared

aperture with a thickened peristome. In some of the individu¬

als, a rest mark and an additional band of shell representing

the second season’s growth can be seen.

The growth of Acella is shown graphically in Fig. 2.

The shell of this species shows variation in convexity of

whorls and shape of the aperture. An analysis of this varia¬

tion, shows that it has a correlation with the type of plant habi¬

tat. In the material studied by the writer, two growth forms

can be seen. One is the form produced when the individuals

live on rushes (Scirpus), This narrow growth form has flat¬

sided whorls and a proportionately narrower aperture. Meas¬

urements of the individuals from Dead River, Illinois, collect¬

ed from rushes, show the following average :

Morrison— Life History of Acella.

409

Aperture length 9.7 mm. Breadth 3.2 mm.

Length-breadth ratio 33.0%

This average includes the measurements from 22 specimens.

The wider growth form is produced when Acella grows on

other plants, such as: yellow and white pond lilies, burreed

(Sparganium), and pondweed (Potamogeton). This wider

growth form has slightly more convex whorls, and a wider

aperture, with the outer portion of the peristome evenly arched.

Measurements of the wide form are as follows:

Fishtrap Lake, Vilas Co., Wisconsin, Nov. 16, 1929.

Average of 61 individuals.

Aperture length 8.9. Breadth 3.4 mm.

Length-breadth ratio 38.2%

Channel between Fishtrap and High Lakes, Vilas Co., Wis.

February 1, 1931. Average of 2 individuals.

Aperture length 7.3 mm. Breadth 2.8 mm.

Length-breadth ratio 38.3%

Fishtrap Lake, Vilas Co., Wis. May 10, 1930.

Average of 37 individuals.

Aperture length 9.2 mm. Breadth 3.5 mm.

Length-breadth ratio 38.0%

Fishtrap Lake, Vilas Co., Wis. June 17, 1930.

Average of 6 individuals.

Aperture length 8.8 mm. Breadth 3.3 mm.

Length-breadth ratio 37.5%

Channel between Fishtrap and High Lakes, Vilas Co., Wis.

June 21, 1930. Average of 36 individuals.

Aperture length 7.6 mm. Breadth 2.8 mm.

Length-breadth ratio 36.8%

High Lake, Vilas Co., Wis. July 31, 1930.

Average of 1 individual.

Aperture length 10.0 mm. Breadth 4.0 mm.

Length-breadth ratio 40.0%

The wide growth form is illustrated by fig. 30-33, 43-50 ; Plate

XII. The narrow form is illustrated by fig. 26-29, 42; Plate

XII. The difference is strikingly seen on comparison of fig¬

ures 42 and 43.

Summary

The snail Acella has a life-span of only one year. The eggs

are laid in the spring, a month or so after the ice leaves the

410 Wisconsin Academy of Sciences, Arts, and Letters .

Plate XI.

Fig. 1. Newly laid eggs of Acella. May 21, 1930.

Still unsegmented after 28 hours.

Fig. 2. Eggs of Acella laid May 17. Average development at 5 days.

Fig. 3. Eggs laid May 17. Maximum development at 5 days.

Fig. 4. Eggs laid May 17. Juveniles just before hatching. June 2,

1930.

Figs. 5-6. Experimentally raised individuals. June 6, 1930. One week

after hatching.

Figs. 7-8. Juvenile individuals collected by Baker in Oneida Lake, N. Y.

July 17, 1916. N. Y. S. C. F. #834 g.

Fig. 9. Juvenile individual collected by Baker in Oneida Lake, N. Y.

July 24, 1916. N. Y. S. C. F. #1021 d.

A convenient scale for figures 1-4 is furnished by the individual egg

capsules, which are approximately 1 mm. long. Figures 5-9 enlarged 10

diameters.

Morrison— -Life History of Acella.

411

TRANS. WIS. ACAD., VOL. 27 PLATE XI

6

9

412 Wisconsin Academy of Sciences y Arts , and Letters,

Figs. 1-11.

Figs. 12-15.

Figs. 16-20.

Figs. 21-25.

Figs. 26-29.

Figs. 30-33.

Figs. 34-41.

Fig. 42.

Fig. 43.

Fig. 44.

Figs. 45-50.

Plate XII.

From colony #1, High Lake, Vilas Co., Wis. July 31, 1930.

From colony #2, High Lake, Vilas Co., Wis. August 2,

1930.

Fishtrap Lake, Vilas Co., Wis. Aug. 15, 1930.

Harris Lake, Vilas Co., Wis. Aug. 16, 1930.

Dead River, Lake Co., Illinois. Oct. 13, 1929.

Fishtrap Lake, Vilas Co., Wis. Nov. 16, 1929.

Experimentally raised. Hatched from eggs, June 22, 1930.

Fig. 34. Dead, Aug. 22, 1930.

Fig. 35. Dead Sept. 30, 1930.

Fig. 36-41. Dead Oct. 5, 1930.

Dead River, Lake Co., Ill. Oct. 13, 1929, specimen showing

narrow growth form as found on Rushes (Scirpus).

Fishtrap Lake, Vilas Co., Wis. Nov. 16, 1929, specimen

showing wider growth form as found on Burreed, Pond-

weed, etc.

High Lake, Vilas Co., Wis. July 31, 1930, specimen in sec¬

ond season.

Channel between Fishtrap and High Lakes, Vilas Co., Wis.

June 21, 1930, specimens in second season showing extreme

amount of erosion.

All figures slightly enlarged.

TRANS. WIS. ACAD., VOL. 27

PLATE XII

Morrison — Life History of Acella.

413

lakes. The juvenile individuals hatch and grow to full size by

early fall. They overwinter as adults, lay eggs the following

spring and die by mid-summer.

Acella does not migrate to deep water, but remains in the

zone of vegetation near shore at all times of the year. When

the vegetation has been killed down by winter conditions, the

snags and logs serve as a substitute habitat on which to live

and lay eggs.

The shell of this species shows variation directly produced by

the habitat. The individuals living on rushes (Scirpus) have

narrower apertures, with almost parallel margins, while those

from other plants show greater convexity of the whorls and

wider apertures, with more evenly arched outer lips.

Bibliography

1. Baker, Frank Collins. 1911. The Lymnaeidae of North and Middle

America. Recent and Fossil. Spec. Pub. No. 8, Chicago Acad. Sci.

pp. 191-198; pi. XVIII, fig. 1; pi. XXVI, fig. 1-4.

2. Baker, Frank Collins. 1916. The Relation of Mollusks to Fish in

Oneida Lake. Technical Pub. No. 4, N- Y. State College of Forestry;

pp. 89, 44, 54, 77, 78, 98, 99, 107, 130, 136, 138-141, 148, 217, 248,

271, 283, 284, 287.

3. Baker, Frank Collins. 1917. Notes on Acella haldemani (Desh.) Bin-

ney. Nautilus 30 : 135-138.

4. Baker, Frank Collins. 1928. The Fresh Water Mollusca of Wiscon¬

sin. Part I. Gastropoda. Bull, 70, Part I, Wis. Geol. Nat. Hist.

Survey (published by the Wis. Acad. Sci. Arts and Letters) ; pp.

265-270; pi. X, fig. 6-8; pi. XI, fig. 8.

5. Sargent, H. E. 1896. Annotated list of the Mollusca found in the

vicinity of Clearwater, Wright Co., Minnesota. Part 2. Nautilus

9 : 127.

6. (Kirkland, R. J.) 1899. An extract from a letter, printed in Nauti¬

lus 12 : 119.

DISSOLVED OXYGEN AND OXYGEN CONSUMED IN THE

LAKE WATERS OF NORTHEASTERN WISCONSIN

C. JT3DAY AND E. A. RlRGE

Notes from the Limnological Laboratory of the Wisconsin Geological

and Natural History Survey.* No. XL1X.

Introduction

Quantitative determinations of the dissolved oxygen content

and of the oxygen consumed by the lake waters of the Highland

Lake District of northeastern Wisconsin were made during a

general chemical survey of a considerable number of these lakes

between 1925 and 1931. These studies were confined chiefly to

the summer season, that is, between late June and the first of

September. During the investigation more than 2,000 dissolved

oxygen determinations were made, and of this number 1,150

were surface samples. The other 850 samples represented 273

series which covered the entire depth of a considerable number

of lakes ; all of the lakes in the district which were known to

have a maximum depth of 18 m. or more were included in these

series. Similar observations were made on a number of the

shallower lakes also, in which the maximum depth ranged be¬

tween 4 m. and 18 m. The series taken on the various lakes

consisted of 2 to 14 samples each, the number of samples de¬

pending upon the maximum depth of the lake and also upon

the status of the dissolved oxygen in the hypolimnion of the

different bodies of water. Where there was a marked decrease

in the quantity of dissolved oxygen in the lower water, samples

were taken at one meter intervals through the region of rapid

decrease and at two or three meter intervals below this stratum.

Determinations of oxygen consumed were made during the

summers of 1929, 1930 and 1931.

Methods

The standard Winkler method was used for the determination

of dissolved oxygen. For purposes of comparison a number

* The 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.

416 Wisconsin Academy of Sciences , Arts, and Letters .

of duplicate determinations were made in which the standard

Winkler method was used for one sample and the Rideal-Stew-

art modification for the other; the differences were within the

limits of error of the two methods and the standard method

was adopted for the routine determinations because it required

less time. The nitrite content of these lake waters rarely ex¬

ceeded a trace, most frequently not even a trace, so that the

modified method for dissolved oxygen was unnecessary.

The method given in Standard Methods of Water Analysis

(A. P. H. A., 1925 edition) was employed in making the oxygen

consumed determinations.

I. Dissolved Oxygen

Surface samples. Surface samples of water for dissolved

oxygen determinations were obtained from 510 lakes during the

period of this investigation. Only a single observation was

made on the surface water of 281 of these lakes, but two or

more surface samples were secured from each of the other

lakes at different times during the progress of this work. These

samples were taken at a depth of 2 cm. to 5 cm. below the sur¬

face of the water so that they came from the stratum which

was freely exposed to the air through the action of the wind

and also from the region where photosynthesis was taking

place.

In spite of the fact that these samples were obtained from

the surface stratum which was in circulation and which was

thus freely exposed to the air where any deficiency or excess of

oxygen tension could be equalized, there was nearly a fourfold

difference in the amount of dissolved oxygen in them. Some

of this difference was due in part to differences in the tempera¬

ture of the water, which affected its oxygen holding capacity,

but the major differences were due to other factors, such as

the respiration of the living organisms, the decomposition of

organic matter, and photosynthesis.

The variations in the quantity of dissolved oxygen in the sur¬

face samples are shown graphically in Figure 1. This diagram

includes 1047 surface samples and the range is from 5.0 mg. to

10.9 mg. of oxygen per liter of water. The maximum number

of samples (108) falls in the 8.0-8.1 mg. group, while 605 of

Juday & Birge — Oxygen in Wisconsin Lake Waters . 417

Fig. 1. The dissolved oxygen in the surface samples of water from the

lakes of northeastern Wisconsin. The vertical scale represents the number

of samples in the various groups and the horizontal spaces show the

amount of dissolved oxygen in milligrams per liter of water, ranging from

5.0 mg. to 10.9 mg., inclusive.

them, or more than 57 per cent of the total number, fall between

7.2 mg. and 8.5 mg. per liter.

The smallest amounts of dissolved oxygen in surface waters

were found in the samples from small bog lakes and bog pools

or ponds. These waters usually contain a relatively large

amount of dissolved organic matter and, in addition, the water

is kept in intimate contact with a large amount of decaying

bog material around the margin and on the bottom. In such

bodies of water, therefore, decomposition is usually taking place

at a vigorous rate. Their small size also makes them less sub¬

ject to disturbance by the wind and thus tends to prevent a

vigorous circulation and aeration of the upper stratum. The

water usually gives a rather strong acid reaction, the hydro¬

gen ion concentration ranging from pH 4.4 to 6.7. A consid¬

erable number of chlorophyll bearing organisms was found,

however, the number varying from 260 to more than 1700 cells

and colonies per cubic centimeter of water in these bug lakelets

and ponds.

Figure 2 shows the distribution of the individual surface

samples on the basis of their percentage of saturation with

oxygen; this diagram covers the range from 64 per cent to

108 per cent. The samples above and below these percentages

are so widely scattered that they were omitted from the dia¬

gram. The maximum number of samples, namely 99, falls in

the 86-87 per cent group. The 88-89, 90-91, and 92-93 per cent

418 Wisconsin Academy of Sciences , Arts , and Letters.

groups contain 94, 85, and 86 samples respectively, and there

are 77 samples in the 84-85 per cent group. These five columns

contain 441 samples out of the 951 represented in the diagram,

or a little more than 46 per cent of the total number included.

Half of the surface samples, namely 477, fall between 83 per

cent and 93 per cent of saturation.

Fig. 2. The percentage of oxygen saturation in the surface samples of

water. The vertical scale shows the number of samples in each group and

the horizontal spaces indicate the percentage of saturation at 2 per cent

intervals from 64 per cent to 109 per cent, inclusive; the first column rep¬

resents 64 per cent and 65 per cent, the second column 66 per cent and 67

per cent and so on to 108 per cent and 109 per cent in the last column.

Note the maximum of 99 samples in the 86-87 per cent column. The fig¬

ure includes 951 samples.

It will be noted that there is a marked difference between

the number of samples in the 68-69 per cent column and that

in the 70-71 per cent column; there are 8 in the former as com¬

pared with 27 in the latter. Another abrupt change is shown

between the 92-93 per cent column and the 94-95 per cent col¬

umn ; the former represents 86 and the latter 49 samples.

Below the column representing 70 per cent of saturation,

there are 4 samples from bog lakelets or ponds which fall be¬

tween 34 per cent and 53 per cent, 7 lake samples between 55

per cent and 59 per cent, and 23 lake samples between 60 per

cent and 69 per cent, inclusive.

In 32 samples the oxygen varied from 106 per cent to 129

per cent of saturation. Thirty-nine samples fall between 101

per cent and 105 per cent, inclusive; adding these to the 32

above 105 per cent gives a total of 71 samples in which the

Juday & Birge— Oxygen in Wisconsin Lake Waters. 419

quantity of dissolved oxygen exceeds the amount required for

saturation. This is approximately 7 per cent of the total num¬

ber of surface samples, namely, 1,056. The percentages of sat¬

uration found in some of the lakes are given in Tables I, III

and VIII.

Table I shows the variations in the oxygen content of the

surface waters of a few lakes that were visited several times

during the progress of the investigation. These results show

the range of variation that is to be expected from irregular

visits to such bodies of water. A much more detailed study of

each lake would be required to ascertain the seasonal and diur¬

nal changes which take place in the dissolved oxygen content

of the surface water. The various bodies of water represented

in this table were selected for the purpose of illustrating the

results obtained on the different types of lakes and lakelets.

Adelaide Lake is rather small (22 ha.), but has a maximum

depth of 21.5 m.; the water is soft and has a certain amount

of brown color, ranging from 32 to 45 on the platinum-cobalt

scale. Bear Lake has an area of about 104 ha., but it is shal¬

low (maximum depth 10 m.) so that the water is not thermally

stratified in summer. The water is soft and the brown color

is generally low (16). The open water in the Cardinal Bog

is only about 25 m. in diameter and it is surrounded by a wide

margin of typical bog; the maximum depth of the water is 5.8

m. The water is soft and the brown color ranges from 32 to

49. Crystal Lake has an area of 31 ha. and a maximum depth

of 21 m. ; its water is clear, transparent and soft, with no brown

color at all or only a trace.

Lake Mary has an area of 1.2 ha. and a maximum depth of

22 m. Its water is soft and rather highly colored, the color

ranging from 100 to 122 on the platinum-cobalt scale. Mus-

kellunge Lake has an area of about 375 ha. and a maximum

depth of 21 m. The water has a medium quantity of fixed or

bound carbon dioxide and very little brown stain, the latter

ranging from 0 to 10. Nebish Lake has an area of 47 ha. and

a maximum depth of 15.8 m. The water is soft and has a

brown color ranging from 0 to 14. Silver Lake has an area of

87 ha. and a maximum depth of 19.5 m. The water has a fairly

large amount of fixed carbon dioxide for this region and very

little or no brown color at all.

420 Wisconsin Academy of Sciences , Arts , and Letters .

Trout Lake, with an area of about 1683 ha., is the second

lake in size in the Highland Lake District; it has a maximum

depth of 35 m., which makes it the deepest lake in the district.

The water carries a relatively large amount of fixed carbon di¬

oxide and it has very little brown color (0-14). Weber Lake

has an area of 15.6 ha. and a maximum depth of 13.5 m. ; its

depth is sufficient, however, to produce a thermal stratification

of the water in summer. The water is very soft and has very

little or no brown color. Wild Cat Lake has an area of 102 ha.

and a maximum depth of 12 m. ; the water is thermally strati¬

fied in the deepest part of the lake in summer. The water con¬

tains a larger amount of fixed or bound carbon dioxide than

any other lake in this district and the brown color ranges from

16 to 26 on the platinum-cobalt scale.

The lowest percentage of oxygen saturation in surface water

was found in the Cardinal Bog where only 34 per cent of the

amount required for saturation was noted in the surface sample

on August 20, 1927. The largest amount found in this bog

pool reached only 73 per cent of saturation, so that there was

somewhat more than a twofold difference between maximum

and minimum. In the Forestry Bog the range was from 45 per

cent to 84 per cent, just a little less than a twofold difference.

In Little Star Bog it was from 60 per cent to 98 per cent; the

open water in this bog has an area of about 2 ha. so that the

surface water is more thoroughly aerated.

Low percentages of saturation were also noted in the sur¬

face samples of Lake Mary; though none of them fell below

70 per cent, only 2 out of the 6 samples were above 80 per cent.

These low percentages may be accounted for, in part at least,

by the fact that this lakelet is partially surrounded by boggy

shores.

In the other bodies of water represented in Table I, 12 sur¬

face samples fell between 68 per cent and 80 per cent of satura¬

tion, 76 between 81 per cent and 100 per cent and 6 above 100

per cent. The maximum percentage (120) was found in Bear

Lake on July 1, 1928; the lowest percentage in this lake was

68 per cent, so that there was almost a twofold range between

maximum and minimum. The surface samples included in

Table I were taken at different hours of the day, but most fre¬

quently during the forenoon, so that they do not represent the

daily maximum quantities in most cases ; neither do they repre-

Juday & Birge — Oxygen in Wisconsin Lake Waters . 421

sent the minimum amount that would be found in the early

morning.

The mean quantity of dissolved oxygen in the surface waters

of the 510 lakes was 7.6 mg. per liter; the range was from a

minimum of 3.4 mg. to a maximum of 12.4 mg. per liter of

water. The mean percentage of saturation of all of .these sur¬

face samples was 82 per cent;, the percentage varied from a

minimum of 34 per cent noted in a surface sample from the

Cardinal Bog to a maximum of 129 per cent in a sample from

the surface of Little St. Germain Lake.

During the summers from 1907 to 1910 observations were

made on the dissolved oxygen content of 60 of these northeast¬

ern lakes ; 70 determinations were made on surface waters dur¬

ing this period. In 59 of the 70 surface samples the quantity

of oxygen ranged from 7.5 mg. to 8.9 mg. per liter ; in the en¬

tire number of samples the amount varied from 6.8 mg. to 9.6

mg. per liter, while the mean quantity was 8.2 mg. In per¬

centages the range was from 71 per cent to 102 per cent of

saturation, with a mean of 88 per cent. The mean percentage

of saturation in these 70 samples, therefore, was 6 per cent

larger than that of the samples obtained between 1925 and

1931. No bog samples, however, were included in the former

observations.

In the summers of 1907 to 1909 surface samples were ob¬

tained from 62 lakes in northwestern Wisconsin and they in¬

cluded 64 determinations of dissolved oxygen. In 45 cases the

quantity of oxygen fell between 8.0 mg. and 9.4 mg. per liter,

while the mean quantity in the 64 surface samples was 8.7 mg.

per liter. The percentage of saturation in the surface waters

of the various lakes varied from 68 per cent to 168 per cent;

that is, the maximum was about two and a half times as large

as the minimum. The mean percentage for the 64 samples was

99 per cent, or 17 per cent above the mean percentage found

in the surface samples taken between 1925 and 1931 in the

northeastern lakes.

Oxygen determinations were made on 120 surface samples

obtained from. 24 lakes in southeastern Wisconsin, exclusive of

Lake Men dot a, between 1905 and 1910. Of this number 101

samples yielded between 8.0 mg. and 9.4 mg. per liter ; the range

for all samples was from a minimum of 6.7 mg. to a maximum

of 14.2 mg. per liter. More than half (66) of the samples con-

422 Wisconsin Academy of Sciences , Arts, and Letters .

tained 96 per cent to 104 per cent of the quantity of oxygen re¬

quired for saturation; the mean for the 120 samples was just

a little over 100 per cent, with a minimum of 77 per cent and a

maximum of 131 per cent. Thus the mean percentage of oxy¬

gen saturation in these surface samples from the southeastern

lakes was a little more than 18 per cent higher than that in the

surface samples of the northeastern lakes and somewhat more

than 1 per cent above that in the northwestern lakes.

Vertical Distribution of Dissolved Oxygen

Series of oxygen samples covering the entire depth of the

water were obtained from 76 of the northeastern lakes. In the

shallowest ones, those having a maximum depth of 4 m. to 5 m.,

only surface and bottom samples were taken in some instances,

but in the deeper lakes the number of samples in a series varied

from 3 to 14, depending upon the depth of the lake and the

scarcity or abundance of oxygen in the hypolimnion. Includ¬

ing those of 1931, 261 series of samples were taken ; the larg¬

est number from a single lake, namely 32, was obtained from

Trout Lake.

For convenience in the discussion of the results, the lakes

are separated into four groups on the basis of their depth. The

maximum depth and the area of the various lakes are shown

in Table II. The first group includes the 17 lakes which do not

exceed 9.5 m. in depth, the second group 15 between 10 m. and

14.5 m. in depth, the third group 19 between 15 m. and 19.5 m.,

and the fourth 25 that are 20 m. to 35 m. deep. The vertical

distribution of the dissolved oxygen in some of these lakes is

given in Table III.

Group L Jag Lake in the first group has a maximum depth

of only 4 m. and the bottom water contained substantially the

same amount of oxygen as the surface. The next in order of

depth are the Cardinal Bog (5.5 m.) and Little Star Bog (6

m.). The open water in the Cardinal Bog is only about 25 m.

in diameter and is well protected from wind by the surround¬

ing forest, so that its water is subject to only slight distur¬

bances by air currents. As a result there is a marked thermal

stratification of the water in spite of the shallowness of the

pool; the temperature at a depth of 5.5 m., for example, was

only 4.7° C. on July 18, 1926, and 5.2° on August 18, 1931. A

Juday & Birge — Oxygen in Wisconsin Luke Waters . 423

lower temperature than this has been found in summer in only

one other northeastern lake, namely Lake Mary, where a me¬

dian stratum usually has temperatures ranging between 4.0°

and 4.5°.

The upper 2 m. of the Cardinal Bog contained only 50 per

cent to 53 per cent of the quantity of oxygen required for sat¬

uration on July 18, 1926 and less than 1 per cent of saturation

was found at 3 m. to 5 m. ; only a trace of oxygen was found

below 3 m. on August 18, 1931. Little Star Bog has an area

of about 2 ha., so that the wind produces a fair circulation of

the water; the upper stratum contained from 60 per cent to

98 per cent of the amount of oxygen required for saturation,

while the quantity varied from 34 per cent to 48 per cent of

saturation at a depth of 5 m. Yolanda is also a bog lakelet and

its water possessed only a trace of oxygen at 3 m. and none be¬

low this depth on August 23, 1931.

The vertical distribution of the oxygen in three of the lakes

belonging to the shallow group is shown in Figures 3 to 5.

These diagrams show that there was a decrease in the quantity

of dissolved oxygen in the lower water of these three lakes.

In Dorothy Dunn Lake the decrease took place between 3 m.

and 6 m., corresponding to the fall in the temperature of the

water. Only 3 mg. of oxygen per liter were found at 6 m.,

which was only 30 per cent of saturation at that depth. (See

Fig. 3).

Fig. 3. The dissolved oxygen (02) and the temperature (T) of Dorothy

Dunn Lake on August 28, 1926. The vertical spaces represent the depth

in meters and the horizontal spaces show the amount of oxygen and the

temperature of the water. The quantity of oxygen is given in milligrams

per liter and in percentage of saturation. The scale at the top of the di¬

agram indicates the quantity of oxygen in milligrams per liter of water

and the temperature in degrees centigrade, while that at the bottom of the

diagram represents the percentage of oxygen saturation.

424 Wisconsin Academy of Sciences , Arts , and Letters .

In Finley Lake (Fig. 4) the water was well supplied with

oxygen down to a depth of 7 m. ; the quantity at this depth was

above 82 per cent of saturation in each of the three summers

in which observations were made on this lake. There was a

more or less marked decrease at 8 m. ; this decrease was most

marked in 1927 when the quantity fell to 8.1 mg. per liter on

July 28 at 8 m. The oxygen content of the surface water was

smallest in 1929. The water of this lake is fairly transparent,

the readings with a Secchi disc varying from 3.7 m. to 6 m.

Readings with a pyrlimnometer in this lake on July 19, 1929

showed that more than 1.5 per cent of the energy delivered to

the surface of the lake by the sun penetrated to a depth of 7 m. ;

at this depth the energy is in the form of light and would repre¬

sent twice this amount if expressed in terms of illumination.

The energy is sufficient to enable the chlorophyllaceous organ¬

isms to carry on photosynthesis at this depth and help to keep

the lower water well supplied with oxygen.

Fig. 4. The dissolved oxygen (02) and the temperature (T) of Finley

Lake on July 28, 1927, July 19, 1928, and August 10, 1929, indicated in

milligrams per liter of water and in degrees centigrade, respectively. Com¬

pare with Fig. 5.

Figure 5 shows the oxygen results obtained on Bragonier

Lake. This body of water is less than half a kilometer from

Finley Lake and it is smaller than the latter, having an area

of 19.2 ha. as compared with 62.8 ha. in Finley. As a result

of its smaller size and its protection from wind by the sur¬

rounding forest, the epilimnion of Bragonier is only 1 m. to 2

m. thick in July and August as compared with 4 m. in Finley.

Bragonier also has a brown stained water with the color rang¬

ing from 22 to 45 on the platinum-cobalt scale at the surface

and from 60 to 126 at 8 m. ; in Finley Lake the brown color

Juday & Birge — Oxygen in Wisconsin Lake Waters . 425

does not exceed 8. The vegetable stain in Bragonier Lake adds

materially to the quantity of organic matter in the water and

thus increases the demand for oxygen in the process of decom¬

position; the average amount of organic carbon in the surface

water of Bragonier Lake was 6.1 mg. per liter as compared

with 4.5 mg. in Finley Lake. As a result of the brown stain

in the water of Bragonier Lake the sun’s energy is reduced to

less than 0.2 per cent at a depth of 4 m.

FIG. 5. The dissolved oxygen (02) and the temperature (T) of Brag¬

onier Lake on July 28, 1927, July 19, 1928 and August 10, 1929, indicated

in milligrams per liter of water and in degrees centigrade. Compare with

Fig. 4.

The oxygen was substantially uniform in quantity in the up¬

per 4 m. of Bragonier Lake in 1927 and in 1928, but there was

a marked decrease at 5 m. and only a trace or none at all was

found at 8 m. Similar results were obtained in 1929 except

that the decrease in the quantity of oxygen was noted in the

3 m.-4 m. stratum instead of in the 4 m.-5 m. stratum as in

the former years. There was a large increase in the free carbon

dioxide below 4 m. on July 28, 1927 ; the amount of fixed or

bound carbon dioxide at 8 m. on this date was a little more

than twice as large as that at the surface. In general the lower

water of Bragonier Lake is eutrophic in character, while that

of Finley Lake is oligotrophic.

Group II. With the exception of Diamond Lake, the lakes

included in Group II show a more definite thermal stratifica¬

tion in summer than those in Group I. Diamond Lake has an

area of 48.4 ha. and a maximum depth of only 12.3 m., so that

from 80 per cent to 87 per cent of its water is kept in circula¬

tion during the summer ; as a result, the difference in tempera-

426 Wisconsin Academy of Sciences , Arts , and Letters .

ture between surface and bottom water is not as great in this

lake as it is in the other members of this group. Figure 6

shows that the water was well aerated at all depths in the three

years represented in the diagram. On July 24, 1929, the amount

of dissolved oxygen at 10 m. was 109 per cent of that required

for saturation, while the quantity at 11 m. was 99 per cent.

The curves for 1927 and 1929 coincide down to a depth of 7

m., but in the latter year there was a more marked increase

below this depth than in the former year. The water of Dia¬

mond Lake is transparent, the disc readings ranging from 7.6

m. to 9 m. and about 2 per cent of the sun’s energy penetrates

to the bottom in the deepest water. This light is sufficient to

enable the phytoplankton to carry on the process of photosyn¬

thesis in the lower water and thus furnish enough oxygen to

keep the lower stratum well supplied with this gas.

Fig. 6. The dissolved oxygen (02) and the temperature (T) of Diamond

Lake on July 15, 1927, August 29, 1928 and July 24, 1929.

Blue and Weber lakes (Figs. 7 and 8) contained an abun¬

dance of dissolved oxygen down to a depth of 8 m., more than

90 per cent of the amount required for saturation being pres¬

ent in this stratum. Both lakes showed an excess of oxygen

at 9 m. and 10 m. ; the quantity of this gas rose to 106 per cent

and 102 per cent of saturation at these depths, respectively, in

Blue Lake on August 6, 1929, and to 111 per cent and 101 per

cent in Weber Lake on August 21, 1929. The percentage of

saturation at the various depths is shown for both lakes in

Juday & Birge — Oxygen in Wisconsin Lake Waters . 427

Fig. 7. The dissolved oxygen (Os) and the temperature (T) of Blue

Lake on August 6* 1929. The quantity of oxygen is indicated both in milli¬

grams per liter of water and in percentage of saturation. The scale at

the top of the diagram indicates the quantity of oxygen in milligrams per

liter and the temperature in degrees centigrade* while that at the bottom

shows the percentage of oxygen saturation.

Fig. 8. The dissolved oxygen and the temperature of Weber Lake on

August 21* 1929. The oxygen is shown in milligrams per liter and in per

cent of saturation. The quantity of oxygen was above the saturation point

at 9 m. and 10 m. Compare with Fig. 7.

Figures 7 and 8. There was a rather marked decrease in the

quantity of oxygen below 10 m, in both lakes. The water of

these two lakes is transparent* so that photosynthesis can take

place at considerable depths. More than 3 per cent of the sun's

energy penetrates to a depth of 10 m. in Weber Lake and a

little less than 3 per cent is found at that depth in Blue Lake.

A more pronounced decrease of dissolved oxygen in the lower

water was noted in Midge Lake on July 31* 1929 (Fig. 9) ; the

428 Wisconsin Academy of Sciences , Arts, and Letters .

quantity of oxygen in this lake ranged from 83 per cent of

saturation at the surface to 10 per cent at the bottom. Long

Lake (Fig. 10) showed a still more marked decrease in oxy¬

gen in the lower water; only a trace was found between 8 m.

and 10 m., and none below the latter depth. Even the sur¬

face water reached only 76 per cent of saturation on August

23, 1928. There was a marked increase in the free carbon di¬

oxide in the lower water correlated with the decrease in oxy¬

gen.

Group III . This group includes 19 lakes and they belong to

three general types. Two of these lakes had an abundance of

dissolved oxygen at all depths ; 5 showed a more or less marked

0 3 10 15 20 25

Fig. 9. The dissolved oxygen and the temperature of Midge Lake on

July 31, 1929. The oxygen was well below the saturation point at all

depths, especially in the lower strata.

Fig. 10. The dissolved oxygen and the temperature of Long Lake (south

of Crystal Lake) on August 23, 1928. No dissolved oxygen was found

below 10 m.

Juday & Birge— Oxygen in Wisconsin Lake Waters. 429

excess of oxygen in the thermocline or mesolimnion, with a

decrease in the amount below this stratum; in 12 of them

there was no increase in the thermocline and there was a

marked decrease in the amount of oxygen in the lower stratum

or hypolimnion.

Island and Little Long lakes are the two members of this

group which possessed a good supply of dissolved oxygen at all

depths. In Island Lake (Fig. 11) the oxygen content of the

surface water was 8.7 mg. per liter, or 91 per cent of satura¬

tion, on August 25, 1926; at 14 m. (1 m. above the bottom) it

was 4.7 mg. per liter, or 42 per cent of saturation.

Fig. 11. The dissolved oxygen and the temperature of Island Lake (near

Forest Lake) on August 25, 1926. Dissolved oxygen was abundant in the

hypolimnion.

The surface sample of Little Long Lake (Fig. 12) contained

7.6 mg. of oxygen per liter on August 22, 1931, equivalent to 83

per cent of saturation; the 17 m. sample yielded 3.1 mg. of

oxygen per liter, or 24 per cent of saturation. On July 24,

1930 the percentages of saturation at surface and bottom were

75 per cent and 31 per cent respectively. Attention may be

called to the notch at 5 m. in the two oxygen curves in Figure

12; the quantity of oxygen fell from 7.1 mg. per liter at 3 m.

to 5.0 mg. at 5 m. and then rose to 6.1 mg. and 6.3 mg. per liter

at 7 m. and 8 m. respectively. A similar decrease was found

at 5 m. in the series taken in this lake on July 24, 1930.

Pallette Lake (Fig. 13) represents the type in which there

is an excess of oxygen in the thermocline. On August 22, 1928,

the sample from a depth of 8 m. yielded 13.6 mg. of oxygen per

430 Wisconsin Academy of Sciences , Arts , and Letters .

Fig. 12. The dissolved oxygen and the temperature of Little Long Lake

on August 22, 1931. Note the irregular distribution of the oxygen below

3 m.

liter of water and that from 9 m. contained 12.6 mg. per liter;

these amounts represented saturations of 139 per cent and 119

per cent, respectively. The curve for oxygen saturation in

Figure 13 shows that all the samples between 7 m. and 10 m.

contained more oxygen than was required for saturation. The

phenolphthalein alkalinity amounted to 2.6 mg. per liter at 8

m. and 0.9 mg. at 9 m. A marked difference was noted in the

hydrogen ion concentration at 8 m.; there was a change from

pH 6.6 at 7 m. to pH 9.1 at 8 m. and back again to pH 6.9 at

9 m., while the surface water was pH 6.7 and that at 16 m.

was pH 6.3. Excess oxygen was also found in the region of

the thermocline of Pallette Lake on July 24, 1925 (123 per cent

at 8 m.), on August 18, 1927 (112 per cent at 8 m.) and on

July 21, 1931 (118 per cent at 7 m.) ; on the latter date the

water was alkaline to phenolphthalein between 5 m. and 8 m.,

but this was accompanied by only a slight change in the hydro¬

gen ion reading. In 1927 the water was alkaline to phenolph¬

thalein at 8 m. and 9 m. and the hydrogen ion readings changed

from pH 6.5 at 7 m. to pH 7.8 at 8 m., pH 8.2 at 9 m. and back

to pH 6.7 at 10 m.; the surface was pH 6.4 and 12 m. and 15

m. pH 6.0.

Excess oxygen was also obtained in the thermocline of An¬

derson, Day, Pauto and Silver lakes, but the percentages in

Juday & Birge — Oxygen in Wisconsin Lake Waters. 431

Fig. 13. The dissolved oxygen and the temperature of Pallette Lake on

August 22, 1928. There was a considerable excess of oxygen at 8 m.

these four lakes were not as large as the maximum in Pallette

Lake. In Anderson Lake the dissolved oxygen amounted to

12.8 mg. per liter at 7 m., 14.2 mg. at 8 m. and 13.4 mg. at 9

m. on August 8, 1929, representing respectively 123 per cent,

128 per cent and 116 per cent of saturation at these depths.

There was a distinct phenolphthalein alkalinity at 7 m. and 8

m. and corresponding to this was a change in hydrogen ion

readings from pH 7.5 at 6 m. to pH 8.4 at 7 m. and 8 m.

In two of the four series of observations made on Day Lake,

an excess of oxygen was noted in the thermocline. On August

4, 1928 the sample from 8 m. yielded 10.0 mg. of oxygen per

liter, or 102 per cent of saturation and that from 9 m. gave

10.8 mg., or 103 per cent. A series taken on July 23, 1929

yielded 10.8 mg. per liter at 7 m. and 11.4 mg. 8 m., represent¬

ing 104 per cent and 106 per cent of saturation respectively.

The series taken in 1926 and 1927 did not show excess oxygen

at any depth.

The highest percentage of dissolved oxygen obtained in the

thermocline of Pauto Lake was 111 per cent, which was found

in a 6 m. sample on July 10, 1929; 108 per cent was noted at

7 m. on this same date. An excess of oxygen was found at 8

m., 9 m. and 10 m. on July 30, 1927, with 108 per cent at 9 m.

as the maximum. In neither of these series was there any

change in the phenolphthalein alkalinity or in the hydrogen ion

concentration corresponding to the excess oxygen. A series of

432 Wisconsin Academy of Sciences, Arts, and Letters .

samples obtained from Pauto Lake on July 10, 1928 did not

show an excess of oxygen at any depth.

There was a marked decrease in the quantity of dissolved

oxygen in the lower water of these 5 lakes, but in 4 of them

the amount did not fall below 0.5 mg. per liter at the bottom in

any of the series. In Pallette Lake, on the other hand, only a

trace was found at the bottom in the series represented in Fig¬

ure 13.

The third type of lake in this group, namely those which

have a bottom stratum with comparatively little or no dissolved

oxygen in late August, are represented in Figures 14 to 16.

Two Sisters Lake (Fig. 14) shows a much more marked de¬

crease in oxygen in the lower stratum than Island and Little

Long lakes; although the sample at 18.5 m. in Two Sisters

Lake yielded a little more than 1 mg. of oxygen per liter on

August 13, 1929, a series of samples taken on August 27, 1907

showed only a trace of this gas at 17 m. and none at 19 m.

The free carbon dioxide showed a large increase in the hypolim-

nion, with a miximum of 15.6 mg. per liter at 18.5 m.

Fig. 14. The dissolved oxygen and the temperature of Two Sisters Lake

on August 13, 1929. There was a marked decrease in the quantity of oxy¬

gen below 9 m.

The bottom water of Papoose Lake (Fig. 15) contained only

a small amount of dissolved oxygen during the latter part of

August, both in 1926 and in 1929. The curves show that there

Juday & Birge — Oxygen in Wisconsin Lake Waters. 433

was a larger amount of oxygen between 9 m. and 15 m. in 1929

than in 1927. The saturation curve for 1926 is not shown in

the diagram because it falls on the temperature curves in the

epilimnion; the percentage of saturation throughout the epi-

limnion in this year was below that of 1929, ranging from 79

Fig. 15. The dissolved oxygen and the temperatures of Papoose Lake on

August 27, 1927 and on August 15, 1929. Very little oxygen was found be¬

low 15 m. in both years.

Fig. 16. The dissolved oxygen and the temperature of Nebish Lake on

August 29, 1931. Only a small quantity of oxygen was found below 10 m.

434 Wisconsin Academy of Sciences, Arts, and Letters.

per cent at the surface to 76 per cent at 8 m. Big Lake repre¬

sents a somewhat more advanced stage in the disappearance of

the oxygen in the lower stratum; usually only a trace or no

oxygen at all was present at 15 m. in this lake in late August

and none below this depth.

Nebish Lake (Fig. 16) also shows a total disappearance of

the dissolved oxygen in the greater part of the hypolimnion in

late August ; very little oxygen was found below 10 m. on Aug¬

ust 29, 1931. There was a very large increase in the free car¬

bon dioxide below 8 m. on this date, the amount rising to 29

mg. per liter of water at 14.5 m.

Group IV. This group includes 25 lakes with a maximum

depth of 20 m. or more (Table II). The oxygen results ob¬

tained on 10 of them are shown in Figures 17 to 27, inclusive;

several series of oxygen determinations are included in Tables

III, VI and VIII. The upper stratum or epilimnion of these

lakes contained from 7 mg. to about 9 mg. of dissolved oxygen

per liter, which represented from 70 per cent to 100 per cent

of saturation; a few of the surface samples yielded an excess

of oxygen.

Excess oxygen was found in the thermocline of six lakes be¬

longing to this group. It was noted in 5 of the 11 series from

Crystal Lake, in one out of 5 series from Big Carr, in one out

of 4 series from Black Oak and in one out of 7 series from

Clear Lake. Only one series has been taken on Catfish Lake

and on Yawkey Lake, and excess oxygen was obtained at 8 m.

in the former and at 10 m. in the latter. The excess was rela¬

tively small in all cases, however, ranging from 101 per cent

to 111 per cent of the amount required for saturation. The

maximum percentage, namely 111 per cent, was observed in

Crystal Lake at 10 m. on August 12, 1931.

The various lakes in this group show marked differences in

the quantity of dissolved oxygen that persists in the hypo¬

limnion, or lower stratum of water during the summer; they

form a fairly continuous series ranging from those which have

an abundant supply of oxygen (4 mg. to 8 mg. per liter) in the

lower water during late August to those in which the hypo¬

limnion is practically devoid of free oxygen by the latter part

of August. The results shown in the diagrams (Figs. 17 to 27)

were selected for the purpose of illustrating the different stages

Juday & Birge — Oxygen in Wisconsin Lake Waters. 435

in the disappearance of the dissolved oxygen in the hypolimnion

during the course of the summer.

Crystal Lake (Fig. 17) has soft water, since it has only an

average of about 1 mg. of fixed or bound carbon dioxide per

liter, and it supports a relatively small crop of phytoplankton.

The number of chlorophyll bearing organisms ranged from a

minimum of 137 cells and colonies per cubic centimeter of wa¬

ter in early July to a maximum of 1035 cells and colonies per

cubic centimeter in late August. The lake also supports a

rather large crop of Daphnias and these cladocerans are an im¬

portant factor in keeping the crop of phytoplankton down to

the limits indicated above. The water is clear and the most

transparent of any that has been found in the northeastern

group of lakes. Disc readings of 12 m. or more have been ob¬

tained and more than 1 per cent of the sun’s energy that is de¬

livered to the surface of the lake penetrates to a depth of 18 m.

So much sunlight reaches the lower water that the moss Dre-

panocladus aduncus var. aquations grows abundantly upon the

bottom at depths of 17 m. to 20 m.

Fig. 17. The dissolved oxygen and the temperature of Crystal Lake on

August 21, 1928. The quantity of oxygen was above the saturation point

at 10 m. Compare with Fig. 13 and Fig. 18.

Figure 17 shows that the lower water of Crystal Lake con¬

tained an abundant supply of dissolved oxygen on August 21,

1928. The water at 19 m. (1 m. above the bottom) yielded 8.5

mg. of oxygen per liter, which was equivalent to almost 75 per

cent of saturation; the quantity at 10 m. was 104 per cent of

436 Wisconsin Academy of Sciences, Arts, and Letters .

saturation and that at 12 m. was 100 per cent. With such a

small supply of phytoplankton as this lake supports, a propor¬

tionately small amount of organic material derived from this

source sinks into the lower stratum and decays there, and this

material is probably the most important factor in the depletion

of the dissolved oxygen in the hypolimnion. The shores are

sandy and there is neither an inlet nor an outlet, so that very

little organic matter reaches the lake from outside sources. It

seems probable also that an important factor in maintaining the

supply of oxygen in the lower stratum is the photosynthesis

carried on in this region both by the phytoplankton and by the

moss Drepanocladus which thrives on the bottom in the deeper

water. Thus the relatively small quantity of organic matter

that reaches the hypolimnion from the epilimnion makes a cor¬

respondingly small demand for oxygen in its decomposition,

and this small demand, together with the oxygen that may be

liberated in the process of photosynthesis carried on in this

region, leaves an abundant supply of oxygen in the bottom

stratum throughout the summer. The free carbon dioxide fur¬

nishes a good index of the comparatively small amount of de¬

composition in the lower water ; it shows only a small increase

in the bottom water.

Similar results were obtained on Crystal Lake on August

20, 1929 and on August 12, 1931. In the former year the per¬

centage of saturation varied from 91 to 93 per cent in the epi¬

limnion, from 98 to 104 per cent in the thermocline and from

71 to 83 per cent in the hypolimnion; in the 1931 series the

quantity of oxygen ranged from 91 to 92 per cent in the epi¬

limnion, from 100 to 111 per cent in the thermocline and from

73 to 83 per cent in the hypolimnion.

Big Carr Lake (Fig. 18) has soft water (1.5 mg. of fixed

or bound carbon dioxide per liter) and a fair degree of trans¬

parency, so that approximately 2 per cent of the sun's energy

penetrates to a depth of 10 m. The phytoplankton is somewhat

more abundant than in Crystal Lake ; the number of organisms

ranges from 283 to 1135 cells and colonies per cubic centimeter

of water in the upper 10 m., with an average of about 730. A

slight excess of dissolved oxygen was obtained at a depth of

7 m. on July 18, 1927 and the bottom sample yielded 3.6 mg.

per liter, representing a little more than 30 per cent of satura-

Juday & Birge— Oxygen in Wisconsin Lake Waters. 437

tion. In the 5 series of samples taken on Big Carr Lake, the

minimum noted at the bottom was 2.4 mg. per liter, or 21 per

cent of saturation, on July 22, 1925. A maximum of 8.5 mg.

of free carbon dioxide per liter was found at 21 m. on this

date.

FlG. 18. The dissolved oxygen and the temperature of Big Carr Lake on

July 18, 1927. The quantity of oxygen was a little above the saturation

point at 7 m. Compare with Fig. 17.

The results of the two series of oxygen samples obtained

from Dead Pike Lake are shown in Fig. 19. The water of this

lake has an average of about 11.5 mg. of fixed carbon dioxide

per liter and it contains a moderate amount of vegetable stain,

so that the color readings range from 60 to 68 on the platinum-

cobalt scale. The epilimnion contained from 73 per cent to

75 per cent of the quantity of oxygen required for saturation

in the series taken on August 9, 1927, the thermocline from 44

per cent to 64 per cent and the hypolimnion from 30 per cent

to 41 per cent; the curves for the percentages have been omit¬

ted from the diagram because they fell too close to the tem¬

perature curves in the region of the epilimnion. The quantity

of free carbon dioxide at the bottom amounted to 8.5 mg. per

liter in 1927 and 7.0 mg. in 1928.

438 Wisconsin Academy of Sciences , Arts , and Letters .

Fig. 19. The dissolved oxygen and the temperature of Dead Pike Lake

on August 9, 1927, and on July 28, 1928. Note that the amount of oxygen

was substantially the same in the two years. Compare with Fig. 12.

Fence Lake (Fig. 20) is third in size among the lakes be¬

longing to this group. Its water contains 16 mg. to 17 mg. of

fixed carbon dioxide per liter and the hydrogen ion ranged

from pH 7.7 at the surface to pH 6.8 at 27.5 m. The upper

5 m. of the epilimnion contained a slight excess of dissolved

oxygen (101 to 102 per cent of saturation) and this was cor¬

related with a rather large crop of phytoplankton ; the surface

water yielded 2845 cells and colonies of chlorophyll bearing or¬

ganisms per cubic centimeter and that at 5 m. 2337 per cubic

centimeter, with 2658 per cubic centimeter at 10 m. The lower

part of the epilimnion yielded 97 per cent to 98 per cent of the

amount of oxygen required for saturation, but there was a

marked decrease in the thermocline; the hypolimnion yielded

from 32 per cent to 47 per cent of saturation. The free carbon

dioxide showed an increase in the thermocline and rose to a

maximum of 7.8 mg. per liter at 27.5 m.

One series of oxygen samples was obtained from Crawling

Stone Lake and the results are shown in Figure 21. This lake

ranks fifth in size among those included in the group. The

Juday & Birge — Oxygen in Wisconsin Lake Waters . 439

Fig. 20. The dissolved oxygen and the temperature of Fence Lake on

August 23, 1929. The quantity of oxygen was slightly above the satura¬

tion point in the upper 5 m.

water contains about 11 mg. of fixed carbon dioxide per liter

and the hydrogen ion readings in this series varied from pH

7.4 at the surface to pH 7.0 at 23 m. On August 10, 1927 the

quantity of oxygen amounted to about 75 per cent of satura¬

tion in the epilimnion, declined to about 35 per cent in the ther-

mocline and gave an average of about 32 per cent in most of

the hypolimnion, with a decrease to 12 per cent of saturation

at 23 m. The free carbon dioxide in this series rose to a maxi¬

mum of 9 mg. per liter of water at 23 m.

Thirty-two series of oxygen determinations covering the en¬

tire depth of Trout Lake were made between June 1925 and

the close of the field work in 1931. The number of series per

annum varied from 2 in the summer of 1930 to 9 in 1925. The

32 series consisted of 259 samples, or an average of 8 samples

for each series. Each year the first series was taken in late

June or early July and the last series was taken as late in

August as practicable; one series was obtained as late as Sep-

440

Wisconsin Academy of Sciences, Arts, and Letters.

Fig. 21. The dissolved oxygen and the temperature of Crawling Stone

Lake on August 10, 1927. Note that the hypolimnion contained a smaller

quantity of oxygen than that of Fence Lake, Fig. 20.

tember 21 in 1925. The various series were distributed in such

a way each year that they would give a general idea of the

changes that took place in the quantity of oxygen at different

depths during the summer period of stagnation.

With an area of 1683 ha., Trout Lake is the second lake in

size in the Highland Lake District of northeastern Wisconsin.

It is exceeded in size by Lac Vieux Desert which has an area

of 1934 ha., but which has a maximum depth of only 6 m. The

water of Trout Lake contains an average of 19 mg. of fixed

carbon dioxide per liter and the hydrogen ion readings during

the summer fall between pH 7.3 and pH 8.2 at the surface and

between pH 6.5 and pH 7.3 at the bottom. There is only a

small amount of vegetable stain in the water, the color read¬

ings varying from 6 to 14 on the platinum-cobalt scale. About

1 per cent of the sun’s energy that is delivered to the surface

of the lake reaches a depth of 8 m.

Figures 22 and 23 show the changes which took place in the

dissolved oxygen of Trout Lake during the summer stagnation

period in 1928 and in 1931. On June 24, 1928 the dissolved

Juday & Birge — Oxygen in Wisconsin Lake Waters. 441

Fig. 22. The dissolved oxygen and the temperature of Trout Lake on

June 24, 1928 and on August 25, 1928. Note the marked decrease in quan¬

tity of oxygen, especially in the lower water, during these two months.

Compare with Fig. 23.

oxygen in the upper 10 m. varied from 86 per cent to 91 per

cent of the amount required for saturation ; between 10 m. and

15 m. the quantity was between 86 per cent and 71 per cent of

saturation, and between 15 m. and 30 m. it varied from 71 per

cent to 60 per cent, with a minimum of 53 per cent at 31 m.

On August 25, 1928 the quantity of oxygen in the upper 10 m.

was between 85 per cent and 83 per cent of saturation ; between

10 m. and 15 m. it amounted to 83 per cent to 53 per cent and

between 15 m. and 30 m. it declined from 52 per cent at the

former depth to 28 per cent at the latter, with a minimum of

17 per cent at 32 m. Thus there was only about half as much

oxygen below 25 m. on August 25 as on June 24, but 2.1 mg.

442 Wisconsin Academy of Sciences , Arts, and Letters .

0 5 10 15 20 _ 25 30

p. If T? "

20% 40% 60% 80% 100%

Fig. 23. The dissolved oxygen and the temperature of Trout Lake on

July 1, 1931 and on August 27, 1931. Note the decrease of dissolved oxy¬

gen below 5 m. during this period. Compare with Fig. 22.

per liter was still present at 32 m. on August 25. This is the

largest amount that has been found in the bottom water in late

summer.

Figure 23 shows that the changes which took place during the

summer of 1931 differed somewhat from those noted in 1928.

In 1931, for example, the percentage of saturation on August

27 was somewhat higher at the surface than on July 1 and there

was a more marked decrease in the quantity of dissolved oxygen

below 25 m. in 1931 than in 1928. The decline in percentage

oetween 10 m. and 25 m., however, was substantially the same

in the two years. Only 0.6 mg. per liter of water remained at

31 m. on August 27, 1931. The smallest amount of oxygen that

has been found in the bottom water of Trout Lake in late sum¬

mer, was obtained on August 22, 1930, namely 0.3 mg. per liter

at a depth of 32 m. ; this represents a sevenfold difference be-

Juday & Birge — Oxygen in Wisconsin Lake Waters. 443

tween the maximum and minimum amounts found at the bot¬

tom in these summer observations covering a period of seven

years.

Oxygen in excess of the amount required for saturation was

noted in 8 of the 259 samples; 3 of these samples were taken

at the surface, 2 from 5 m., 1 from 8 m. and 2 from 10 m. The

surface, 5 m. and 8 m. samples showed an excess of oxygen in

the series taken on August 27, 1929. The maximum percentage

was observed at a depth of 5 m. on July 23, 1925, namely 115

per cent of saturation and the second highest, 110 per cent, was

found at 10 m. on the same date.

Black Oak Lake (Fig. 24) shows a more advanced stage in

the depletion of the dissolved oxygen in the bottom stratum in

late summer. The average quantity of fixed carbon dioxide in

the upper water in summer is between 9 mg. and 10 mg. per

liter and the hydrogen ion readings in this stratum vary from

pH 7.0 to pH 7.8. The color of the surface water ranges from

0 to 14 and the sun’s energy is reduced to 1 per cent of the

amount delivered to the surface of the lake at a depth of about

10 m.

Fig. 24. The dissolved oxygen and the temperature of Black Oak Lake

on August 24, 1928. There was a small excess of oxygen at 10 m. and

none below 22 m. Compare with Fig. 25.

444 Wisconsin Academy of Sciences , Arts , cmd Letters.

The epilimnion of Black Oak Lake yielded about 87 per cent

of the quantity of oxygen required for the saturation of the

water, but there was a slight excess in the thermocline (108

per cent at 10 m.) ; below 10 m. there was a rapid decrease in

the amount of oxygen, the quantity falling to about 1 mg. per

liter at a depth of 20 m. in the different series. By late August

the quantity of oxygen at 22 m. and below that depth ranged

from just a trace or none at all up to 0.2 mg. per liter. A

maximum of 11 mg. of free carbon dioxide per liter was found

at 25 m. on August 24, 1928.

Presque Isle Lake is one of the larger members of this group.

It has fairly hard water for this region, the fixed or bound

carbon dioxide amounting to 23 mg. to 25 mg. per liter in the

upper water. The color of the surface water ranged from 8 to

15 and the hydrogen ion from pH 7.7 to pH 8.5.

Fig. 25. The dissolved oxygen and the temperature of Presque Isle Lake

on August 11, 1927. Very little oxygen was found below 13 m. Compare

with Fig. 24.

Juday & Birge — Oxygen in Wisconsin Lake Waters. 445

The oxygen in the epilimnion on August 11, 1927 amounted

to only 68 per cent to 78 per cent of saturation (Fig. 25) ;

there was a rapid decrease in the quantity in the thermocline

and in the upper part of the hypolimnion, so that only 7 per

cent of saturation was left at a depth of 18 m. None was found

at 20 m. or below this depth. The entire hypolimnion, there¬

fore, contained only a small supply of dissolved oxygen. Free

carbon dioxide was abundant in the hypolimnion also, the

amount reaching a maximum of 11 mg. per liter at the bottom

in one series of samples.

Adelaide Lake is one of the small members of this group. Its

water is relatively soft, the upper water containing from 8 mg.

to 4 mg. of fixed carbon dioxide per liter. The hydrogen ion

concentration varied from pH 6.3 to pH 8.8 at the surface. The

color of the surface water ranged from 24 to 38 and the sun's

energy is reduced to about 1 per cent at a depth of 4 m.

The upper 3 m. of Adelaide Lake (Fig. 26) contained from

78 per cent to 80 per cent of the amount of oxygen required

Fig. 26. The dissolved oxygen and the temperature of Adelaide Lake on

August 21, 1926. Very little oxygen was found below 6 m. Compare with

Fig. 25.

446 Wisconsin Academy of Sciences , Arts, and Letters.

for saturation on August 21, 1926, but there was a very rapid

decrease in the amount between 3 m. and 6 m., with only a

trace left at 10 m. and none below this depth. More than half

of the maximum depth of the lake, therefore, was devoid of free

oxygen on this date. An abundant supply of free carbon di¬

oxide was found in the hypolimnion, with a maximum amount

of 19.5 mg. per liter of water at 20 m.

Lake Mary (Fig. 27) shows the most extreme case of oxygen

depletion among the lakes of Group IV. This lakelet has an

area of slightly more than one hectare, but the maximum depth

is 22 m. It is well protected from wind, so that only a very thin

upper stratum is kept in circulation during the summer. The

upper water contains less than 4 mg. of fixed carbon dioxide per

liter; the hydrogen ion ranges from pH 6.0 to pH 6.2 at the

surface and from pH 5.7 to pH 5.9 at the bottom. The color

of the surface water ranged from 100 to 123 on the platinum-

cobalt scale and the sun’s energy is reduced to 0.5 per cent at

a depth of only 2 m.

The upper meter was the only stratum that was well aerated

on August 29, 1929 (Fig. 27). The quantity of oxygen at the

Fig. 27. The dissolved oxygen and the temperature of Lake Mary on

August 29, 1929. Very little oxygen was found below 3 m. Strongly

eutrophic in character.

Juday & Birge — Oxygen in Wisconsin Lake Waters . 447

surface was equivalent to a little more than 90 per cent of the

amount required for saturation and that at 1 m. was 87 per

cent of saturation ; the sample at 2 m. contained 33 per cent of

the amount necessary for saturation and that at 3 m. less than

2 per cent. In all of the series taken on Lake Mary, very little

or no dissolved oxygen was obtained below a depth of 3 m. Ob¬

servations made on May 7, 1929 indicate that the vernal over¬

turning and circulation of the water are not complete, so that

the oxygen deficiency in the summer is due in part to the winter

stagnation and in part to the summer stagnation as a result

of the incomplete mixture of the water in the spring.

The phenolphthalein titrations of samples from the lower

water represent a large amount of free carbon dioxide in that

stratum; the titrations of the bottom samples of the various

series, for example, represented from 32 mg. to 52 mg. of free

carbon dioxide per liter. Whether all of this acidity is due to

free carbon dioxide, or whether part of it is due to free carbon

dioxide and part to organic acids has not been determined up

to the present time.

Attention may be called to the unusual character of the tem¬

perature curve in Lake Mary (Fig. 27). The coldest stratum

of water was found at an intermediate depth and not at the

bottom in summer. Temperature readings of 4.0° C. were ob¬

tained at 10 m. and 12 m. on August 29, 1929. Similar results

were obtained in 4 other sets of temperature readings taken in

Lake Mary during the months of July and August between

1926 and 1928, inclusive. In each of these 5 series the coldest

water was found between 8 m. and 12 m. The readings at the

bottom (22 m.) were from half a degree to three-quarters of a

degree higher than the minimum found in the 8 m.-12 m.

stratum.

General Discussion

Sources and Distribution of Oxygen

Lakes receive their supplies of dissolved oxygen from two

primary sources, namely, (1) from the atmosphere and (2)

from the photosynthetic activities of the aquatic plants which

populate their waters. The former may be regarded as the

more important source, particularly with reference to supply¬

ing an entire lake with dissolved oxygen when its water is in

448 Wisconsin Academy of Sciences, Arts, and Letters .

complete circulation. Considerable amounts of oxygen may be

obtained from photosynthesis, as illustrated in the diagram for

Pallette Lake (Fig. 13), but the quantity obtained from this

source depends upon the abundance of the aquatic plants and

upon the extent of their activities. They are usually most

abundant from spring to early autumn, so that they make their

largest contributions of oxygen during this part of the year.

A certain amount of oxygen is liberated by bacteria in the

process of denitrification, which is carried on under certain con¬

ditions, but the quantity derived from this source is too small

to play any important role in the household economy of a lake.

In lakes of the temperate class, such as those of northeastern

Wisconsin, the dissolved oxygen is distributed throughout the

lake chiefly by means of the general movements of the water.

These bodies of water are subject to four seasonal changes each

year, namely, spring and autumn periods of overturning and

circulation which are separated by summer and winter periods

of stratification and stagnation. In the process of overturning

and circulation the whole body of water is freely exposed to

the air at the surface of the lake, so that a general aeration of

the water takes place at these times ; the quantity of dissolved

oxygen becomes substantially uniform from surface to bottom.

During the summer and winter periods of stratification, how¬

ever, there is no complete circulation of the water and the oxy¬

gen supply of the lower water is limited to the amount that it

possesses at the time that stratification takes place. Any de¬

crease in the quantity of dissolved oxygen in the lower stratum

when the water is thermally stratified remains as a deficiency

until the subsequent period of overturning and circulation. A

certain amount of oxygen may be transferred by diffusion, but

this process goes on so slowly in water that it may be disre¬

garded as a factor in the transference of oxygen from one

stratum to another.

Classification of Lakes

The biological processes that take place within a lake make

a constant demand upon the supply of dissolved oxygen therein.

Most of the aquatic organisms require free oxygen for their

respiration and the decomposition of organic matter at all

depths consumes an additional amount. The extent of the con-

Juday & Birge— Oxygen in Wisconsin Lake Waters. 449

sumption of the dissolved oxygen in the lower stratum or hypo-

limnion of a lake during the summer period of stratification

depends upon various physical, chemical and biological factors.

The preceding diagrams (Figs. 3 to 27) show that only a rela¬

tively small amount of the oxygen supply of the lower stratum

is consumed in some lakes, while in others there is very little

dissolved oxygen left in this stratum by the latter part of

August.

Thienemann (1921) has divided lakes into three general

types on the basis of the chemical and biological conditions that

obtain in the hypolimnion in late summer, and the quantity of

dissolved oxygen in this stratum is an important factor in his

system of classification. Naumann (1921) proposed a more

elaborate system of classification in which the three main types

were divided into a number of subclasses, but these northeast¬

ern Wisconsin lakes show such a great diversity of character¬

istics that it is not worth while to attempt a classification of

them beyond the three general types; in fact, it is doubtful

whether a separation into more than two general types has any

particular value.

These three general types are designated as oligotrophic, eu-

trophic and dystrophic lakes. Oligotrophic lakes possess a good

supply of dissolved oxygen at all depths during the two periods

of stagnation, even including the hypolimnion in late summer.

They receive relatively small amounts of organic matter from

their drainage basins and their waters do not produce an un¬

usual amount of such material, so that the demand for dissolved

oxygen in the process of decomposition is not great enough to

consume all of the free oxygen even at the bottom. Their wa¬

ters are transparent owing to the scarcity of plankton and other

suspended material. Crystal Lake (Fig. 17) is an outstanding

example of this type of lake; Big Carr (Fig. 18) and Fence

(Fig. 20) also belong in this group. Trout Lake (Fig. 23) may

be regarded as a member of this group, but it lies near the low¬

er limit of this type of lake on account of the small quantity of

oxygen found below a depth of 30 m. in late August of some

years.

Eutrophic lakes are characterized by a marked paucity of

dissolved oxygen in the lower water in late summer. In lakes

of this type the physical and chemical conditions are favorable

450 Wisconsin Academy of Sciences, Arts, and Letters.

for the production of a large amount of organic matter and

the decomposition of this material uses up the free oxygen in

some or frequently in practically all of the hypolimnion in sum¬

mer. Owing to the abundance of plankton and other suspended

material, the water has a low degree of transparency. Exam¬

ples of this type are found in Long (Fig. 10), Nebish (Fig.

16), Black Oak (Fig. 24) and Presque Isle (Fig. 25) lakes.

Dystrophic lakes are characterized by their brown colored

waters. The color is due to the presence of vegetable extrac¬

tives (humic substances) which are derived from peat bogs or

from marshes that are tributary to such lakes. These stains

give their waters a low degree of transparency. The European

lakes on which the definition of this type is based, are said to

be poor in phytoplankton as well as in electrolytes, especially in

phosphorus and calcium, and in available nitrogen compounds.

Both Thienemann and Naumann state that the waters of the

European dystrophic lakes range from yellow to brown in col¬

or, but they do not give any definite standard of color for the

minimum limit of the group. Neither are any definite limits

given by Lonnerblad (1931) who has made a special study of

the dystrophic lakes in the vicinity of Aneboda, Sweden.

During this investigation color readings have been made on

the waters of 530 lakes situated in northeastern Wisconsin and

the colors ranged from zero up to 340 on the platinum-cobalt

standard. The waters of two lakes gave readings of 314 and

340, respectively, 11 fell between 200 and 300 and 61 between

100 and 200, making a total of 74 lakes with color readings of

100 or more; an additional 18 lakes gave readings between 90

and 99. It will probably be best, however, to limit the present

consideration to those lakes with color readings of 100 or more

on the platinum-cobalt scale, although those with readings as

low as 60, or even 50, show the brown color distinctly in the

lake itself. Lake waters having colors of 100 to 120 resemble

weak tea in appearance, so that there is no question whatever

about an abundant supply of humic substances in them ; in fact

100 is probably too high rather than too low for the minimum

limit of these brown colored waters.

With respect to the electrolytes found in these 74 Wisconsin

lakes having colors of 100 or more, the conductivity or specific

conductance varied from a minimum of 11 up to a maximum

of 81 when expressed in terms of reciprocal megohms ; this rep-

Juday & Birge — Oxygen in Wisconsin Lake Waters . 451

resents more than a sevenfold difference. A conductivity of 11

reciprocal megohms indicates a very soft water, with few elec¬

trolytes in solution, while one of 81 reciprocal megohms indi¬

cates that the water has a fair degree of hardness, especially

in the region in which these lakes are situated. In the former

lake, for example, the calcium amounted to 0.6 mg. per liter

and in the latter to 11.2 mg. A maximum of 12.2 mg. of calci¬

um per liter was found in one of these brown water lakes with

a color of 118.

The phosphorus content of these brown waters was substan¬

tially as large as in the lakes with color readings below 20. The

quantity of soluble phosphorus ranged from a trace up to 0.012

mg. per liter of water, with an average of 0.008 mg. per liter

for 62 of the 74 lakes; no soluble phosphorus determinations

were made on the other 12 lakes. The quantity of nitrate nitro¬

gen, also, was as large in these brown water lakes as in those

having little or no color ; it varied from a minimum of 0.01 mg.

to a maximum of 0.045 mg. per liter. Likewise the hydrogen

ion concentration showed about the same range in the brown

water lakes as in those with little or no stain, varying from

pH 4.9 to pH 9.1. The waters of 24 of these lakes were near

the neutral point, the readings falling between pH 6.8 and pH

7.2, inclusive.

The vegetable extractives which produce the brown color in

these Wisconsin lakes are chiefly carbon compounds, so that

these humic substances increase the organic carbon content of

the water in a marked degree. The quantity of organic carbon

in these brown waters varied from a minimum of 9.2 mg. to a

maximum of 25.8 mg. per liter; the organic nitrogen content

ranged from 0.88 mg. to 1.75 mg. per liter, so that the carbon-

nitrogen ratio varied from 10 to 36. The lake waters in which

the brown color fell between 0 and 20 on the platinum-cobalt

scale yielded from 1.3 mg. to about 8.0 mg. of organic carbon

per liter and had carbon-nitrogen ratios of 8 to 15. There is

thus some overlapping of the carbon-nitrogen ratio at the low¬

er limit of the high colored waters and at the upper limit of

those with little or no brown color and this is accounted for by

the fact that some of the high colored lakes contained large

crops of phytoplankton in addition to the humic substances. This

plankton material was rich in organic nitrogen, so that the car-

452 Wisconsin Academy of Sciences , Arts , and Letters .

bon-nitrogen ratio in them was brought down below the upper

limit of that in lakes having little or no stain in their waters.

The European lakes with high colored waters are said to be

plankton poor, but this is not true of a considerable number of

the high colored lakes of northeastern Wisconsin. In the latter

the centrifuge phytoplankton in the upper water ranged from

2300 to 5900 cells and colonies per cubic centimeter in 6 lakes

with color readings of 200 or more, and from 2100 to 11,800

cells and colonies per cubic centimeter in 18 of the 62 lakes

having color readings between 100 and 200. Lakes that sup¬

port phytoplankton crops of this size can hardly be regarded

as plankton poor. In the other 59 members of this group of

lakes having brown waters, the number of centrifuge phyto¬

plankton organisms ranged from 300 to 2000 cells and colonies

per cubic centimeter. The number fell below 1000 per cubic

centimeter in 17 of the 74 lakes; these may be regarded as

rather poor in plankton. Little Long Lake (Fig. 12) yielded

the minimum number of 300 cells and colonies per cubic centi¬

meter in the upper 5 m.

The phytoplankton crop of very few of the lakes having little

or no stain in their waters exceeded that of the more produc¬

tive ones in the high colored group. In Trout Lake, with a col¬

or not exceeding 14 on the platinum-cobalt scale, the centrifuge

phytoplankton in 23 catches from the upper 5 m. gave an aver¬

age of 1200 cells and colonies per cubic centimeter of water. In

Black Oak Lake, the average was 1260 per cubic centimeter in

3 series, in Muskellunge Lake 1900 in 8 series and in Little

St. Germain Lake 2480 in 4 surface samples taken in different

years. Mann Lake, with a color ranging between 20 and 34,

yielded the largest number of phytoplankton organisms, name¬

ly 40,500 cells and colonies per cubic centimeter of water on

August 14, 1928 ; a second maximum of 12,300 per cubic centi¬

meter was obtained from this lake on August 26, 1927.

Lonnerblad (1931) states that some of the dystrophic lakes

which he studied, were oligotrophic and some eutrophic in char¬

acter in so far as the dissolved oxygen was concerned. The

same is true of the brown water lakes of northeastern Wiscon¬

sin. Little Long Lake (Fig. 12) contained a good supply of

dissolved oxygen throughout the hypolimnion on August 22,

1931, so that it possessed one of the important characteristics

of an oligotrophic lake. Dead Pike Lake (Fig. 19) has not

Juday & Birge — Oxygen in Wisconsin Lake Waters . 453

been included in the brown water group, but its water has a

distinct brown color, with readings of 60 to 70, and its lower

water was well supplied with free oxygen in late summer.

Lake Mary (Fig. 27), on the other hand, is an extreme case

of the eutrophic type. Three small lakes in the same vicinity,

namely Helen, Rose and Yolanda, which have highly colored

waters with readings ranging from 95 to 160, are also marked¬

ly eutrophic in character. These 4 lakelets may perhaps be

regarded as typical dystrophic lakes, yet they support fairly

large crops of phytoplankton. The upper water of Lake Mary

yielded from 1000 to more than 6000 phytoplankton cells and

colonies per cubic centimeter, while the other 3 were not so

productive, yielding from 800 to 4300 phytoplankton organisms.

The organic carbon content of these 4 lakelets ranged from a

minimum of 12 mg. in Helen to a maximum of 17 mg. per liter

in Mary. These results may be compared with those of Trout

Lake where the color does not exceed 14 and where the or¬

ganic carbon averages about 4 mg. per liter of water.

In view of the marked chemical and biological differences

between the brown colored lakes of northeastern Wisconsin and

those of Europe, the separation of this group from the other

two types is of very doubtful value in the case of the Wiscon¬

sin lakes.

Oxygen Deficit

Thienemann (1928) computed the total oxygen content of a

number of lakes from series of oxygen determinations and dis¬

cussed the relation between the quantity in the hypolimnion

and that in the epilimnion. He also considered the relation of

the total quantity of oxygen found in a lake to the amount re¬

quired to saturate the water at all depths; the difference be¬

tween these two amounts is a measure of the deficiency or ex¬

cess of oxygen in the different strata of a lake. The vertical

distribution of the oxygen and the deficits and excesses at the

various depths are given in Table III for some of the lakes of

northeastern Wisconsin. These data enable one to compute the

total quantity of oxygen present when the volume of the lake

is known. The results of such computations are given in Table

IV. The quantity of oxygen required to saturate the water at

the different temperatures is taken from the table published

by Birge and Juday in 1914; the values given in this report

454 Wisconsin Academy of Sciences , Arts, and Letters.

were based upon those given by Fox (1907) and they are some¬

what higher than those published by Birge and Juday in 1911.

The total amount of oxygen in a given stratum of water has

been determined by multiplying the mean quantity of oxygen

per cubic meter in that stratum by the volume of the stratum;

the amount required for the saturation of the stratum has also

been determined in the same manner.

It will be noted in this table that the quantity of oxygen ac¬

tually found at the different depths in the various series was

below the theoretical point of saturation except in a few cases

where the oxygen liberated in the process of photosynthesis in

the region of the thermocline brought the amount up to the

saturation point or higher. As already pointed out the defi¬

ciency is due chiefly to the demand for oxygen in the respira¬

tory processes of aquatic organisms and in the decomposition

of organic matter. It is due in part also to the elevation of

these lakes above sea level. The theoretical amount required

for saturation is based upon an atmospheric pressure of 760

mm. at sea level, but the lakes of northeastern Wisconsin are

about 500 m. above sea level, so that they receive a correspond¬

ingly smaller atmospheric pressure. Thus the oxygen tension

in the air and that in the lake water come to an equilibrium

below the amount required for saturation at sea level ; the sat¬

uration point for an altitude of 500 m. is a little more than

94 per cent of that at sea level.

Table IV shows the quantity of oxygen in the several strata

of these lakes as well as the total amount in each of the lakes.

It also shows the quantity of oxygen required for the satura¬

tion of the strata and of the entire lake. In only two of the

lakes was there an actual excess of oxygen in any of the strata ;

the 6 m.-lO m. stratum of Pallette Lake had 16 per cent more

than the quantity required for saturation and the 8 m.-lO m.

stratum in Weber Lake yielded a small excess of oxygen.

The data given in Table IV are summarized in Table V, with

the addition of the absolute deficit of oxygen for comparison

with the actual deficit and also a column showing the relation

between the quantity of oxygen in the hypolimnion and that in

the epilimnion. In 1929 Alsterberg suggested that the summer

oxygen deficit may be considered from two standpoints, namely

(1) that of the actual deficit and (2) that of the absolute de¬

ficit. The actual deficit is the difference between the quantity

Juday & Birge — Oxygen in Wisconsin Lake Waters . 455

of oxygen present at a certain depth and the amount required

to saturate the water at the temperature observed at that depth ;

the absolute deficit represents the difference between the quan¬

tity of oxygen found at any depth and the amount required to

saturate the water at a temperature of 4° C., which is 9.26 cc.

or 13.23 mg. per liter minus a correction for the elevation of the

lake above sea level. The lakes of northeastern Wisconsin have

an elevation of about 500 m., so that the quantity of oxygen

required to saturate the water at 4° at this altitude is approxi¬

mately 12.5 mg. per liter without any correction for tempera¬

ture and humidity. Since the temperature of the water in the

lakes of northeastern Wisconsin is above 4° in summer, the

oxygen deficit on the absolute basis is always larger than that

on the actual basis.

The actual percentage of saturation, when the entire lake is

considered, ranges from a minimum of 16 per cent in Lake

Mary to a maximum of almost 95 per cent in Weber Lake.

Adelaide Lake ranks second in the minimal percentage of actual

deficiency, with approximately 43 per cent of saturation. Two

of the lakes are above 90 per cent of actual saturation and 5

fall between 80 per cent and 90 per cent ; thus 7 of the 14 lakes

included in the table have more than 80 per cent of the quan¬

tity of oxygen required for actual saturation.

The percentage of saturation on the absolute basis ranges

from 14 per cent in Lake Mary to 70 per cent in Weber Lake.

Only 5 of the 14 lakes are above 60 per cent on this basis, while

5 are between 50 and 60 per cent.

The ratio of the quantity of oxygen in the hypolimnion to that

in the epilimnion, as shown in the last column of Table V, de¬

pends upon the relation between the volumes of these two strata

and also upon the quantity of oxygen in them. Two of the

lakes, namely Diamond and Finley, do not have a well marked

hypolimnion and no ratios are given for them. The thickness

of the epilimnion in the other lakes ranges from a minimum

of 2 m. in Lake Mary to a maximum of 10 m. in Black Oak and

Crystal. Differences in the thickness of the epilimnion in the

various lakes, together with differences in amount of oxygen in

the epilimnion and hypolimnion, produce wide differences in

the ratio of the oxygen content of the latter to that of the

former stratum. These ratios range from a minimum of 0.08

in Bragonier Lake to a maximum of 1.22 in Big Carr Lake,

456 Wisconsin Academy of Sciences , Arts , and Letters .

which represents a fifteenfold difference. The low ratio in Web¬

er Lake is due to the small volume of the hypolimnion as com¬

pared with that of the epilimnion.

Oxygen Gradient

Maucha (1931) has recently presented a discussion of the

vertical distribution of dissolved oxygen in lakes from a new

standpoint. In his computations he takes into account the at¬

mospheric pressure corrected for the elevation of the lake, the

vapor pressure, and the hydrostatic pressure at the different

depths. Three formulae are involved in these computations.

The first formula relates to the determination of the general

barometer reading at the elevation of the lake in question.

1. h = 18400 (1 + 0.004t) X (log 760 - log bx).

In this formula h is the elevation of the lake in meters, t the

temperature of the surface water in degrees centigrade and

b1 the barometer reading in millimeters.

The following formulae are used for the computation of the

saturation values at the various depths:

o2 b + mp — 760

2. g = — X - - X 100

o'2 b + mp — f

b + mp — 760 b — f

3. g' = - — X - - - X 100

b + mp — f 760 — f

In these two formulae, o2 is the dissolved oxygen present at a

given depth, o'2 the amount of oxygen required to saturate the

water at the observed temperature, b the general barometer

reading at the elevation of the lake expressed in millimeters,

m the depth of the water in meters, p the hydrostatic pressure

of 1 m. of water (73.5 mm.) and / the vapor pressure of sat¬

urated air at the temperature of the water at the various

depths. The pressure of saturated aqueous vapor at different

temperatures is given in the Smithsonian Physical Tables.

While hydrostatic pressure is taken into account in these com¬

putations, it does not play any important role in reality except

where the water obtains a supply of oxygen from the photo¬

synthetic activities of plants. In general the summer supply of

oxygen for the hypolimnion is obtained from the atmosphere at

Juday & Birge ■ — Oxygen in Wisconsin Lake Waters . 457

Fig. 28. The oxygen saturation values of Crystal Lake on August 21,

1928. The vertical spaces show the depth in meters and the horizontal

spaces indicate the saturation values. The solid line indicates the actual

saturation value and the broken line the theoretical value. Note that the

former exceeds the latter at 10 m. and 12 m. Compare with Fig. 17.

the surface of the lake during the vernal period of circulation,

so that hydrostatic pressure does not play any part in determin¬

ing the amount of oxygen that is absorbed by the water at this

time. When this aerated water is carried down to a depth of

80 m., for example, in the process of circulation, where the hy¬

drostatic pressure is equivalent to about 3 atmospheres, no more

oxygen is absorbed at that depth because there is no extra sup¬

ply from which the water may obtain oxygen. Any oxygen

liberated in the process of photosynthesis at a depth of 10 m.,

for instance, is readily retained in that stratum if the water

is not disturbed by circulation, because the hydrostatic pres¬

sure at that depth is equivalent to about one atmosphere thus

making a total pressure of two atmospheres at that depth.

The results obtained for 6 of the lakes of northeastern Wis¬

consin are given in Table VI and in Figures 28 to 88. The

column marked g in this table represents the actual oxygen

saturation values as found in the series of determinations, while

458 Wisconsin Academy of Sciences, Arts, and Letters .

Fig. 29. The oxygen saturation values of Pallette Lake on August 22,

1928. The actual value (solid line) exceeds the theoretical value (broken

line) between 7 m. and 10 m. Compare with Fig. 13 and Fig. 28.

g' represents the values for water that is saturated with oxygen.

These oxygen values have been designated as oxygen gradients

by Maucha, or simply gradients. The difference between these

two gradients shows to what extent the oxygen supply of the

water is affected by the biological processes that take place

within a lake.

In Crystal Lake (Fig. 28) the quantity of oxygen found in

the upper 8 m. on August 21, 1928 was a little less than that

shown by the theoretical gradient, but it exceeded the latter in

the 10 m.-12 m. stratum and then fell below the theoretical

value between 15 m. and 19 m. Pallette Lake (Fig. 29) yielded

similar results on August 22, 1928, but there was a larger ex¬

cess of oxygen in the 7 m.-lO m. stratum and a much more

marked decrease of oxygen below 10 m. The actual and theo¬

retical oxygen gradients in Silver Lake (Fig. 80) are so near

each other in the upper 6 m. that they can not be platted sepa¬

rately; an excess of oxygen is shown between 7 m. and 10 m.,

with a marked decline in the quantity of oxygen below the lat¬

ter depth. The excess oxygen in these three lakes was due to

Juday & Birge — Oxygen in Wisconsin Lake Waters . 459

Fig. 30. The oxygen saturation values of Silver Lake on August 28,

1931. The actual value (solid line) exceeds the theoretical value (broken

line) between 7 m. and 10 m. Compare with Fig. 29.

Fig. 31. The oxygen saturation values of Muskellunge Lake on August

26, 1931. This is a typical eutrophic lake. Note the large difference be¬

tween the actual value (solid line) and the theoretical value (broken line)

below 7 m. Compare with Fig. 28.

460 Wisconsin Academy of Sciences, Arts, and Letters .

the photosynthetic activities of the various aquatic plants. Mus-

kellunge Lake (Fig. 31) shows the eutrophic type of dissolved

oxygen gradient. The two gradients are substantially the same

down to a depth of 7 m., but the actual gradient shows a marked

decline below this depth and falls to zero at 15 m. The irregu¬

larity in the actual gradient curve between 7 m. and 9 m. is

similar to that shown by Maucha for the lower water of Grosser

Ploner See.

Figures 32 and 33 show the change that took place in the

oxygen supply of Trout Lake between June 24 and August 25,

1928. The curves indicate a marked decrease in the oxygen

supply below a depth of 5 m. during this period of two months ;

the greatest decline was found below 20 m.

Fig. 32. The oxygen saturation values of Trout Lake on June 24, 1928.

The actual value (solid line) is below the theoretical value (broken line)

at all depths. Compare with Fig. 33 and Fig. 22.

Fig. 33. The oxygen saturation values of Trout Lake on August 25,

1928. Compare the difference between the actual value (solid line) and

the theoretical value (broken line) in this figure with that in Fig. 32. See

Fig. 22.

Juday & Birge — Oxygen in Wisconsin Lake Waters. 461

The general status of the oxygen of the hypolimnion is indi¬

cated by dividing the sum of the actual saturation values in

that stratum by the sum of the theoretical values for that

stratum; such results are given in Table VII where the ratio

of the actual to the theoretical values range from 92.2 in Crys¬

tal Lake to 11.5 in Muskellunge Lake. The former is oligo-

trophic and the latter eutrophic in character. Maucha’s compu¬

tations show a maximum ratio of 102.1 in Seneca Lake, New

York. No other ratio given in his tables is as high as that of

Crystal Lake.

II. Oxygen Consumed

The determinations of oxygen consumed or oxygen absorbed

were made by the usual method of adding sulphuric acid and

potassium permanganate to the samples and digesting them

for 30 minutes in a bath of boiling water; ammonium oxalate

was then added to the digested samples and they were titrated

with potassium permanagate. In this procedure only the more

readily oxidized carbon is affected, so that the results show only

a part of the organic carbon that is present in the water ; it is

a fairly large proportion of the organic carbon, however.

Determinations of oxygen consumed were made on 509 sam¬

ples of lake water during the summers of 1929, 1930 and 1931.

Of this number 365 were surface samples obtained from 290

different lakes; 144 samples represent 28 series taken on 21

different lakes. The results obtained on some of the lakes are

given in Table VIII. In the surface samples the quantity of

oxygen consumed ranged from a minimum of 1.2 mg. in Dor¬

othy Dunn Lake to a maximum of 34.5 mg. per liter in Helmet

Lake. Only 4 samples showed less than 2.0 mg. of oxygen con¬

sumed per liter of water and the maximum number of samples,

namely 52, fell between 4.0 mg. and 4.9 mg.; there were 46

samples in the 3.0-3.9 mg. group and 45 in the 5.0-5.9 mg.

group. In 283 surface samples, or 77 per cent of the total

number, the oxygen consumed varied between 2.0 mg. and 10.0

mg. per liter; the amount exceeded 17.0 mg. in 20 samples, but

only 4 exceeded 25.0 mg.

The majority of the lakes from which these surface samples

were secured have waters that are more or less deeply stained

by humic substances derived from peat bogs and marshes ; some

462 Wisconsin Academy of Sciences , Arts , and Letters.

of them, however, do not show any trace of brown color. Thus

the brown color in the various samples ranged from zero in

several of them up to 268 in Helmet, which is a small bog lake.

The color readings in 62 samples did not exceed 9 on the plati¬

num-cobalt scale; in such waters the brown color is not per¬

ceptible in the lake itself. The largest number of samples in

the different color groups, namely 93, gave readings between 10

and 19, while 59 samples were between 20 and 29; thus 206

surface samples, or 56 per cent of the total number, possessed

only a comparatively small amount of stain or none at all. The

readings for 71 samples fell between 30 and 49 and these wa¬

ters show a more or less distinct brown color, especially those

with readings above 40; 88 of the surface samples gave color

readings of 50 or more.

The relation between the color of the water and the quantity

of oxygen consumed is shown in Table IX, which includes 354

of the surface samples ; 20 samples gave color readings varying

from 150 to 340, but determinations of oxygen consumed were

made on only 11 of them. These 11 samples are not included

in the table because they are distributed over such a wide color

range. The samples are grouped at color intervals of 10 up to

59, but above the latter group it is necessary to combine them

into larger color groups in order to obtain a fair mean.

There are rather wide differences between minimum and

maximum amounts of oxygen consumed in the various color

groups, but the means for the different groups show a gradual

increase in the quantity of oxygen consumed with increasing

0 10 20 30 40 50 60 70 80 90 100 110 120 130

Fig. 34. The relation between the color of the water and the quantity of

oxygen consumed in 354 surface samples. The vertical spaces represent

the number of milligrams of oxygen consumed per liter of water and the

horizontal spaces show the color of the water based on the platinum-cobalt

standard. See Table IX.

Juday & Birge — Oxygen in Wisconsin Lake Waters. 463

color; the mean quantity rose from 3.7 mg. per liter in the 0-9

color group to 15.9 mg. in the 110-149 group. The maximum

amount of oxygen consumed was noted in the surface water

of Helmet Lake, namely 34.5 mg. per liter with a color of 260 ;

the next in rank was Red Bass Lake where the oxygen con¬

sumed was 31.6 mg. per liter and the color 166.

The correlation of color and oxygen consumed is shown

graphically in Figure 34 in which the mean quantity of oxy¬

gen consumed in the different color groups has been platted at

middle of each group. The curve shows a steady rise from the

0-9 group up to the 50-59 group; beyond the latter group the

rise is not so marked, but it continues up to the limit of the

last group.

Quantitative determinations of the organic carbon were made

on 282 of the surface samples that were analyzed for oxygen

consumed. The results obtained for a considerable number of

these samples are given in Table VIII. The organic carbon in

these 282 surface samples varied from a minimum of 1.2 mg,

to a maximum of 25.3 mg. per liter of water. Figure 35 repre¬

sents the mean quantity of organic carbon platted against the

mean quantity of oxygen consumed in the different groups of

samples. The curve shows a fairly regular increase in the

amount of oxygen consumed correlated with the increase in the

amount of organic carbon found in the samples. In general an

increase of 1.0 mg. of organic carbon per liter is correlated with

an increase of approximately 0.9 mg. of oxygen consumed per

liter. The individual samples, however, show a certain amount

of variation ; in some of them the amount of oxygen consumed

is larger than that of the organic carbon, in others the reverse

is true, and in still others the two quantities are about the

same.

In the lower part of the curve, where the amount of oxygen

consumed and that of organic carbon do not exceed 8.0 mg.

per liter, the mean quantity of the former is somewhat larger

than that of the latter, but above the 8.0 mg. point the two are

more evenly balanced. The general mean of oxygen consumed

in the 282 samples represented in this diagram is 9.0 mg. and

that of organic carbon is 8.7 mg. per liter, so that the varia¬

tions tend to balance each other. The fact that the mean quan¬

tity of oxygen consumed and that of organic carbon are so

nearly the same in these surface samples indicates that about

464 Wisconsin Academy of Sciences, Arts, and Letters.

Pig. 85. The relation between the amount of oxygen consumed and the

quantity of organic carbon in 282 surface samples. The vertical spaces

show the number of milligrams of oxygen consumed per liter of water and

the horizontal spaces indicate the number of milligrams of organic carbon

per liter of water.

40 per cent of the organic carbon in them, on an average, is

oxidized by the potassium permanganate, since two atoms of

oxygen are required to oxidize one atom of organic carbon.

The relation of the quantity of oxygen consumed to that of

the dissolved oxygen present in the surface samples shows

marked differences in the different lakes. In those lakes show¬

ing a relatively small amount of oxygen absorbed, the dissolved

oxygen is in excess of the oxygen consumed; examples of this

relation are given in Table VIII, such as the surface samples of

Black Oak, Clear, Crystal, Day, Fence, Trout and Weber lakes.

In other lakes the quantity of dissolved oxygen is substantially

Juday & Birge — Oxygen in Wisconsin Lake Waters . 465

the same as that of the oxygen consumed; the surface samples

of Adelaide, George and Little Papoose lakes are representa¬

tives of this type. In the more highly .colored waters, on the

other hand, the quantity of oxygen consumed is much larger

than that of the dissolved oxygen ; Black, Findler, Helen, Mary,

Summit and Yolanda lakes are good examples of this group.

The humic substances in the colored waters give them a larger

oxygen consuming capacity and as a result there is a greater

demand for dissolved oxygen in them than in the waters that

have little or no stain. This increased demand for free oxygen

usually keeps the quantity of dissolved oxygen well below the

saturation point in these colored waters. In Black Lake, for

example, the percentage of oxygen saturation in the surface

water was only 69 per cent, in Circle Lily Lake 65 per cent, in

Findler Lake 62 per cent, and in Summit and Yolanda lakes

72 per cent.

Three general types of vertical distribution of the oxygen

consumed have been found in the northeastern lakes. In one

group the quantity of oxygen consumed was substantially the

same from surface to bottom. Such results are shown in Table

VIII for Anderson, Bragonier, Long and Nokomis lakes. The

second type is represented by lakes that show a smaller amount

of oxygen consumed at the bottom than at the surface; Black

Oak and Trout lakes belong to this class. In the third type the

quantity of oxygen consumed is larger at the bottom than at

the surface; George, Mary and Papoose lakes are representa¬

tives of this class. The vertical distribution of the organic car¬

bon is similar to that of the oxygen consumed in Anderson

Lake; that is, the quantity is substantially uniform from sur¬

face to bottom, but there are more or less marked differences in

the vertical distribution of the two in the other lakes. These

differences are more prominent in the lower than in the upper

water.

Figure 34 shows that there is a close correlation between the

color and the amount of oxygen consumed in the surface sam¬

ples and similar results were obtained in two of the series ; in

Lake George there was a marked increase in the amount of

oxygen consumed at the bottom which corresponded to a five¬

fold rise in the color of the water and a similar change was

noted in the bottom water of Papoose Lake. (See Table VIII) .

The increase in the color of the water in these two lakes may

466 Wisconsin Academy of Sciences , Arts , and Letters.

be attributed to the presence of vegetable extractives at these

depths. Very different conditions were noted in the lower wa¬

ter of three other lakes, however. In Anderson Lake the color

rose from zero at the surface to 180 and 240, respectively, at

15 m. and 18 m., yet there was no corresponding increase in

the oxygen consumed. This lack of correlation is due to the

fact that the color of the lower water was due to the presence

of iron; 3.0 mg. of iron per liter were found at 15 m. and 7.0

mg. per liter at 18 m. in this series of samples. A similar in¬

crease in the color of the lower water without a corresponding

increase in the oxygen consumed was noted in Mary and Noko-

mis lakes. The series of samples from Nokomis Lake yielded

1.2 mg. of iron per liter at 18 m. and 2.2 mg. at 20 m., so that

iron was the factor involved in the increased color of the lower

water of this lake. No iron determinations were made on the

series of samples from Lake Mary, but it seems probable that

it was responsible for most of the increase in the color of the

lower water of this lake also.

Summary

1. Quantitative determinations of dissolved oxygen were

made on the waters of 510 lakes and lakelets situated in north¬

eastern Wisconsin.

2. The quantity of dissolved oxygen in the surface samples

ranged from 3.4 mg. to 12.4 mg. per liter of water, or from 34

per cent to 129 per cent of saturation.

3. The smallest amounts of oxygen in the surface waters

were found in bog lakes and lakelets.

4. Some of the lakes had an abundance of dissolved oxygen

at all depths during the summer period of stratification (oligo-

trophic lakes), while the lower stratum of others was charac¬

terized by a marked deficiency of oxygen in summer (eutrophic

lakes). (Figs. 17-27).

5. Excess oxygen was found in the thermocline of a few of

these lakes (Fig. 13).

6. The oxygen gradients showed striking differences in the

different types of lakes. (Figs. 28-33).

7. Determinations of oxygen consumed were made on the

waters of 290 lakes and lakelets.

Juday & Birge — Oxygen in Wisconsin Lake Waters. 467

8. The quantity of oxygen consumed varied from 1.2 mg. to

34.5 mg. per liter of water.

9. The quantity of oxygen consumed was correlated rather

closely with the amount of vegetable extractives or humic sub¬

stances in the water which give it a brown color. (Fig. 34) .

10. The quantity of oxygen consumed was also closely corre¬

lated with the amount of organic carbon present in the water.

(Fig. 35).

11. An average of about 40 per cent of the organic carbon

present in the water was oxidized by potassium permanganate

in the oxygen consumed procedure.

Literature

Alsterberg, Gustaf. 1929. Ueber das aktuelle und absolute Oz-Defizit der

Seen im Sommer. Botaniska Notiser, 1929 : 354-376.

Birge, E. A. and Juday, C. 1911. Inland lakes of Wisconsin. The dis¬

solved gases and their biological significance. Bui. 22, Wis. Geol. &

Nat. Hist. Survey. 259p.

Birge, E. A. and Juday, C. 1914. A limnological study of the Finger

lakes of New York. Bui. Bur. of Fisheries, 32 : 527-609.

Lonnerblad, Georg. 1931. Ueber den Sauerstoffhaushalt der dystrophen

Seen. Lunds Universitets Arsskrift, N. F. 27 (14) : 1-53.

Maucha, R. 1931. Sauerstoffschichtung und Seetypenlehre. Verh.

Internal. Ver. theoret. und angew. Limnologie, 5 : 75-102.

Naumann, Einar. 1921. Einige Grundlinien der regionalen Limnologie.

Lunds Universtitets Arsskrift, N. F., 17 (8) : 1-22.

Standard methods of water analysis. 1925. Amer. Pub. Health Assoc.

Thienemann, August. 1921. Seetypen. Die Naturwissenschaften, 1921,

Heft 18 : 1-3.

Thienemann, August. 1928. Der Sauerstoff im eutrophen und oligotro-

phen See. Die Binnengewdsser , 4 : 1-175.

468 Wisconsin Academy of Sciences, Arts, and Letters .

Table I. Variation in the quantity of dissolved oxygen in the surface waters of the

various lakes , together with the hydrogen ion concentration and the fixed or bound car¬

bon dioxide . The results for oxygen are indicated in milligrams per liter of water and

in per cent of saturation . The fixed carbon dioxide is indicated in milligrams per liter of

water.

Juday & Birge— Oxygen in Wisconsin Lake Waters. 469

Table I.— -Continued

470 Wisconsin Academy of Sciences , Arts , and Letters .

Table I. — Continued

Juday & Birge— Oxygen in Wisconsin Lake Waters. 471

Table II. Lakes on which series of oxygen determinations were made ; the area of

each lake is given in hectares and the maximum depth in meters. The last column shows

the number of series taken on the various lakes.

GROUP III

472 Wisconsin Academy of Sciences , Arts, and Letters.

Table II. — Continued

GROUP IV

Juday & Birge — Oxygen in Wisconsin Lake Waters. 473

Table III. Dissolved oxygen deficiency or excess at different depths in

14 lakes of northeastern Wisconsin. The column marked 02 shows the

amount actually found at the different depths , 0'2 indicates the amount of

oxygen required to saturate the water at the observed temperatures and D

represents the deficiency or excess of the oxygen at the various depths; the

amount of oxygen is indicated in these three columns in milligrams per

liter of water. The minus sign indicates a deficiency of oxygen and the

plus sign an excess. Tr. means trace .

474 Wisconsin Academy of Sciences , Arts, and Letters ,

Table III. — Continued

Juday & Birge — Oxygen in Wisconsin Lake Waters . 475

Table III.— Continued

476 Wisconsin Academy of Sciences , Arts, and Letters .

Table IV. Amount of dissolved oxygen in 14 lakes of northeastern Wisconsin

The volume of water in the different strata is expressed in cubic meters. O2 represents

the amount of oxygen present, O' 2 the amount required for saturation at the observed

temperatures and D the deficit or excess of oxygen in the various strata ; the results for

these three items are indicated in kilograms of oxygen. The minus sign indicates a

deficit and the plus sign an excess of oxygen in the stratum in question . The results

shown in this table are based on those given in Table III.

Juday & Birge — Oxygen in Wisconsin Lake Waters . 477

Table IV. — Continued

478 Wisconsin Academy of Sciences , Arts , and Letters .

Table V. This table shows the total quantity of oxygen present in the

various lakes and the actual and absolute amounts required for complete

saturation , expressed in kilograms , as well as the percentages of satura¬

tion. The last column shows the relation of the quantity of dissolved

oxygen found in the hypolimnion (H) to that present in the epilim-

nion (E).

Juday & Birge — Oxygen in Wisconsin Lake Waters. 479

Table VI. Temperature , dissolved oxygen and saturation values for

5 lakes of northeastern Wisconsin. 02 is the oxygen present and 0'2 the

quantity of oxygen required for saturation, expressed in milligrams per

liter; g is the actual and g' the theoretical saturation value. The eleva¬

tions of the lakes and the mean barometer readings used in computing the

saturation values are given in Table VII.

480 Wisconsin Academy of Sciences , Arts, and Letters ,

Table VI — Continued

Juday & Birge — Oxygen in Wisconsin Luke Waters . 481

Table VII. This table shows the ratio of the sum of the actual saturation values of

the hypolimnion to the sum of the theoretical saturation values in that stratum , together

with the elevations of the surfaces of the lakes and the mean barometer readings for such

elevations.

482 Wisconsin Academy of Sciences , Arts, and Letters .

Table VIII. Color, oxygen consumed, dissolved oxygen and organic car¬

bon from a number of northeastern lakes which represent the different

types. The color determinations are based on the platinum-cobalt scale and

the readings were made with the standard instrument used by the U. S.

Geological Survey. The oxygen consumed and organic carbon are ex¬

pressed in milligrams per liter of water , and the dissolved oxygen in milli¬

grams per liter and in per cent of saturation.

Juday & Birge — Oxygen in Wisconsin Lake Waters . 483

Table VIII. Continued

484 Wisconsin Academy of Sciences , Arts , and Letters ,

Table VIII. Continued

Juday & Birge — Oxygen in Wisconsin Lake Waters . 485

Table VIII. Continued

486 Wisconsin Academy of Sciences , Arts , and Letters .

Table IX. T/ie color of the surface samples and the minimum , maxi¬

mum and mean quantities of oxygen consumed by the waters belonging to

the different color groups. The color readings are based on the platinum-

cobalt standard and the oxygen consumed is expressed in milligrams per

liter of water .

RACES, ASSOCIATIONS AND STRATIFICATION OF

PELAGIC DAPHNIDS IN SOME LAKES OF WISCONSIN

AND OTHER REGIONS OF THE UNITED STATES

AND CANADA.

Richard Woltereck

Professor of Zoology , University of Leipzig

Notes from the Limnological Laboratory of the Wisconsin

Geological and Natural History Survey. No. L.

Introduction

There are three very different groups of aquatic animals

which show most clearly the recent differentiation of races after

their immigration into the postglacial lakes of Europe and

North America. These are the coregonids among the fishes,

the pulmonates among the mollusks and some cladocerans

(Daphnia and Bosmina) among the pelagic Crustacea. The dif¬

ferentiation of inheritable races of these animals is one of the

last evolutional changes in the animal kingdom except the dif¬

ferentiation of some parasites and of the domesticated animals.

Local differentiates in postglacial lakes can not be much older

than 10,000 to 20,000 years, because these bodies of water did

not become habitable until after the retreat of the ice.

The factors determining the differentiation of these three

kinds of animals have been the same presumably, namely,

changed environment (acting directly or indirectly), isolation

and the internal tendency or potentiality to react by proper al¬

terations to changed external influences.

A comparison of the recent races of coregonids, pulmonates

and daphnids shows some important differences, but principally

a striking resemblance in the general process of differentiation.

The local races and the so called species and subspecies of the

freshwater coregonids are well known; there are hundreds of

them in the United States and Canada and some dozens in Eu¬

rope. We know their geographical origin; they were derived

488 Wisconsin Academy of Sciences , Arts , and Letters .

from marine fishes which migrated into the lakes only tempo¬

rarily at the spawning season at first and later became estab¬

lished permanently in the lakes. Here they have become differ¬

entiated into local and ecological forms, such as inhabitants of

the pelagic zone, of the deep water and of the shore region.

Many of the observed differences are certainly inheritable char¬

acters, while others may be only modifications ; it is a pity that

experimental investigation of these races is almost impossible

since they will not survive in aquaria or small ponds for any

considerable period of time.

The differentiation of some of the freshwater pulmonates is

fairly well known relative to their paleontological history be¬

cause their shells are found in the aquatic sediments of earlier

periods, especially of the Pleistocene. In this group of animals

the recent differentiation and multiplication of species and

races, in comparison with the periods before the existence of

the present postglacial lakes, are especially well known. If

we take as examples the lacustrine pulmonates of Illinois, in¬

vestigated by F. C. Baker, we find 31 recent species and some

older ones in the latest deposits of that region, but only 13

species in the deposits of the early Wisconsin period, and be¬

tween 2 and 9 species in the deposits of earlier Pleistocene in¬

tervals (Peorian, Sangamon, Yarmouth). The few species of

Stagnicola, Fossaria, Helisoma and Physella living at the be¬

ginning and during the earlier part of the Pleistocene have

been split up and differentiated from each other since the re¬

cent lakes came into existence. There are, in addition to the

larger systematic units with more or less prominent differences

in internal structure, a certain number of smaller local differ¬

ences relating to dimensions and shapes; these latter are

described by Baker as variations. He found similar varieties

in artificial lakes also, in habitats where the changes have taken

place during a period of less than a century. Again we do not

know which of these smaller local peculiarities are modifica¬

tions only and which are more than that, but probably Baker

is right in considering some of these young differentiations as

species, or at least races, in the making.

The differentiation of local races in Daphnidae and Bosmini-

dae has peculiar features as compared with those of the core-

gonids and mollusks. There are innumerable local differentiates

W oltereck—Pelagic Daphnids in American Lakes. 489

and we know by experimental cultivation under definite and

varied conditions that these races in most instances are heredi¬

tarily different. Almost all of these innumerable races of Cla-

docera of postglacial lakes belong to only four species : Daphnia

longispina and D. longiremis in Europe, Asia and America,

Daphnia pulex1 in America and Bosmina coregoni in Europe.

Other variable species, usually distinguished but not sharply

distinguishable from Daphnia longispina or Daphnia pulex,

such as D. cucullata and D. retrocurva, indicate only groups or

series of differentiates within these species, which are them¬

selves connected by transitional forms. So there are only a

very few exploding species which have produced all of this im¬

mense richness of endemic differentiates in postglacial lakes.

The Pelagic Daphnids of North America

The various forms of daphnids that are distributed all over

the earth fall into two very different groups2. The first group

contains the above named three species and their subspecies, all

very near to Daphnia pulex and to D. longispina . These two

species are found in all continents, so that they seem to be

rather old; their undifferentiated forms (“primitiva” Burck-

hardt) are of preglacial origin; D. pulex appears as large,

heavy animals living in all kinds of freshwater ponds, bogs and

pools, sometimes together with D. magna, while D. longispina

prefers the open water of the same ponds and of freshwater

lakes. Following the same line of development from littoral

to pelagic organisms, many different daphnids of the plankton

of American and European lakes have been derived from this

D. longispina, such as hyalina, galeata, mendotae and so forth,

and also the whole series of forms called D. cucullata (in Eu¬

rope and Asia) and the northern species D. longiremis and D.

cristata ; the latter is found only in Europe and it is closely re¬

lated to longiremis .

The second group is very different from this one ; it contains

Daphnia magna and some related species, some of them living

as pelagic animals in tropical and subtropical lakes of Asia,

Africa and Australia. These pelagic “M-daphnids”, being mor-

1 This species is abundant in all continents, but it is split into many very dif¬

ferent shapes and races only in North America.

2 See Woltereck 1919. The nomenclature applied in this paper and in other

publications of the author will be treated in a special article next year.

490 Wisconsin Academy of Sciences, Arts, and Letters.

phologically and geographically very different from the other

group (“P-daphnids”), have nothing to do with postglacial

lakes and are not found in North America, northern Europe

or northern Asia3 The continents and their lakes are divided

between the derivatives of Daphnia longispina and D. pulex.

The former has been well developed in all of these continents,

but we find their terminal pelagic races only in northern Eura¬

sia as the cucullata series. Daphnia pulex has no pelagic de¬

velopment in Europe and Asia, but a very strange and mani¬

fold pelagic differentiation in North America. The third spe¬

cies, Daphnia longiremis, is found in the high northern lati¬

tudes of Scandinavia, in Alaska, and in the cold water of the

hypolimnion of some Wisconsin lakes. (See Plate XVIII).

The Daphnia pulex Series

The differentiation of the American pulex derivatives shows

the same peculiar trend as the differentiation of Daphnia longi-

spina-> hyalina ( galeata , etc.)-* cumllata — > longiremis and

as the other series Daphnia magna -» barhata -» lumholtzi -»

cephalata ; that is, in the direction from littoral life to pelagic

life. This tendency in the American Daphnia pulex we see de¬

veloped or developing in various ways and with very different

morphological results. (See Plates XIII and XIV).

1. There are many populations of Daphnia pulex and some of

Daphnia pulex obtusa living in the plankton of large and of

small lakes without any alteration of the body except a certain

transparency. This single physiological and ecological differ¬

entiation of the originally littoral (benthonic) animals occurs,

for instance, in Lake Erie, and in Devils, Nebish, Clear and

many other Wisconsin lakes.

2. There are some differentiates of Daphnia pulex that show

no other visible alteration than a modest crest as a beginning

elongation of the head (Plate XIII, fig. 2). A race like this is

living in Nebish Lake, for instance, together with the usual

short headed form of this water flea.

3. In the western part of the United States, there are popu¬

lations of Daphnia pulex, first described by Forbes from the

Yellowstone Park lakes as D. clathrata and D. arcuata, with

3 A species of the M-group occurs also, as I found recently, on Kauai, Hawai¬

ian Islands.

Woltereck—. Pelagic Daphnids in American Lakes . 491

elongated shape, slightly elongated head and with an alteration

of the postabdomen (claw). I found similar forms in Clear

Lake, California, and in some other lakes in California. The

Clear Lake Daphnia has characters of elathrata and of arcuata

together, so that it is not possible to apply one of these names

to this form; it may be best to distinguish all races related to

these forms as the pulicoides series or subspecies of Daphnia

puiex . The diagnostic characters of these subspecies are : elon¬

gated shape of the shell, in some races also of the head, long

terminal spine which may be straight or curved, large eye and

ocellus, claw with fewer (2-5) and longer teeth in the distal

pecten as compared with the main type of Daphnia puiex .

4. In Wisconsin and other middle western lakes, in the Great

lakes and also in many of the eastern lakes of the United States

and Canada, the most interesting derivatives of Daphnia puiex

are found; they are small animals with elongated shell and in

most cases elongated head (helmet) , with a small eye and with¬

out any eye spot or ocellus (pigment of the primary eye), and

with a pecten that contains more numerous and shorter teeth

than the main type of Daphnia puiex . Daphnids of this kind

have been described as Daphnia retrocurva by Forbes and as

Daphnia breviceps by Birge ; the latter are races without a hel¬

met. In some lakes of Wisconsin, Minnesota, and Canada, there

are many other shapes of the head, such as round helmets not

curved and long, pointed helmets like the European cucullata,

but combined with the claw of Daphnia puiex . All of these

small transparent puiex forms without an eye spot appear to be

members of the same group which we can not call retrocurva

or breviceps . So it may be advisable to apply a comprehensive

name and call all of these races the parapulex series of Daphnia

puiex . By using these two names for the two different series,

we avoid the use of a number of new systematic names for the

races which are on the same level as retrocurva and so forth.

Retrocurva and breviceps or brachycephala remain as impor¬

tant indications of certain shapes which occur in Daphnia puiex

as well as in Daphnia longispina (hyalina) and in Daphnia Ion -

giremis, but they can not be used as specific names.

The most remarkable daphnids of this series are the retro¬

curva forms of some of the smaller lakes, such as lakes Mendo-

ta and Wingra; in the Great lakes the development of the hel¬

met is much inferior (Plate XIV, figs. 6-9) to that developed

492 Wisconsin Academy of Sciences , Arts , and Letters .

by the races of the Wisconsin lakes. The extreme elongations

of these heads are to be compared only with the extreme pel¬

agic helmets of some races of cucullata (derived from longi -

spina) and of cephalata (derived from magna). These races

are indeed the “terminal races” (Grinnell) of the polymorphic

genus Daphnia, each one a terminus in a different series, par¬

allel to the others, but independent of them.

The center of the differentiation of these retrocurva forms

seems to be located in southern Wisconsin, where the pelagic

life may be somewhat older than that of the Great lakes, the

latter presumably becoming inhabitable for pelagic algae and

Crustacea much later than the smaller and shallower lakes.

Another shape of the parapulex daphnids has been developed

in Minnesota, where daphnids with very much elongated but

not distinctly curved helmets (Plate XIV, figs. 12-13) are found

in Bemidji and Vermilion lakes. A similar shape is found in

some very distant lakes in eastern Canada in material which

was kindly shown me by Dr. Klugh of Kingston.

Daphnids of the same group with shorter helmets, not curved

beyond the dorsal side, are rather common in Wisconsin and in

other lakes as well as in some of the Great lakes. Races like

breviceps are found in a large number of the smaller lakes of

Wisconsin, often together with a helmeted race of the same

group, the breviceps being found chiefly in the deeper strata of

these lakes as indicated in the tables.

Besides the regional “centers” of development of certain

shapes in these daphnids, we have to consider the fact that the

whole, or at least the main, differentiation of the parapulex

forms is confined to the region of the northern glaciation in and

around Wisconsin, reaching to about Oneida Lake in the east

and to some parts of Minnesota in the west, but not extending

very far south of this area. On the other hand the differentia¬

tion of the pulicoides forms ( clathrata , etc.) seems to be con¬

fined entirely, or at least mainly, to a western region, out of

the reach of the northern glaciation and outside of the lakes

remaining after the retreat of the last enormous ice sheet

(Plate XIII, figs. 3a-5b).

A close examination of the preglacial distribution of the dif¬

ferent types of American daphnids is very desirable and may

afford a better understanding of these trends of evolution.

W oltereck— Pelagic Daphnids in American Lakes . 493

Since I am now preparing a monograph of the pelagic daphnids

of the world and realize especially the great importance of their

distribution in North America, I should be greatly pleased to

receive plankton samples collected during the summer months

from as many North American lakes as possible4*

The Daphnia longispina Series

The American races of Daphnia longispina are not quite so

interesting as the D . pulex series; the difference between the

American and the European development of this species is not

very large, and in this case in favor of Eurasia where the ex¬

tremes of this development are found. If we exclude the special

differentiation of the D. cucullata series, the number of the

pelagic differentiates of Daphnia longispina hyalina is larger in

North America than in Europe. More important is the fact

that the character of the American shapes of this species is dif¬

ferent from that of the European shapes, although these ani¬

mals are living on both continents in about the same types of

postglacial lakes and under about the same range of conditions.

The predominant forms in Europe are those with pointed heads

(galeata) very often with procurved helmets (procurva) , round

heads of the pellucida type, and the true hyalina form. (Plates

XV and XVI).

In about a hundred American lakes, most of them in Wis¬

consin, we find arched heads like the “lancet arch” of the archi¬

tects, or others like the “rampart arch” ; some have slender and

pointed heads which are retrocurved and with an elongated

rostrum (Plate XVI, fig. 26, “nasuta”) ; there are also strangely

curved heads and helmets like a rococo ornament and low, tent¬

like heads of some races living in the Great lakes and in the

deeper strata of smaller lakes (Plate XVI, figs. 27, 28).

More important than these single differences, but much more

difficult to explain, is the fact that there is a common style of

all of these different American shapes and another general style

of all of the Eurasian types. After examining more than a hun¬

dred American races as well as a large number of European

and some Asiatic types in the past 20 years, I feel able to rec¬

ognize every race of Daphnia longispina as either American or

Eurasian without knowing its locality. One type of D. longi-

4 Address : Biolog. Laboratorium, Seeon be! Obing, Bavaria, Germany.

494 Wisconsin Academy of Sciences, Arts, and Letters.

spina is common to some European and American lakes ; it has

a round head without any helmet and has been called “primi¬

tiva” by Burckhardt. It is found not only in this, but in all

species of the P-series of Daphnia. There is, however, a physio¬

logical and ecological peculiarity of some of the “primitiva”

races of D. longispina in North America. They are very small

dwarf-like races living in the deep water of some lakes that are

not eutrophic, being found in the cold, dark hypolimnion near

the bottom. The same small primitiva type of Daphnia longi¬

spina may live in small shallow lakes and in ponds, but in a

quite different way, namely near the surface as they do in

some European ponds and small artificial lakes, in municipal

parks, for instance. (Plate XV, figs. 15, 16).

In some warm California lakes (Clear, S. Andreas, Upper

and Lower Crystal and others), I found the same primitiva

type of D. longispina without any elongation of the head, and

a form like this is living in the warm Lake Victoria Nyanza in

East Africa and in other tropical lakes. (Plate XVI, fig. 22).

I saw the same form in material collected by Prof. J uday in the

very deep Lake Atitlan in Guatemala, while in another lake of

the same region, the shallow Lake Amatitlan, a race with elon¬

gated head (galeata) was found. (Plate XVI, fig. 23).

Similar daphnids of the same species, without any elongation

of the head, are also common in the cool lakes of the mountains

of North America as in those of our European Alps. It seems

that they have this kind of shape in all lakes, hot or cold, when

they are living and migrating within a thick layer of water,

but they develop elongated, procurved or retrocurved heads

only in lakes where they have to live and migrate in a distinctly

stratified medium and within rather narrow layers of water.

The effect of these elongations and curved rudders seems to be

that the originally steep direction of swimming (jumping) be¬

comes more or less horizontal. The animals living in narrow

layers only 2 m. or 3 m. thick are moving upwards and down¬

wards in directions of small elevation ; if they have to migrate

in about 12 hours from a depth of 50 m. to the surface, the ele¬

vation of their movement has to be a much steeper one.

The Daphnia longiremis Series

These animals, easily distinguished by their elongated swim¬

ming antennae, have been found in northern Norway and Swed-

Woltereck — Pelagic Daphnids in American Lakes . 495

en by Sars and Lilljeborg. I found them in plankton material

from Karluk Lake, Alaska, and Dr. Birge found the same form

in Wisconsin lakes many years ago. I saw these strange little

daphnids in the plankton of 8 Wisconsin lakes out of about 40

examined and in one out of 6 Indiana lakes. In all cases they

were found only in the cool water below the thermocline. They

live near the bottom if there is enough oxygen, and near the

thermocline if there is a lack of oxygen in the deeper strata.

If all of the hypolimnion is without oxygen for some time dur¬

ing the year, such lakes are not inhabitable for cold water or¬

ganisms like Daphnia longiremis . (Plate XVII).

In lakes where the water is well supplied with oxygen from

thermocline to bottom these races have no elongation of the

head, while other local races living for some months in a nar¬

row layer between the thermocline and the oxygen-free depth,

develop prolongations of the head, either straight or retro-

curved helmets (Plate XVII, figs. 33-85). This indicates again

that not the high temperature of the water but its stratification

and the necessary change in the direction of swimming are the

factors determining the peculiar prolongations and the curved

“rudders” attached to the head of so many pelagic daphnids

and bosminids. (See Woltereck 1928).

Associations of Pelagic Daphnids in Some American Lakes

There is a very conspicuous difference between the lakes of

the old and the new world, even if we compare only lakes of

about the same size, the same history (as glacial relicts) and

the same conditions of temperature, light, oxygen, pH and so

forth. In European lakes of this kind, we find only one or two

differentiates of pelagic daphnids ; if there are two, they belong

almost always to different series, such as longispina and cucul -

lata. Most of the Wisconsin lakes that I have been able to ex¬

amine closely enough contain more than two differentiates ; th^

average of 22 lakes from which I have examined living material

of all strata, is three to four different races for one lake. All

of these lakes have a maximum depth of only 20 m. to 30 m.

except Trout Lake which is 35 m. deep.

One reason for the greater number of races in these small

American lakes and for only one or two races in similar and

also in much larger and deeper European lakes, is the fact that

496 Wisconsin Academy of Sciences , Arts, and Letters .

there are only two pelagic species or series (longispina and

cucullata) in Europe, while there are three pelagic species in

America, namely longispina, pulex and longiremis . But this

explanation is not sufficient; there is another and more inter¬

esting difference. If we have two different races in a single

European lake, one is invariably a longispina race and the other

a cucullata race. Many American lakes of the same postglacial

character, on the other hand, contain two races of longispina or

two races of pulex (parapulex). Crawling Stone Lake, for

instance, which has a maximum depth of 28 m., contains one

pulex, two different parapulex races, one longispina and one

longiremis race. It is not possible at the present time to ex¬

plain this kind of polymorphism in American lakes of the usual

postglacial type.

I have examined plankton material from a large number of

other lakes in Wisconsin, Indiana, Michigan, Minnesota, New

York, Ontario (Canada) and from the Great lakes, but not

complete series from all depths, only occasional samples; from

this material I have obtained the impression that the usual

association of pelagic daphnids in the eastern part of the United

States is one smaller race of parapulex (mostly retrocurva) ,

and one larger race of longispina (mostly mendotae or galeata) .

This is similar, ecologically, to the common association of one

longispina with one smaller cucullata in Europe. Many Ameri¬

can lakes contain, in addition to this “standard association”, one

of the common pulex races and in some instances a race of D.

longiremis is found. The Great lakes, which have not been very

thoroughly investigated so far, seem to contain a number of

races with different and characteristic forms in the different

parts of each lake, such as found in Lake Erie for example. In

Ontario, Oneida, Nipigon and Simcoe lakes, I found the same

association of parapulex and longispina (galeata or mendotae ).

In Clear Lake (California) I obtained two races of longispina

(one large and helmeted, the other very small and short — primi-

tiva), and one D. pulex (pulicoides). In San Andreas and the

two Crystal lakes near San Francisco, I saw one large and hel¬

meted D. longispina, a small primitiva form of the same species

— both very near to the Clear Lake populations — , one large

D. pulex (obtusa), and another pulex of the pulicoides type. In

many of the lakes in the Sierra and Rocky mountains, simpler

associations of Daphnia longispina and D. pulex are found.

Woltereck — Pelagic Daphnids in American Lakes. 497

Some of the Wisconsin associations are represented in Tables

I to XII. Future work may account for some of these different

types of communities, but we do not understand most of these

differences at present, in spite of the fact that the physical

and chemical conditions of the Wisconsin lakes are so well

known. One example will show that it is still too early to dis¬

cuss the relation between environmental conditions and these

associations. I wished to understand why Daphnia longiremis

is present in some lakes and not in others when the conditions

of temperature and oxygen are about the same. I was glad to

see that the first 7 lakes where I found this small species in the

hypolimnion, all contained hard water, with between 10 mg. and

25 mg. of fixed or bound carbon dioxide per liter ; but the next

lake in which I found a colony of this same species was Big

Carr, which has very soft water, or an average of less than 2

mg. of fixed carbon dioxide per liter. The shape of the animals

of this race is very different from that of all of the other races ;

Big Carr Lake contains a race similar to Lilljeborg’s D . longi¬

remis from Storsjon in Jemtlandia (Table XX, Plate XIV, fig.

13). This lake is situated in the granite region of northern

Sweden and presumably contains very soft water. But the

presence of this species in some lakes and its absence in others

is not understood at present.

Stratification of Pelagic Daphnid Populations in Some

Wisconsin Lakes

I had a good opportunity to examine the vertical distribution

of the various daphnids in 16 very different lakes in northeast¬

ern Wisconsin with a most effective instrument for such an in¬

vestigation, namely a plankton trap. Most of the catches were

obtained by Prof. Juday personally or by one of his trained

assistants; the night catches were secured by Mr. Baum and

myself. Each catch represents the pelagic animals of 10 liters

of water, and all of the daphnids present in a sample have been

counted and not estimated. These observations were made dur¬

ing my visit at the Trout Lake Limnological Laboratory of the

Wisconsin Survey.

The results of these catches and countings are represented

in Tables I to XII, which show the vertical distribution of the

different populations of daphnids in the several Wisconsin

498 Wisconsin Academy of Sciences , Arts , and Letters .

lakes. In the different series, the catches were taken at inter¬

vals of 2 m. to 5 m. ; the numbers of young, adult (egg bear¬

ing), and the total number of daphnids in each 10 liter catch,

are given in the tables. There were almost no males during

the time of this investigation, which extended from August 20

to September 9, 1931. The maxima of each race in every ver¬

tical series are indicated in the tables in italics ; the position of

the thermocline is shown by a broken line.

The stratification of the different daphnids living in the same

body of water seems to be sufficiently expressed by the location

of the maxima ; the total “zone of habitation” of every race can

be estimated if we neglect the very small numbers (less than

10 per cent of the maximum) which are found in the lower

strata and which possibly represent dying or moulting speci¬

mens that sink below their normal habitat.

The results shown in the tables may be distinguished as fol¬

lows:

1. They confirm plainly the earlier statements made by Juday

(1904) that the helmeted daphnids in Wisconsin lakes live

above the short headed ones, and similar observations made la¬

ter by Dr. Ter-Poghossian (1928) and others in lakes near See-

on in Bavaria. A report on the “zonare Verteilung der helrn-

losen und der helmtragenden Biotypen von Daphnia” has been

published by Woltereck (1930). The new investigation of the

vertical distribution in some Wisconsin lakes has produced an

unexpected variety of other results. Some of these results will

be published after the examination of the stratification of the

nannoplankton algae is completed, but some others may be dis¬

cussed in this connection.

2. There are great differences in the thickness of these racial

“zones of habitation” in the different lakes; the zones range

from a thickness of about 15 m. or 20 m. for Daphnia longi -

spina in Fence and Trout lakes to only 1 m. or 2 m. for Daph¬

nia longiremis in Muskellunge Lake.

3. The latter case is especially interesting because this is a

population of cold water animals that usually lives near the

bottom, but it is compelled to live temporarily in a very thin

stratum of water just below the thermocline in Muskellunge

and some other lakes owing to a lack of oxygen in the deeper

strata. The main point is that these populations of D, longi-

Walter eck— Pelagic Daphnids in American Lakes . 499

remis develop elongations of the head, namely straight or retro-

curved helmets, while the other races of the same species liv¬

ing in lakes that possess enough oxygen to enable them to oc¬

cupy the whole hypolimnion, are of the same short headed shape

as the races living in lakes of Alaska and northern Scandina¬

via.

4. We found populations of pelagic daphnids in the hypolim¬

nion which belong to three different types, but which are very

similar in shape. After Juday, Kikuchi (1930) was the first

investigator who described special populations of Cladocera and

other plankton organisms that were confined to the hypolimnion

of some Japanese lakes during the summer months; he found

round headed populations of Daphnia longispina in this situa¬

tion.

In Day Lake, Wisconsin, there is a population of small

D. longispina (primitiva) with round heads living between the

thermocline and the bottom, that is, between 9 m. and 15 m.

In Dead Pike Lake, there is a population of large D. longispina

(apicata) living between 10 m. or 14 m. and 21 m. Also there

are many populations of Daphnia longiremis, all living below

the thermocline ; they are always found near the bottom if there

is enough oxygen in the deepest layers, as in Trout Lake, Wis¬

consin and in James Lake, Indiana, for instance. In these lakes

they seem to be concentrated especially in the deepest holes.

In Trout Lake we found 300 individuals in a 10 liter catch taken

at 31 m. (1 m. above the bottom) in the afternoon, while in the

late evening of the same day when the buoy marking the 32 m.

water could not be found, we obtained only 36 individuals in a

10 liter catch taken at 30 m.

5. Some series of catches taken about 9 p. m. in Trout Lake

and in Day Lake show the beginnings of an upward migration

of some but not of all populations in these lakes. The maxima

of Daphnia longispina (galeata) and of parapulex (retrocurva)

in Trout Lake are nearer the surface at this time; the first

moves upward from 5 m. to 3 m. and the latter from 12 m. to

5 m., while the position of the deep water population of Daph¬

nia longiremis is unchanged. (See Table IX).

In Day Lake the maximum of D. longispina (nasuta) is trans¬

ferred from 7 m. at 4 p. m. to 5 m. at 9 p. m. and 17 specimens

were found in the 10 liter catch made at the surface in the

500 Wisconsin Academy of Sciences , Arts , and Letters .

evening where none was found at this depth in the afternoon.

The maximum of D. longispina mendotae moved upward some¬

what in the evening, that is, from 10 m. at 4 p. m. to between

7 m. and 9 m. at 9 p. m. In this lake also the maximum of the

primitiva form of D. longispina moved from 14 m. at 4 p. m.

to 11 m. at 9 p. m. ; a very large maximum (swarm ? ) was

found at the latter depth in the evening. No specimens were

present in the 7 m. catch, which was in the upper part of the

thermocline, or in the catches taken nearer the surface. (See

Table X.)

6. Two other results, or questions for further investigation

rather than definite results, can only be indicated at present

because the number of ascertained facts concerning them is now

too limited to permit an explanation of these phenomena.

In some lakes, such as Trout, Fence and Big Carr, we find

Daphnia longispina galeata occupying a thick stratum of wa¬

ter; in Trout Lake and Fence Lake they extend from the sur¬

face to a depth of 20 m. and in Big Carr Lake from the sur¬

face to 12 m. All of these populations are living both above

and below the thermocline, and this is true also of the same

species in many European lakes.

In Trout Lake there are two distinct maxima of this race,

but I could not distinguish any morphological difference be¬

tween those living in the epilimnion and those found in the

hypolimnion. There may be a physiological difference, and

surely there must be an ecological difference between the ani¬

mals living in the warm upper water and those found in the

cool water below the thermocline. In Day Lake there are two

different races of the same helmeted Daphnia longispina ; one

is a slender form (nasuta) living above the thermocline and the

other is a heavy form (mendotae) which lives in the thermo¬

cline, with a maximum at 10 m. at 4 p. m. and between 7 m.

and 9 m. at 9 p. m.

The questions that we cannot answer at present regarding

these forms are as follows : Have these two races been differ¬

entiated within this same lake, at first ecologically, then physio¬

logically and at last morphologically? Or have they been intro¬

duced into these lakes from elsewhere as already differentiated

races ? Further investigation, both ecological and experimental,

are necessary before these questions can be answered.

WolterecJc- — Pelagic Daphnids in American Lakes . 501

7. The same applies to another observation and question.

Different species which live in the same lake under identical

conditions, near the bottom or near the thermocline for in¬

stance, show the same or a very similar shape of the head in

many cases. We have experimental evidence in European forms

that such characteristic shapes are not only modifications but

inheritable peculiarities of the different races.

I wish only to mention this fact and to indicate as a strik¬

ing example the two races Daphnia longiremis and D . longi-

spina which live at the same level in Dead Pike Lake and which

have the same short, tent-like head. (See Plate XVII, fig. 31).

Other examples are the local races of Daphnia pulex (parapu-

lex) and of D. longispina (hyalina) in Silver, Muskellunge and

some other lakes. These inheritable local characters induced

by certain conditions as well as the complexities concerning the

general “style” of such features and shapes, and their regional

distribution in America, constitute the most important subjects

of future investigations in this field of limnology. Today we

are only at the beginning of this kind of ecological, and at the

same time genetical, research.

Some General Results

Taxonomic. All American (and European) races of pelagic

daphnids have been derived from two main species that are

very nearly related, namely Daphnia pulex and Daphnia longi¬

spina. Each of these species has been split into numerous races

which form a few very distinct series or subspecies. Daphnia

pulex occurs in America mainly in four series, as follows:

Daphnia pulex pulex; Daphnia pulex obtusa ; Daphnia pulex

parapulex (including retrocurva and many others) ; Daphnia

pulex pulicoides (including clathrata and others).

Biogeographic, a. The third American species, namely Daph¬

nia longiremis, which is closely related to D. longispina, is an

arctic form. It is found in northern Scandinavia and in Alas¬

ka, and it has developed some local races which are now living

in the cold deep strata of some Wisconsin and Indiana lakes.

It is very probable that these forms occur also in Canadian

lakes.

b. The pelagic series of Daphnia pulex (parapulex and puli¬

coides) have been developed only in North America; the dif-

502 Wisconsin Academy of Sciences , Arts , and Letters .

ferentiation area of parapulex is located in the postglacial lakes

of the northern and eastern part of the continent, while the

differentiation center of the pulicoides series seems to be located

in the western part of the United States.

c. The geographical distribution of five of the main groups

of daphnids is shown in Plate XVIII.

E cologic, a . There are some definite and different associa¬

tions of pelagic races of Daphnia in American lakes ; from three

to five races are found in the associations noted in Wisconsin

lakes, but only one or two occur in the associations observed in

European lakes.

6. Every lake that has stratified water in summer, contains

during that time a clear stratification of the different popula¬

tions or races of Daphnia. Some associations belong to the

epilimnion, others to the thermocline and still others to the

hypolimnion.

c. The differences between the populations living in the dif¬

ferent strata of the same lake may be only ecological and

physiological in character, or they may be morphological char¬

acters of ecological value in relation to the necessary direction

of swimming as determined by the shape of certain appendages

of the head.

Genetic . a . Three orders of inheritable characters may be

distinguished : Local peculiarities of the races ; regional quali¬

ties of groups of local types; continental “style” of all Ameri¬

can and of all European races of Daphnia longispina for in¬

stance.

b. Racial characters in pelagic daphnids show a close corre¬

lation with the special conditions of life in most cases (see

above). This correlation proves that alteration of racial quali¬

ties in daphnids is either caused or directed by certain environ¬

mental factors.

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Woltereck, R. 1930. Alte und neue Beobachtungen iiber die geo-

graphische und die zonare Verteilung der helmlosen und helmtragen-

den Biotypen von Daphnia. Internat. Rev. ges. Hydrobiol. u. Hydrog.

24 : 358-380.

Table I. Vertical distribution of Daphnia in Nebish Lake on August 29, 1931.

The figures in italics represent the maxima.

504 Wisconsin Academy of Sciences , Arts, and Letters .

Table II. Vertical distribution of Daphnia in Little Long Lake on August 25, 1931

The figures in italics represent the maxima .

Table III. Vertical distribution of Daphnia in Weber Lake on August 25, 1931.

The figures in italics represent the maxima.

Table IV. Vertical distribution of Daphnia in Silver Lake on August 28, 1931.

The figures in italics represent the maxima.

Wolterech — Pelagic Daphnids in American Lakes . 505

Table V. Vertical distribution of Daphnia in Dead Pike Lake on September 2

1931. The figures in italics represent maxima.

Table VI. Vertical distribution of Daphnia inBig Carr Lake on September 5 , 1931.

The figures in italics represent maxima.

Table VII. Vertical distribution of Daphnia in Presque Isle Lake on August 31 ,

1931. Cloudy. Figures in italics represent maxima.

506 Wisconsin Academy of Sciences, Arts, and Letters,

Table VIII. Vertical distribution of Daphnia in MuskellungeLake on August 26,

1981. The dissolved oxygen amounted to 0.21 mg. per liter at 12m.) none was found

at 15m. and below that depth. Figures in italics represent maxima.

Table IX. Vertical distribution of Daphnia in Trout Lake on August

21, 1931, at 2:30 and 9:00 p. m. respectively . Figures in italics represent

maxima.

Distribution at 2:30 p. m.

Table X. Vertical distribution of Daphnia in Day Lake on August 20, 1981 at J+\00 p, m. and 9:00 p. m. respectively. The figures in

italics represent the maxima.

Walter eck — Pelagic Daphnids in American Lakes . 507

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Vertical distribution of Daphnia in Fence Lake on September 1, 1931. The figures in italics represent the maximal

508 Wisconsin Academy of Sciences , Arts , and Letters .

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510 Wisconsin Academy of Sciences, Arts , and Letters.

Plate XIII

FIGS. 1-5. Examples of racial series (“subspecies”) of Daphnia pulex

with ocellus and without helmet. Summer forms only; young females, x 50.

Camera drawings.

Fig. la. Daphnia pulex pulex. Usual form without helmet. Nebish Lake,

Wisconsin.

Fig. lb. Daphnia pulex pulex. Usual form of the claw.

Fig. 2 Daphnia pulex pulex (cristata). With short crest. Nebish Lake,

Wisconsin.

Fig. 3a. Daphnia pulex obtusa. A small pellucid race. Lake San

Andreas, California.

Figs. 3b, 3c. Daphnia pulex obtusa. Two views of claw.

Fig. 4. Daphnia pulex pulicoides. Elongated form, large eye. Lake San

Andreas, California.

Fig. 5a. Daphnia pulex pulicoides (cristata). With short crest (from a

fall catch, summer form possibly a little more elongated). Clear Lake,

California.

Fig. 5b. Daphnia pulex pulicoides (cristata). The claw of this race is

near D. clathrata Forbes, while the head is near D. arcuata Forbes.

TRANS. WIS. ACAD., VOL. 27

PLATE XIII

3 o .

512 Wisconsin Academy of Sciences , Arts, and Letters .

Plate XIV

Figs. 6-14. Examples of racial series (“subspecies”) of Daphnia pulex

without ocellus and with helmet, which has been secondarily reduced in D.

breviceps Birge (figs. 10, 11). Summer forms only; young females with

first brood of eggs, x 35. Camera drawings.

FIG. 6. Daphnia pulex parapulex (retrocurva Forbes). Specimen from

the original material of Dr. Forbes used for describing this race as

“ Daphnia retrocurva”. Lake Mendota, Wisconsin.

Figs. 7-9. Three local races of the same subspecies selected out of a

large number of differentiates: Fig. 7. Lake Erie. Fig. 8. Presque Isle

Lake. Fig. 9. Kawaguesaga Lake, Wisconsin.

Figs. 10-11. Daphnia pulex parapulex (breviceps Birge). From Weber

and Fence lakes, Wisconsin.

Figs. 12-13. Daphnia pulex parapulex (elongata). From Vermilion and

Bemidji lakes, Minnesota. The shape of these races of D. pulex parapulex

is very similar to that of some races of D. longispina cucullata in Europe.

Fig. 14. Typical claw of Daphnia pulex parapulex.

TRANS. WIS ACAD., VOL. 27

PLATE XVI

514 Wisconsin Academy of Sciences , Arts , and Letters ,

Plate XV

Figs. 15-21. Examples of racial series (“subspecies”) of Daphnia long-

ispina without or with a rounded helmet and with ocellus. Summer forms

only : young females with the first brood of eggs, x 50. Camera drawings.

Figs. 15-16. Dap hnia longispina longispina (primitiva Burckhardt).

From Fair View Pond, Decatur and from Clear Lake, California. From

fall catches, but presumably without alteration in summer. Similar races

in Day Lake, Wisconsin and other races have the same shape in August-

September.

FIGS. 17-19. Daphnia longispina elongata. Three different dolichocephata

types of races. Fig. 17. Forma mendotae Birge, very common in many

races of the middle west and eastern parts of the United States; this

specimen from Day Lake, Wisconsin. Fig. 18. Forma indianae; found in

some lakes of northern Indiana. This specimen from Tippecanoe Lake,

Indiana. Fig. 19. Forma not yet named; found in some western lakes.

This specimen from Clear Lake, California.

Fig. 20. Daphnia longispina elongata. With only a short helmet oi

crest in summer ( mesocephala type). From Weber Lake, Wisconsin.

Fig. 21. Typical claw of all races of D. longispina.

TRANS. WIS. ACAD., VOL. 27

516 Wisconsin Academy of Sciences , Arts, and Letters .

Plate XVI

Figs. 22-28. Examples of racial series (“subspecies”) of Daphnia longi-

spina apicata. With pointed crests or helmets and with ocellus. Claw as

in fig. 21. Summer forms only; young females with the first brood of

eggs, x 50. Camera drawings.

Fig. 22. Daphnia longispina. Without crest or helmet. From the tropi¬

cal Lake Atitlan, Guatemala. Forma primitiva as in figs. 15-16 and as in

Day Lake, Wisconsin, Victoria Nyanza Lake, East Africa, and in many

other lakes and ponds.

Fig. 23. Daphnia longispina apicata. From the tropical Lake Amatitlan,

Guatemala.

Figs. 24-26. Examples of dolichocephala races of D. longispina apicata.

Fig. 24. Typical forma galeata from Crawling Stone Lake, Wisconsin.

Fig. 25. Forma curvata from Lake Erie. Fig. 26. Forma nasuta from Day

Lake, Wisconsin.

Figs. 27-28. Examples of mesocephala and tent-like types of D. long¬

ispina apicata. Fig. 27. From Dead Pike Lake, Wisconsin. Fig. 28. From

Lake Michigan.

TRANS. WIS. ACAD., VOL. 27

PLATE XVI

518

Wisconsin Academy of Sciences , Arts , and Letters .

Plate XVII

Figs. 29-35. Examples of racial differences (not yet named) of Daphnia

longiremis. Summer forms only; young females with the first brood of

eggs, x 50. Camera drawings.

Fig. 29. Primitiva type from Karluk Lake, Alaska.

Fig. 30. Race with similar head and very long spine from Silver Lake,

Wisconsin. The elongated antennae and the peculiar curve of the shell are

the same in all races of this species.

Fig. 31. Race with tent-like head from Dead Pike Lake, Wisconsin.

Fig. 32. Larger race with elongated rostrum (nasuta) from Trout Lake,

Wisconsin.

Fig. 33. Similar race with slightly elongated head from Presque Isle

Lake, Wisconsin.

Fig. 34. Race with elongated and curved helmet (retrocurva) from

Muskellunge Lake, Wisconsin.

Fig. 35. Race with elongated and rounded helmet from Big Carr Lake,

Wisconsin.

TRANS. WIS. ACAD., VOL. 27

PLATE XVII

520 Wisconsin Academy of Sciences , Arts, and Letters .

Plate XVIII

The areas indicated on the map show the

present geographical distribution of the vari¬

ous types of Daphnia. Areas A and B are oc¬

cupied by derivatives of Daphnia pulex ; A

representing the pulicoides series, and B the

parapulex series. Areas C and D are occupied

by derivatives of D. longispina; C representing

the cucullata series, and D the longiremis se¬

ries. The distribution of the derivatives of D.

magna is shown by area E.

TRANS. WIS. ACAD., VOL. 27

PLATE XVIII

SOLAR RADIATION AND INLAND LAKES. FOURTH

REPORT. OBSERVATIONS

OF 1931

E. A. Birge and C. Juday

Notes from the Limnological Laboratory of the Wisconsin Geological

and Natural History Survey. No. LI.

This paper reports the work of the summer of 1931 on the

transmission and absorption of solar radiation by the waters

of the lakes of the Highland Lake District. The percentile rela¬

tion is determined between the amount of radiation delivered

to a unit of area at the surface of the lake and that delivered

to a similar unit at different depths of the lake. From these

data are platted transmission curves of radiation, both those

observed and those computed for zenith sun. The main task of

1931 was to extend observation to greater depths of water and

to smaller quantities of radiation, by employing a sensitive

galvanometer in addition to the millivoltmeters already in use.

Much work was done on the form of the solar energy spectrum

and on the changes which it suffers in the lakes. The results of

this part of the study were of a type similar to those reported

in our third paper (Birge and Juday ’SI1) ; they were carried

to greater depths, corresponding to the increased sensitivity of

the receiving instrument. Many observations were also made

on the transmission of total radiation in lakes of various types,

and these are the subject of the present report. They enable

us to give a general notion of the transmission of solar radia¬

tion through the waters of small inland lakes during the period

of summer stratification. This can be done for lakes of widely

different types, which present a more complicated story than

does the ocean or even large and deep fresh-water lakes.

The observations reported in earlier papers, made with ther¬

mopile and millivoltmeters, were carried down to the value of

1.0 per cent (’29) or 0.10 per cent (’30) of the radiation

delivered to the surface of the lake. Under similar conditions

of observation we can now follow transmission until the limit

1 For the sake of brevity our earlier papers on this subject will be cited as

’29, *30, ’31, without using the names of the authors.

524 Wisconsin Academy of Sciences, Arts, and Letters.

of 0.01 per cent has been reached or even passed; the extreme

values observed being less than one-tenth of those reported in

earlier papers. The curves platted on semi-logarithmic paper

extended through two logarithmic cycles in the first report ; in

the second paper three cycles were shown and in the present

paper the curves extend through four cycles and even into the

fifth. The result is that in many of our lakes observation has

been extended into the region where the increased opacity of

the hypolimnion greatly modifies the rate of transmission. In

the diagrams of the first report (’29) very few observations

went beyond the straight-line curves that mark the character¬

istic transmission of epilimnion and the water immediately

below it. In this respect they resemble most of the curves

given for much greater depths of the ocean by Shelford and

Gail (’22) and Poole and Atkins ('28, ’29) ; and those for the

Bodensee by Oberdorfer (’28) . The second report (’30) car¬

ried the curves nearly to the bottom of six lakes whose water

is so transparent that more than 0.10 per cent of incident radi¬

ation is left at such depths. In the present report similar curves

are extended to 0.01 per cent with corresponding fullness of

knowledge regarding lakes of a much less transparent type.

The work of 1930 was devoted mainly to a more careful

study of the effect of the lake waters on the form and com¬

position of the solar energy spectrum and observation rarely

went below the epilimnion (’31 : 391) and thus was limited to

the region of characteristic transmission. In 1931 observation

extended into the hypolimnion of all lakes whether transparent

or relatively opaque. The most important effects of hypolimnion

on radiation could therefore be worked out in lakes of widely

different types. There is still incomparably more to be known

about the transmission of solar radiation in these lakes than

has already been ascertained, but the general outlines of the

story have been traced. Just as our earlier reports showed the

characteristic transmission of radiation in a wide variety of

lakes, so the present report carries the story on to include the

general effect of the hypolimnion, with its increase of color

and turbidity, on the transmission of the sun's radiation.

Our studies have been made on small lakes and after sum¬

mer stratification has been established. All observations are

made in full sunshine unless expressly stated otherwise. In

all tables radiation is stated as a percentage of that delivered

Birge & Juday — Solar Radiation and Inland Lakes . 525

to the surface. In computing results all radiation is assumed to

be direct from sun ; none is regarded as diffuse. Correction is

made for difference of reading of thermopile in air and in

water as stated in earlier reports (’29 : 515). Observed values

are recomputed and stated for zenith sun, in the same way as

in former reports. By this process all observations are reduced

to a common denominator. All of these corrections and adjust¬

ments are approximate rather than minutely accurate.

In the report of 1929 the value of the surface reading was

given in cal/cm2/min. In later papers this has been omitted as

having no special significance. If the amount of radiation pres¬

ent at any depth is stated as a percentage of that delivered to

the surface, it is easy to compute its value in units of any kind.

There is no difficulty in using meter-candles rather than ca¬

lories, so far as the surface illumination is concerned. But for

observations in the depths of lakes in whose water the form of

the solar energy spectrum is modified profoundly and rapidly,

with consequent change in the color of the light, the term

“candle” becomes little more than a conventional expression for

energy ; it does not mean “illumination” of a type discernible by

the eye.

In all of our papers the term “transmission” refers to the

passage of solar radiation through the waters of a lake. The

unit of distance is one meter ; rate of transmission is expressed

by the percentile ratio between radiation at the top and the

bottom of a one-meter stratum.

Results — General Statement

1. By the use of pyrlimnometer and galvanometer the pass¬

age of solar radiation through the water of lakes may be fol¬

lowed until it is reduced to about 0.01 per cent of that incident

on the surface. This limit is approximately the same as that

reported in observations made with other instruments, such as

the photoelectric cell or photography. The pyrlimnometer,

whose central component is a thermopile, registers radiation of

all wave-lengths; other instruments are more selective and are

sensitive to the radiation in the short-wave part of the visible

spectrum.

2. Most of the infra-red radiation and much of the ultra¬

violet are absorbed in the first meter of lake water. For prac-

526 Wisconsin Academy of Sciences , Arts, and Letters .

tical purposes the remainders of such radiation present at one

meter below the surface may be neglected, and all radiation at

that depth and below may be considered as light.

3. If transmission of radiation is measured by instruments

which are sensitive to short-wave light only, the transmission

of the upper meter of the water will be substantially the same

as in the meter below. If the instrument gives full value to

radiation of wave-length 6000 A and more, transmission of light

in the upper meter will be smaller than below on account of

the rapid rise of absorption by water of light in the longer

wave-lengths. If the pyrlimnometer is used transmission in the

upper meter should be stated by itself as it will differ consider¬

ably from that below. In fairly transparent lakes transmission

below one meter will be much the same, whatever the instru¬

ment used.

4. Transmission of radiation through the water of lakes de¬

pends on (1) the effect of water as such; (2) the color of the

water due to stains; (3) the turbidity of the water. Color dif¬

fers greatly in different lakes, but is a fairly constant quality

in any one lake. It renders the water highly selective toward

radiation, cutting off the short-wave radiation. Turbidity de¬

pends on particles suspended in the water and is very variable

both in different lakes and in the same lake at different times

and depths. If particles are large their action on radiation is

nearly or quite non-selective ; if small, the shorter wave-lengths

are more rapidly extinguished. In any ordinary lake all sizes

of particles are likely to be present. Transmission as actually

found is therefore a result of several variable factors, whose

separate influence has not been adequately studied.

5. In general the water of the ocean and of large lakes is free

from stain or has only a slight color. Differences in transmis¬

sion of radiation are primarily due to turbidity and in a large

body of water this is likely to be fairly uniform to depths

reached by observation. A series of observations is likely to

disclose a characteristic transmission , which extends without

notable change to the limit of observation. This limit may be

found in the ocean at 60-75 m. (Poole and Atkins *28, *29) or

even 120 m. ( Shelf ord and Gail *22) ; the same minimum of

radiation may be reached at 20-30 m. in the Bodensee (Ober-

dorfer) . These series both in fresh and salt water, yield sub¬

stantially straight line curves of average transmission when

Birge & Juday - — Solar Radiation and Inland Lakes. 527

platted on semi-logarithmic paper. Differences in the turbidity

of different strata may cause sudden and great changes in these

curves (Poole and Atkins, '28 : 276), but such changes are

not frequently observed.

Results — Small Lakes

1. These general statements must be modified in order to rep¬

resent the situation in small lakes like those of the Highland

Lake District. The color of the water in different lakes is very

different; turbidity may be very slight or present in a high de¬

gree; both color and turbidity may be fairly uniform through

the whole water of the lake or may change greatly with depth.

The effect of currents induced by wind may be almost absent

or may be very great; turbidity due to wave action on the

shores is variable but is usually small and may be almost en¬

tirely absent. The result of these numerous factors is to com¬

plicate the story of transmission of radiation in such lakes.

2. Small lakes have a characteristic transmission, found from

1 m. on for a variable distance. Its value is dependent on color

and turbidity, and if these are uniform it may continue with¬

out marked change to the bottom of the lake or to the limit of

observation. It usually continues through the epilimnion and

for some distance into the hypolimnion. No observable differ¬

ences in transmission are traceable to changes of temperature

and density of water at the thermocline.

3. Transmission in the ocean and in large lakes, so far as

these have been examined, is ordinarily high, rarely below 70.

In small lakes it ranges from 85 or more, as a maximum, to

10-12 or even less. Its value in any lake is primarily deter¬

mined by the color of the water, which in the lakes reported

in this paper, ranges from 0 to 260 on the platinum-cobalt scale.

Color is a fairly constant quality of the water of these lakes,

varying relatively little from year to year. It is ordinarily

higher near the bottom of the lake; but usually such increase

has much less effect on transmission of radiation than is ex¬

erted by the increased turbidity at the same depths. Color in

the upper water of the lakes is of organic origin ; in the deeper

water, where oxygen is absent, it may be due to iron. The re¬

lation between color and transmission, so far as this is known,

is shown in Table IV and Fig. 7.

528 Wisconsin Academy of Sciences , Arts , and Letters .

4. Turbidity depends on particles, either organic or inorganic,

suspended in the water. In the Highland Lake District there

is little inorganic turbidity, since the soil is sandy, there is little

cultivation, and the District is densely covered with second

growth trees. Organic particles consist of plankton and its im¬

mediate derivatives, particles from the bottom ooze, and debris

from marsh and peat. These particles exert a mechanical ob¬

struction to the passage of radiation. Particles of all sizes are

usually present in these lakes; but great growths of plankton

algae are rare. Nannoplankton constitutes the bulk of the liv¬

ing organic material. There is no quantitative measure for

turbidity, as there is for color, and therefore statements re¬

garding its effect on radiation are less definite. Turbidity is the

prime factor which determines the range of transmission in

lakes belonging to the same color group (Table IV), and to

variations in turbidity are due most of the changes in transmis¬

sion as radiation passes downward through the water of lakes.

5. The hypolimnion is ordinarily more turbid than the epi-

limnion; turbidity usually increases with depth and is at a

maximum in any lake just above the bottom of the deepest wa¬

ter. This condition causes most of the changes in transmission

shown in the curves of Figs. 1, 2, and 3.

6. In lakes with very high colored water, radiation is rapidly

reduced to that region of the spectrum to which the water is

most transparent. This may aid in causing a marked increase

of transmission immediately below the depth of, say, 2 in.

Transmission in the 2-3 m. stratum may be twice as great as

between 1 m. and 2 m. This situation is seen in different de¬

grees in the last five lakes of Table I. In such lakes there is

no characteristic transmission, within the limits to which radia¬

tion can be followed by the instruments. This situation has

not been found by us in lakes with color less than 60 ; but the

lakes examined are not numerous enough to permit general

statements as to this limit. This change of transmission is not

present, in observable degree, in lakes with colors from 20 to

36, as shown in Table I and Figs. 1 and 2.

Instruments

The instrument for receiving the sun’s radiation during the

season of 1931 was the pyrlimnometer, as described and figured

Birge & Juday — Solar Radiation and Inland Lakes . 529

in ’31 : 384-387. Its central part is a Moll large surface ther¬

mopile made by Kipp and Sons of Delft, Holland.

The instruments for reading the electric currents from the

thermopile were the two millivoltmeters described in the same

paper. The most sensitive range has a scale of 100 divisions for

0.333 millivolt. There was also a new and much more sensi¬

tive instrument — a galvanometer of the D’Arsonval type, made

by Leeds and Northrup, of Philadelphia. This galvanometer

is of the construction called 2500 A in their catalogue. It is

furnished with a street-tripod and has an attached telescope and

scale. The scale extends to 250 divisions on each side of the

central zero and the sensitivity of the galvanometer is 0.5 mi¬

crovolt for one scale-division. The arrangement of tripod and

galvanometer is that listed under number 2123 in the catalogue.

This galvanometer must be set up on shore and connected

by a cable with the boat from which the pyrlimnometer is op¬

erated. The cable is 183 m. (600 ft.) long and it fixes the limit

of distance between boat and shore, thus determining the depth

to which observation may extend. In the smaller lakes the

maximum depth would be reached, or at any rate a depth so

great that the limit of observation is determined by the instru¬

ments. In larger lakes, whose bottom has a gentle slope, the

possibilities of the pyrlimnometer were not exhausted at the

depth which could be reached.

The tripod and telescope have proved entirely satisfactory as

mounting for the galvanometer ; there has been no trouble from

unsteadiness. A hood of nickle-plated metal was provided for

the galvanometer in order to prevent any possible disturbing

effect of the sun on its readings; but it was not found neces¬

sary to use it, since the banks of the lakes are densely wooded

almost everywhere.

A series of shunts is provided by which five ranges of sensi¬

tivity can be used with the galvanometer ; these were numbered

0, 1, 2, 3, 4, in the order of sensitivity; range 4 giving the

maximum sensitivity of the instrument. Range 0 gave about

the same scale reading as that of range 2 of the millivoltmeter ;

range 4 gave a scale reading about 210 times as great as

range 0.

530 Wisconsin Academy of Sciences , Arts , and Letters .

Observing

Observation regularly began and ended with a reading in the

air and one or more air readings were made during longer

series. All observations were made in direct sunshine. Below

the surface readings were made at the depth of one meter and

below this ordinarily at 3 m., 5 m., etc., until readings became

small, when they were made at every meter. In lakes with high

colored water they would be made at every meter from the

first, and in extreme cases at every half meter. Readings in air

were made with the millivoltmeter and the 20 range, i. e., the

range in which 100 divisions of the scale indicate 20 millivolts.

The reading at one meter was regularly made with the milli¬

voltmeter and the 2 range; and readings were continued with

the 2 mv. range and the 0.333 mv. range so far as these gave

an adequate movement of the needle.

In general change was made from the 0.333 range of the mil¬

livoltmeter to the 3 range of the galvanometer, which gives a

scale-reading 10 times as great for the same amount of radia¬

tion. As readings became smaller change was made to the 4

range of the galvanometer. In all cases of change of range or

instrument readings were made with both types at the same

depth.

Computation of Results

Transmission in the first meter. No attempt was made to de¬

termine the rate of transmission of radiation through the suc¬

cessive parts of the upper meter of the lake: comparison is

made directly between the reading in air and that at the depth

of one meter. Two computations are needed: (1) The ratio

of the scale-reading of the 20 mv. range to that of the 2 mv.

range. This has been determined by numerous comparisons

as 2.5 (’29 : 513). (2) The ratio between readings of the in¬

strument in air and in water : a difference due to the glass cover

of the thermopile. It has been determined and computed as

stated in ?29 : 515; the thermopile gives in air 81 per cent of

the full reading and 95 per cent in water. This correction may

be combined with that for the difference in the scale-reading of

the two ranges by multiplying the reading of the 20 mv. range

by 2.94 instead of 2.5.

Birge & Juday — Solar Radiation and Inland Lakes. 531

Transmission below one meter. In any complete series read¬

ings were likely to be made with two instruments and four

ranges; 2 and 0.333 of the millivoltmeter and 3 and 4 of the

galvanometer. These must be reduced to a common value ; and

for the same amount of radiation the relative values of the

scale-readings are as follows :

One scale-division of 2 mv. equals 6 divisions of 0.333 mv.

One scale-division of 0.333 mv. equals 10 divisions of 3 gal.

One scale-division of 3 gal. equals 3.5 divisions of 4 gal.

It thus appears that scale readings from the 2 mv. range must

be multiplied by 210 in order to express them in terms of the

4 gal. range. Those from the 20 mv. range in air must be mul¬

tiplied by 617 (2.94 X 210) in order to express them on the

4 gal. scale as used in water.

The values of readings below one meter were first computed

as percentages of the one-meter reading; the value of the one-

meter reading was determined as a percentage of that in air;

and from these data was computed the value of subjacent read¬

ings.

Reduction to Zenith Sun. The percentages thus obtained are

platted on semi-logarithmic paper at the proper depths and the

points are connected by straight lines so as to constitute the

curve of observed transmission. This is changed to the curve

for zenith sun by the method stated in ’30 : 289. The altitude

of the sun at the time of observation is taken to the nearest de¬

gree ; and the value of the cosine of the angle of refraction cor¬

responding to this altitude is taken to the nearest centimeter.

At any depth of the observed curve the distance is measured in

millimeters from the 100 per cent line to the intersection of the

curve ; this number is multiplied by the cosine ; and the product

is the distance from the 100 per cent line to the value of the

percentage for zenith sun. In this way a new curve is con¬

structed, that for zenith sun, and the new percentages are taken

from it. In practice the cosine used is ordinarily that for the

mean altitude of the sun during the observations; it has not

been thought necessary to adjust for changes in altitude at

every successive reading.

All percentages given in this paper are expressed in terms of

total radiation delivered to the surface of the lake, including

ultra-violet and infra-red. It has been already stated that all

532 Wisconsin Academy of Sciences, Arts, and Letters .

radiation below one meter may be considered as light. In the

solar energy spectrum the amount of energy between 3000 A

and 7640 a varies with the air-mass and the amount of pre-

cipitable atmospheric water; and may lie between 58 per cent

and 33 per cent. If, therefore, the percentages given in this

paper are doubled they will roughly express the percentile en¬

ergy equivalent of the radiation delivered to the surface in the

form of light.

Chapman (’31 : 308) states that the pyrlimnometer will

“give us the depth to which the heat rays penetrate”. By this

he probably means the heat equivalent, at various depths, of the

radiation which is measured; not the penetration of the infra¬

red or so-called “heat rays”. But such results may be expressed

in any units which are desired; heat, light, or electricity.

Pietenpol (’18 : 574), relying on the “corresponding slopes

of the curves” which show the coefficient of absorption of light

in filtered and unfiltered water, states that the action of unfil¬

tered water on light is non-selective. But Harvey (’28 : 158)

points out that Pietenpol’s measurements of transmission in fil¬

tered and unfiltered water from Lake Mendota indicate that

turbidity has a selective influence. The measurements are as

follows :

Wave length Color Transmission

Filtered TJnfiltered

4900 A Blue 72.5 17.0

5580 A Yellow 83.0 31.5

The rate of transmission of blue in the filtered water is 87 per

cent of that of yellow ; in unfiltered water blue has a transmis¬

sion only 54 per cent of that of yellow. It appears therefore

that the particles in the unfiltered water have a considerable

selective capacity. Probably also much of the difference be¬

tween blue and yellow in the filtered water is due to particles

remaining after filtration.

Accuracy of Observations

In general duplicate readings at the same depth are in good

agreement, differing not at all or by one or two divisions of the

scale. Such correspondence was especially close in the epilim-

nion where the temperature of the water is constant during

the observations. Illustrations may be taken at random from

great numbers. In Big Lake readings at 3 m. were 18 and 19

Birge & Juday — Solar Radiation and Inland Lakes . 533

scale-divisions; in Pauto Lake (Aug. 4) at 1 m. they were 31 -H

and 32 — ; at 3 m., 91 and 94; at 11 m., 51, 56, 57; at 15 m.,

13, 15. The third reading at 11 m. was taken because of the

divergence of the first two. The difference in the numbers for

the several depths is due to the use of different sensitivities.

In the case of Pauto Lake the entire series was read without

difficulty, but in many other cases records of series in the ther-

mocline or below might be marked “galvanometer unsteady”

and the results of such readings may be seen as irregularities

in the curves of Figs. 1-3. It also happens that the sun varies

during the period of observation and that readings are thrown

out of line from this cause. The series in Nelson Lake (Fig.

1) shows the result of selecting concordant readings from a large

number made under conditions of varying sun. Small clouds

were forming and disappearing and in such case the amount of

radiation may vary from minute to minute, even when the sun

is unclouded. Much time was consumed in waiting for favor¬

able minutes and the six determinations platted in this curve

are the most concordant out of 28 that were taken.

Study of the details of the curves given in Figs. 1-3 will show

many cases where we should expect the per cent to be higher or

lower than that recorded. The curve from Big Carr Lake (Fig.

1) shows perhaps the most conspicuous irregularities. Some

of them probably represent facts in the lake at the time of ob¬

servation ; others represent accidents of sun or instruments ; but

none are so great as to cause any serious doubt as to the gen¬

eral value of the transmission. In other cases single readings

seem to be out of line. In Pauto Lake Aug. 4, (Fig. 1) the read¬

ing at 11 m. should probably be higher, perhaps 0.23 per cent

instead of 0.19. There are obvious small irregularities in the

two curves for Clear Lake (Fig. 3) and also in the lower part

of that for Trout Lake, and for these we have no definite expla¬

nation. On the other hand, the sudden drop in transmission

at 13 m. in the curve for Crystal Lake, Aug. 19, (Fig. 1) has

an adequate explanation (p. 541) and no doubt other similar

changes in transmission represent temporary local opacities and

transparencies in the water.

Perhaps it is not out of place to state that we have been sur¬

prised at the general agreement of observations and the satis¬

factory nature of the conclusions to be drawn from them, rather

than disappointed by unexplainable irregularities.

Table I. General results. Transmission as observed and as computed for zenith sun. Char. Trm. = characteristic transmission) galv = gal¬

vanometer; Inst. = instrument) Max. — maximum) mv =millivoltmeter) Obs. = observed) Trp. = transparency) Z. S. = zenith sun.

534 Wisconsin Academy of Sciences , Arts, and Letters ,

Birge & Juday - — Solar Radiation and Inland Lakes . 535

. V

“I

§.s

si

H JJSK

' r-H p> w

«H «H P »r« Pi 4m

O O > OJO W W

536 Wisconsin Academy of Sciences, Arts, and Letters,

Fig. 1. Curves of observed transmission for a series of lakes, ranging

from the most opaque to the most transparent. Note the two curves for

Crystal Lake and for Pauto Lake, showing the general loss of transpar¬

ency as summer advances, while the character of the curve is retained. In

Diamond Lake the marked loss of transmission in the lower water is due

Birge & Juday— Solar Radiation and Inland Lakes . 537

Observations on Lakes, 1931

The general results of the work of 1931 are given in Table

I and in Figs. 1-6. Figures 1-3 show the curves of observed

transmission of radiation and Figs. 4-6 those of transmission

computed for zenith sun. Table I gives a summary of both

sets of curves and also adds other pertinent details. The lakes

are arranged in the table in the order of color and transpar¬

ency, as in our third report (’31). Other facts regarding these

lakes are stated in Table VI, p. 562.

In Figs. 1 and 2 are shown series of observations in lakes

ranging from those with low transmission to those which are

very transparent; and the curves for zenith sun are given for

the same lakes in Figs. 4 and 5. Figures 3 and 6 contain five

sets of observations from lakes whose transmission in the upper

water is very similar, while in the deeper water there is wide

divergence. On the same figures curves are also shown for two

lakes for which there was no place in the other figures.

These transmission curves resemble in general those given in

earlier reports (’29 : 527, ’30 : 303, ’31 : 397) but with two im¬

portant differences. The curves are carried out to smaller val¬

ues of radiation and to greater depths in the several lakes ; and

in many cases they show great changes in transmission as radi¬

ation passes into the deeper water. In the former reports very

few curves extended beyond the region of characteristic trans¬

mission; Marl Lake (’29 : 527) being the most conspicuous case

of a curve where there was marked difference between the epi-

limnion and the deeper water. One main purpose of the work

of 1931 was to ascertain whether such differences are not com¬

mon phenomena in small lakes and this was found to be the

case. In all cases observation went to a considerable distance

below the epilimnion, which even in Trout Lake does not be¬

come more than 8 m. thick during the summer. In most lakes

its thickness is 5 m. or less, especially in lakes with high col¬

ored water.

to proximity to bottom of lake. The same is seen in the curves for Muskel-

lunge and Big lakes, in these cases due to turbidity in bottom water of the

slope of the lake. In both cases characteristic transmission would go on

without such change in the deeper water of the lakes. Big Carr Lake

shows an unusually irregular curve, due to accidents of observation.

Curves for Helmet, Mary, and Turtle lakes show increase of transmission

with depth (see p. 554). Curves for Pauto and Day lakes show curves

for lakes with increasing turbidity in hypolimnion.

538 Wisconsin Academy of Sciences, Arts, and Letters .

The 25 lakes may be classified according to their character¬

istic transmission with zenith sun, as was done in a former

report ('30 : 305). The result is as follows:

Lakes with low transmission (6 — 30), Helmet, Little Long, Little

Pickerel, Mary, Turtle.

Transmission low medium (31 — 50), Adelaide, Big, Bragonier, Midge.

Transmission high medium (51 — 70), Big Carr, Little Tomahawk,

Muskellunge, Nelson, Plum, Silver, Tomahawk, White Sand.

Transmission high (71 — 85), Clear, Crystal, Day, Diamond, Little

Bass, Pauto, Trout, Weber.

Since the earlier observations hardly went beyond the region

of characteristic transmission, a more convenient classification

for the purposes of this report groups the lakes according to

the results reached by the use of the several kinds of instru¬

ments and the types of transmission found. This method gives

five groups as follows :

1. Transparent lakes in which the characteristic transmission

is high and extends with little change either to the bottom of

the lake or nearly to that depth. To this group belong Crystal,

Diamond, and Weber lakes ; they belong to that group on which

detailed report was made by us in 1930 (’30 : 291-302; Fig.

2-7).

2. Lakes in which characteristic transmission continues near¬

ly or quite to the instrumental limit of observation without es¬

sential change; Adelaide, Big Carr, and Nelson lakes. This

statement, in a modified form, also applies to the more opaque

lakes in the list. See group 5 below.

3. Large and deep lakes, as lakes go in the Highland Lake

District, in which observation was limited in depth by the

length of cable from shore. In these lakes observation did not

extend beyond the region of characteristic transmission, and

this, whatever may be its numerical value, should continue be¬

yond the depth reached by observation. Here belong Big, Clear,

Muskellunge, Tomahawk, and Trout lakes. Little Tomahawk

Lake belongs in this class, so far as the character of the curve

is concerned ; but observation ended so close to the bottom that

transmission can not continue much farther without change.

4. Lakes in which there is a notable decrease of transmission

in the deeper water. Here belong Day and Pauto lakes (Fig.

1), Midge, Plum and Silver (Fig. 2), White Sand and Little

Bass lakes (Fig. 3). These lakes offer the most interesting ad¬

dition to our knowledge made by the work of 1931.

.01 Per Cenl

Birge & Juday — Solar Radiation and Inland Lakes . 539

540 Wisconsin Academy of Sciences, Arts, and Letters.

5. In the lakes with most deeply stained water observations

in earlier years hardly went deeper than 2 m. The galvanome¬

ter enabled us to carry readings to greater depths in 1931 and

showed that there might be a considerable increase of trans¬

mission with depth; an increase probably associated with the

reduction of the solar spectrum to that region which passes

most freely through water of the particular color of the lake.

This is also an interesting addition to our knowledge.

Group 1 . Transparent lakes with high transmission .

The observations of 1931 in these lakes were much like those

of 1929, which were reported in detail in the paper of 1930

(pp. 293, 296, 301), and there is little to add to that report.

In all of these lakes there is a perceptible, though small de¬

crease of transmission in the hypolimnion ; in Crystal and Web¬

er lakes this turbidity begins several meters below the epilim-

nion ; in Diamond Lake the hypolimnion occupies only the lower

2 m. or 3 m. of the water and the whole of this is turbid to

much the same degree (’30 : 296) as is the bottom of the hypo¬

limnion in the other lakes. No growth of moss has been noted

on the bottom of Diamond Lake as it has been in Crystal and

Weber.

Crystal and Diamond lakes are the only lakes of this District

whose transmission has been regularly found to be above 80

for any considerable distance. No other Wisconsin lakes have

been found to equal them. Characteristic transmission as high

as 85 has been found in Crystal Lake, and even 90 has been

observed for strata of one or two meters (’29 : 559, ’30 : 293).

But these lakes are less transparent than the ocean. Poole

and Atkins (’29 : 324) found average transmission from 86 to

about 90 in the ocean near Plymouth. They report maximum

transmission up to about 95 and very few cases below 80 ; “the

most turbid water” had an extinction coefficient of 0.228, which

is equivalent to a transmission of about 79.5. On the other

Fig. 2. A second seTies of curves of observed transmission ranging from

low to high. Much the same types are present as in Fig. 1. Silver Lake

has a curve much like that of Day, but shows accidental irregularities in

lower water. Tomahawk Lake shows a great effect of bottom turbidity

found on the slope of the lake. Midge and Plum lakes have curves like

that of Day, but effect of turbidity of hypolimnion is less marked because

of color and turbidity of epilimnion. Bragonier Lake shows very great

and sudden effect of bottom turbidity; see p. 553.

Birge & Juday — Solar Radiation and Inland Lakes . 541

hand, Oberdorfer (’28 : 476) reports transmission from the

Bodensee to depths of 20 m., or more, and therefore comparable

with those from Crystal Lake so far as the depth reached is

concerned. The lake is far larger and deeper and is a charac¬

teristic oligotrophic lake of the first order in area and depth.

The mean transmission of eight series, each in a different

month, is 77.2 ; the maximum, 82.7 ; the minimum, 71.7. Per¬

centages at 1 m. (’28 : 482) were: winter, 81.2-88.0; summer,

69.0-72.5; these are percentages of incident light and would

correspond to about one-half that per cent of total incident

radiation. It appears therefore that the transmission of the

Bodensee closely resembles that of Crystal Lake, but is a little

lower.

Crystal Lake . (Figs. 1, 4.) Two series are shown in Figures

1 and 4; taken on July 10 and on August 19. Readings were

made with the millivoltmeter in the deep water of the lake;

that at 19 m. was about 0.5 m. above the mud.

In both series there is an increase of opacity in the lower

part of the hypolimnion; but transmission went on without

noticeable change from the epilimnion into and below the ther-

mocline ; as is also shown in the series reported in ’30 : 298.

On July 10 the transmission to 19 m. was 83 ; to 15 m. it was

85 ; there was 40 per cent of incident radiation present at 1 m.,

and this was reduced to 4 per cent at 15.3 m. In the August

series transmission was lower, being 78 to 18 m. and there was

38 per cent of incident radiation present at 1 m. and this was

reduced to 3.8 per cent at 12 m. There was an active growth

of Dinobryon in the lake on this date and this was the prob¬

able cause of the lower transmission. It was especially abun¬

dant at the depth of 13 m. as was shown by catches made with

the plankton trap. The effect of this condition on the light is

seen in the curve of observed transmission (Fig. 1). Secchi’s

disc was clearly visible a little above 13 m. and suddenly dis¬

appeared when lowered a few centimeters.

Diamond Lake. (Figs. 1, 4.) This lake presented about the

same condition in each of the three years in which it was vis¬

ited. There was an average transmission of 81 to the depth

of 9 m., followed by an abrupt decline of transmission below

that depth. Transmission from 3 m. to 9 m. was reported as

85 (’30 : 292) ; lower transmission in the upper water reduces

this to about 81. This lake has a flat bottom and the percentile

542 Wisconsin Academy of Sciences , Arts, and Letters .

meter and also in their characteristic transmission, but whose transmis¬

sion in the lower water is widely different. With these are the curves for

two lakes for which there was no place in the other figures. Trout Lake

Birge & Juday — Solar Radiation and Inland Lakes . 543

area and volume fall off rapidly with increase of depth. At 9

m. there is present about 35 per cent of the surface area but

only 8 per cent of the volume lies below that depth. The ef¬

fective length of the lake is about the same as that of Crystal

Lake and the wind would have about the same power of creat¬

ing currents. But in Crystal Lake the 9 m. level has about 55

per cent of the surface area and nearly 35 per cent of the total

volume lies below it. The wind currents therefore concentrate

the debris and plankton of Diamond Lake into a far smaller

volume of water than in Crystal Lake, with consequent greater

reduction of transmission.

Weber Lake. (Figs. 2, 5.) Two series were taken in this

lake, July 8 and July 15, each extending to 11 m. This repre¬

sented the deepest water which could be reached with the cable

from the shore. Characteristic transmission was 76, about the

same as in former years. The more complete curve shown in

’30 : 301 extends quite to the bottom of the lake and is in gen¬

eral identical with those of 1931. In this lake the characteristic

transmission extends without noticeable change into the hypo-

limnion and nearly to the bottom of the lake.

It thus appears that in these lakes observations can be ob¬

tained with the pyrlimnometer, reaching to the bottom of the

lake. In all cases transmission continues with little change

nearly or quite to the bottom ; only a small effect can be traced

to increased turbidity of the hypolimnion, due to debris or

growths of algae, except in the lower 2 or 3 meters.

Group 2. Lakes whose characteristic transmission extended to

the instrumental limit of observation.

In this group and the next one the use of the galvanometer

carried observation to greater depths than were formerly

shows an irregular series, but one which should extend with little change

to greater depths. This is true for Clear Lake also, whose two series show

the kind of resemblance and difference that would be expected from obser¬

vations made at about the same time. In White Sand Lake the whole

hypolimnion is turbid, while in Little Bass Lake transmission goes on with

small alteration almost to the bottom. In Little Tomahawk Lake there is

almost no change in transmission, although the deepest reading is only

about a meter above the mud and within three meters of maximum depth.

Note that White Sand Lake and Clear Lake had about the same amount

of radiation at the surface and at 7 m., while at 13 m. Clear Lake had

nearly 30 times as much as had White Sand.

544 Wisconsin Academy of Sciences , Arts , and Letters .

reached, but the character of the transmission did not suffer

any essential change. There are three lakes in this group ; two

with water of low color (10) and one, Adelaide, whose water

has a color of 36.

Adelaide Lake . (Figs. 1, 4.) This small, deep lake (22 m.)

is one of a group of moraine lakelets with deeply stained wa¬

ter ; Adelaide is the largest of the group and the one with low¬

est color. Its water is relatively clear and color must be the

main factor in cutting off radiation. Observations extended to

8 m., at which depth there remained 0.013 per cent of incident

radiation. The transmission curve shows small irregularities

but the mean transmission of 44 with zenith sun is maintained

to the limits of observation and there seems to be no reason

why it should not extend to considerably greater depths. There

was no distinct sign of a rise in transmission in the lower

meters observed, due to absorption of all radiation except that

most freely transmitted by the water. Thus there is present

a characteristic transmission for this lake ; in lakes with more

deeply stained water the situation is different, as is stated on

a later page.

Big Carr Lake . (Figs. 1, 4.) In this lake and in the next

one described, observations were more irregular than usual.

In Big Carr Lake the irregularities were accidental. The char¬

acteristic transmission as observed was 57, and with zenith sun

was 62, much the same as in earlier years. The last reading

was at 14 m., and gave 0.015 per cent of incident radiation.

The epilimnion is 6 m. thick. The lake is 22 m. deep and radia¬

tion might be transmitted to a considerably greater depth with¬

out much change of rate.

Nelson Lake . (Figs. 1, 4.) Readings in this lake were quite

irregular, due to variations in the sun. There were scattered

cumulus clouds, with wide spaces between them. All readings

were taken in full sun, but with such a sky there is sure to be

variation in radiation. The curve is platted from the most con¬

cordant readings. Characteristic transmission was 48 as ob¬

served, and 56 with zenith sun, extending without notable

change to 11 m., about 4 m. above the bottom. At this depth

the observed per cent of incident radiation was 0.012 with a

transmission at the rate of 45 from 7 m. down. This rate must

have fallen off greatly immediately below 11 m. since the bot¬

tom is so near. The epilimnion was 4 m. thick.

Birge & Juday — Solar Radiation and Inland Lakes. 545

of cos r stated in Table I. The changes thus made in the positions of the

curves for characteristic transmission bring these into place for direct

comparison with those published in earlier reports. The lower parts of

curves, like that for Pauto Lake, are treated in the same way; but where

transmission is rapidly changing, no great reliance should be placed on the

correctness of results. It is especially dangerous to attempt to prolong

such curves.

546 Wisconsin Academy of Sciences, Arts, and Letters .

Group 3 . Larger and deeper lakes.

In these lakes of large area and considerable maximum depth,

the radiation was followed to depths on the slope of the main

basin of the lake, determined by the length of cable connecting

the galvanometer on shore with the boat and pyrlimnometer.

In all cases the result lay within the region of characteristic

transmission and the depth was so small that the limit of ob¬

servation was not reached. Characteristic transmission prob¬

ably went on to greater depths. In most cases turbidity sharp¬

ly cut off transmission near the bottom.

Big Lake. (Figs. 1, 4.) Readings in Big Lake went to 7 m.,

about one meter below the epilimnion. At this depth there was

observed about 0.066 per cent of incident radiation, or about

0.10 per cent wtih zenith sun. The instruments would have

allowed readings to a somewhat greater depth, perhaps to about

9 m., where radiation would have been about 0.010 per cent.

With zenith sun the characteristic transmission is 49 ; 0.10 per

cent would come at 8:5 m. and 0.010 per cent at 12 m. The

maximum depth is 18 m., so that transmission might continue

uniformly to that depth; but the lake is large and turbid, and

the area of deeper water is small. It is therefore quite prob¬

able that transmission began to fall off at a less depth.

Clear Lake. (Figs. 3, 6.) There are two series from this

lake taken on July 13 and July 18, and extending to the depth of

18 m. They are platted together in Fig. 3 and show the kind of

close general resemblance and difference in detail which would

be expected in such a lake from series taken so near each other

in time. In each series there was a more transparent stratum,

2 — 3 m. thick, near the depth of 10 m. where transmission rose

to 80 or slightly more. This region was in the upper part of

the hypolimnion. In the series of July 13 the reading at 18 m.

was 0.016 per cent of incident radiation ; at 17 m. it was 0.063

per cent, indicating a transmission of only about 25 in the

bottom meter of water. This reading, as well as others of a

similar kind, shows that there may be an accumulation of sus¬

pended matter close to the bottom of the water even on slopes

and much above the greatest depth.

The data show that 0.10 per cent of incident radiation would

be found at about 18 m. with zenith sun. The same rate of

transmission would call for 5-6 m. more to reduce the amount

Birge & Juday — Solar Radiation and Inland Lakes. 547

548 Wisconsin Academy of Sciences , Arts , and Letters.

to 0.010 per cent. Since the maximum depth of the lake is 28

m. it is quite possible that transmission may continue at this

rate to 28-24 m. But the lake is large, the bottom is irregular,

and the area of the deep water is small, so that it is not improb¬

able that opacity increases rapidly with depth below 18 m.

Muskellunge Lake. (Figs. 1, 4.) Readings in this lake went

to the depth of 10 m., where there was found 0.18 per cent of

incident radiation. The characteristic transmission as observed

to 9 m. was 62 ; falling off to 45 between 9 m. and 10 m. This

change is due to the influence of bottom debris. The charac¬

teristic transmission with zenith sun was 67 ; and since the

maximum depth of the lake is 21 m. this rate might have been

found to extend several meters farther. The epilimnion is 7 m.

thick.

Tomahawk Lake. (Figs. 2, 5.) This lake was observed to

the depth of 18 m. on the side of the slope extending to the

greatest depth of water at 22 m. The conditions are not unlike

those of Clear Lake. The mean transmission was 70 to the

depth of 11 m. with a great and rapid decrease in the water

below that depth, due to bottom turbidity. The observed trans¬

mission would bring 0.10 per cent of incident radiation at about

13 m. ; and the transmission with zenith sun would place it at

about 16 m. But the great size of the lake and the small vol¬

ume of the deeper water render hazardous any extrapolation

of observed results into deeper water than that investigated.

Trout Lake. (Figs. 3, 6.) In this lake the limiting depth was

15 m. and at that limit there was observed 0.044 per cent of in¬

cident radiation ; at this rate reading might have gone 2 m. — 3

m. further. The characteristic transmission was 71 with zenith

sun, although there was a good deal of irregularity in the read¬

ings; for which see Fig. 3. Ordinarily the transmission in

Trout Lake is less than 70, as is shown in Table III. In the

series of 1931, 0.010 per cent of incident radiation would have

been found at about 17 m. as observed, and at about 20 m. with

zenith sun. This is probably about the maximum clearness of

the water for summer, and since the greatest depth is 35 m.

transmission at this rate might go on to even deeper water.

Fig. 3 shows that there was a considerable decrease of trans¬

mission in the hypolimnion; but no marked decrease near the

bottom.

0,01 Per Cent .03 .04.05.06 .08 010 _ 02 03 04 030.6 03 1.0

Birge & Juday — Solar Radiation and Inland Lakes . 549

Fig. 6. Curves of Fig. 3 computed for zenith sun. See explanation of Fig. 4.

550 Wisconsin Academy of Sciences, Arts, and Letters .

Group k. Lakes in whose hypolimnion there is a notable

decrease of transmission.

These lakes have a wide range of characteristic transmission ;

high (71 — 73) in Day, Panto, and Little Bass lakes; high-medi¬

um (56—60) in Silver, Plum, and White Sand; low medium

(48) in Midge Lake. In these lakes mechanical obstruction to

the passage of radiation is present in the hypolimnion, not

merely in water close to the bottom ooze (as in Diamond Lake)

but also to a considerable distance above it. The part of the

hypolimnion thus affected differs in the different lakes, reach¬

ing a maximum in White Sand Lake ; where there is a sudden

change of transmission at the junction of epilimnion and hypo¬

limnion, with a fairly uniform and low transmission until the

limit of observation is reached.

Table II shows the details of the observed transmission in

these lakes. It brings out facts which are hardly noticeable in

the diagrams. Such are the increase of transmission at the

Table II. Observed percentages and transmission .

Column headed “Epi” gives the thickness of the epilimnion in meters.

Birge & Juday- — Solar Radiation and Inland Lakes, 551

thermocline in Day Lake; the similar increase for a consider¬

able distance in Panto Lake; the greater turbidity of White

Sand Lake at the thermocline.

Day Lake, (Fig. 1, 4.) The lower water of this lake was

much less transparent in 1931 than it was in 1929 (’30 : 295).

The situation appears in the following table :

Percentage of incident radiation observed in Day Lake

at the depths indicated

The characteristic transmission, with zenith sun, to 9 m. was

not very different — - 75 in 1929, 73 in 1931. In 1931 trans¬

mission below 9 m. fell off rapidly. In 1929 there was nearly

three times as much radiation at 10 m. as in 1931; at 12 m.

four times; and at 14 m. nearly 20 times as much. This is a

good illustration of the ordinary constancy of the characteristic

transmission from year to year and of the great differences in

deeper water where turbidity is likely to change.

Little Bass Lake, (Figs. 3, 6.) The values of radiation in

this lake were read with the millivoltmeter and therefore were

followed only to 11 m., where there was found 0.16 per cent

of incident radiation. The lake is about 14 m. deep and radia¬

tion is probably very rapidly absorbed in water below 11 m.

Transmission was little altered between surface and 8 m., where

the temperature of the water was 15.7°, the epilimnion having

22.0°. Between 8 m. and 9 m. transmission was 40, while be¬

low, 9 m.-ll m., it was 50. This reading may be entirely cor¬

rect or there may have been variations in radiation to account

for part of the difference. There were white cumulus clouds in

the sky, with large spaces between them. All readings were

made in full sunshine, but it is probable that the value of radia¬

tion was variable from minute to minute.

Midge Lake, (Figs. 2, 5). Four series of observations were

taken in Midge Lake during the summer, of which that of Aug.

5 is the most complete; the other agreed with it in general.

Radiation was followed to 9 m., the maximum depth being 11.6

m. with very soft oozy bottom. The lake is surrounded by bog

and the water is colored and turbid. The observations of Aug.

5 came in a very warm period of the summer and the tempera-

552 Wisconsin Academy of Sciences, Arts, and Letters.

ture of the water declined from the surface, there being no

real epilimnion. The lake is so small that the wind is unable

to establish a regular epilimnion unless aided by cool days and

nights. The surface temperature was 26.7° ; that at 4 m. was

18.8° ; at 7 m. 8.2°. Transmission was 49 in the 1-3 m. stratum;

46 between 3 m. and 5 m. ; and 40 between 5 m. and 7 m. Be¬

low 7 m. there was a sudden decline to about 31. This indi¬

cates turbidity increasing with depth and having a marked rise

as the bottom is neared. Color also rises and shows a sharp

increase at 9 m. The color readings were: surface, 20; 5 m.,

27 ; 9 m., 40. In this lake was found the minimum per cent

of incident radiation observed during the summer — 0.0035 per

cent at 9 m. The area of the 9 m. level of the lake is about

13 per cent of the surface area, and the volume of water below

9 m. is about 3 per cent of the whole. The small remainder

of radiation left at 9 m. must be extinguished before penetrat¬

ing much farther.

Panto Lake. (Figs. 1, 3, 4, 6.) Three series are included

from this lake in order to show differences in transmission due

to increase of turbidity during the summer. In all cases the

characteristic transmission extended to 11 m. or more, and

therefore well into the hypolimnion ; since the epilimnion at this

time of year is about 5 m. or 6 m. thick. The series taken in

August showed a lower characteristic transmission than those

in July — 73 as compared with 78 or 79 in earlier series. The

percentage of incident radiation present at 1 m. was also

smaller in August — 27 as compared with 38 or 40. Both

facts indicate increased turbidity in the upper water, probably

due to plankton. In all series it appears that the lower water

was turbid, and that the relation between it and the upper wa¬

ter was much the same at all dates. Readings on August 4 went

to 15.7 m., close to the bottom at the place of observation.

There was observed at that depth 0.012 per cent of incident

radiation; with the same transmission and with zenith sun

there would have been found at 16 m. about 0.01 per cent.

Plum Lake . (Figs. 2, 5.) This large and generally shallow

lake has one relatively small area near the center where the

depth is 18 m. The color of the water is 16 and there is much

turbidity at all depths. The characteristic transmission, com-

Birge & Juday — Solar Radiation and Inland Lakes. 553

puted for zenith sun, was 60 (observed transmission, about

53) and extended to nearly 9 m. The epilimnion was 5 m.

thick. At 9 m. there was an abrupt change in transmission,

which was 38 to 11 m., the limit of observation; at this depth

there was left 0.016 per cent of incident radiation.

Silver Lake. (Figs. 2, 5.) Radiation was followed in this

lake to the depth of 15 m. (maximum depth 19.5 m.), where

there was observed 0.006 per cent of incident radiation. Read¬

ings below the surface were made at 1 m., then at 5 m., 7 m.,

etc. Observed transmission 1 m. to 5 m. was 67, and declined

to 59 in the 5 — 9 m. stratum, indicating greater turbidity of

water from the epilimnion down. In this respect the lake shows

a transition from lakes like Pauto, where the upper hypolim-

nion is as transparent as the epilimnion, to lakes like White

Sand, in which the whole hypolimnion is turbid to a fairly uni¬

form degree.

White Sand Lake. (Figs. 3, 6.) The characteristic transmis¬

sion of this lake in 1931 was 70 with zenith sun; this is much

higher than in former years, when it was 50 — 55 (Table V).

Characteristic transmission ended at the thermocline and was

succeeded by a much lower one in the stratum 7— -13 m. Trans¬

mission between 7 m. and 9 m. was especially low, about 38 as

observed. At 13 m. the observed per cent was 0.014, and with

zenith sun 0.01 per cent would be found close to 15 m., about 6

m. above the bottom of the lake. Here the whole hypolimnion

was much more opaque than the epilimnion; the difference be¬

ing due not to color but to turbidity, as was shown by centri¬

fuge catches. The relatively large area of the lake enables the

wind to set up currents of considerable power, which sweep the

lighter particles of debris from the bottom of the shallower

parts of the lake and accumulate them in the deeper water.

This was the best example found in 1931 of a lake whose

whole hypolimnion was made turbid by debris so that there was

a sharp difference in transmission between it and the epilim¬

nion. Among smaller lakes Hillis offered a somewhat similar

case in 1929 (’30 : 300), as also did Finley (’30 : 298) ; but in

these cases color of the lower water had much influence, as well

as turbidity.

Bragonier Lake. (Figs. 2, 5). This small lake may well be

treated here. It is small and shallow and the deeper water has

554 Wisconsin Academy of Sciences , Arts , and Letters .

a very small area and little volume. The shores are largely of

bog and marsh ; the bottom is of soft ooze ; there is much color

and much turbidity in the water. Characteristic transmission

was 41 with zenith sun and went to 4 m. ; below that depth there

was a great and rapid increase of turbidity and also of color,

with corresponding decrease of transmission. Observation end¬

ed at 5.5 m. where there remained 0.017 per cent of incident

radiation.

This lake was visited in 1930 031: 397) and a comparison

of the curve there shown with that in the present paper will

illustrate the kind of additional knowledge brought by the use

of the galvanometer. In the earlier year observation could not

go below the depth of the characteristic transmission. The

whole hypolimnion of the lake may well be regarded as “bot¬

tom water”, in view of its small volume and the great amount

of debris which it contains.

Group 5. Lakes with deeply stained water.

There is a very definite gap in the series of colors reported in

Table I, coming between Adelaide (36) and Turtle (68) ; and

the remainder of the list consists of lakes with higher color

than Turtle. There is a difference also in the type of transmis¬

sion found in these high colored waters and in those which are

more transparent. In the case of the latter type there is a mean

transmission (the “characteristic transmission”) which can be

stated and which usually extends through the epilimnion or

deeper. The transmission in the 1-2 m. stratum may be some¬

what lower than that in the meter below; and it should be, so

far as the general effect of the water on solar radiation is con¬

cerned (’29 : 533) ; but the difference is not great and is often

obscured or eliminated by variable accidents, such as the

amount and position of plankton.

In each of the five lakes with high colored waters there is a

marked difference between transmission in the 1-2 m. stratum

and in that immediately following. The situation as shown by

the transmission computed for zenith sun is given in the notes

to Table I. The following table indicates it as directly observed.

In each lake the transmission rises in the 2-3 m. stratum as

compared with that in the meter above. The difference may be

strikingly large and may increase in lower strata. It is least

Birge & Juday — Solar Radiation and Inland Lakes . 555

Table III.

in Little Pickerel Lake, which is a shallow lake with boggy

shores, and whose water is not only high colored but also turbid

with a great amount of organic debris. The other lakes are

much less turbid, except sometimes the surface meter of Lake

Mary. Turtle Lake, which has the lowest color has also the

smallest increase with depth among the lakes with clearer

water.

We can not speak of a “mean transmission” in the upper wa¬

ter of these lakes as we do for the others on our list. The

transmission in the 1-2 m. stratum of these lakes is in some¬

what the same condition as that in the 0-1 m. stratum of all

lakes. Absorption goes on here much more rapidly than be¬

low; so much more rapidly that no significant mean can be

made by combining the record of this stratum with that of

those below. A reason for the situation lies in the high color

of the water. This absorbs very rapidly the short-wave part

of the spectrum and quickly reduces it to the part which most

readily passes through water in this particular color. This is

one cause, and perhaps the main one, for this rise of transmis¬

sion.

The situation in lakes of this type is shown in ’31 : 415, Fig.

20 and especially in ’30 : 331, 333, Figs. 14, 15. The last named

figures show for individual lakes the relation between the trans¬

mission of total radiation and that of the several colors. In

lakes whose water is colored but not deeply stained, the maxi¬

mum transmission is that of the central or yellow region of the

spectrum; with increase of color the transmission of yellow

falls off, both absolutely and relatively, so that it becomes less

than that of total, which comes to coincide with red. It is a

rough sort of division of the spectrum that is effected by these

light-filters, but it shows that the light must soon be reduced

556 Wisconsin Academy of Sciences , Arts , and Letters .

to that part whose color is approximately that transmitted by

the water. During this process of reduction there must be an

increase of the rate of transmission of the remaining radiation,

whose absorption is minimal for the lake under observation.

Observations with results similar to these have also been

made by Poole (’30 : 142-149) in Lough Bray, Ireland, to the

depth of 4 m. In this lake the per cent of incident light present

at 1 m. was 0.5 ; transmission, 1-2 m. was 7 ; 2-3 m., 28.6. These

observations were made with a neon discharge tube which re¬

sponds to short-wave radiation only. Poole’s explanation of the

rise in transmission is that “longer waves penetrate deeper”,

and this, within limits, would be ours also. So far as we know,

these interesting readings are the only observations of the kind

that have been made in Europe.

There is a possible second cause for the rise in transmission

with depth, and at present we can not differentiate its effects

from those coming from color directly. In these lakelets with

high colored water, temperature and consequently density, fall

off rapidly from the surface. An extreme example was found

in Helmet Lake on July 29, where the temperatures were as fol¬

lows: surface, 29.2°; 1 m., 25.0°; 2 m., 18.9°; 3 m., 12.7°; 4

m., 9.3° ; 5 m., 7.9° ; 7 m., 7.6° ; 9.5 m., 7.4°. With such condi¬

tions of temperature rain or seepage water entering through

marginal bog and bringing color and turbidity would be con¬

fined to the water very close to the surface of the lake.

The difference in transmission between successive meters

seems to rise with color of the water ; but no regular series can

be made out. None would be expected, since more than one

factor is concerned. Mary Lake, for instance may or may not

have a considerable quantity of fine gummy debris in the sur¬

face water, and similar statements might be made for each

lake.

In all of these lakes transmission was followed about as far

as the instruments could go and in each case it is safe to ex¬

tend the curve of observed radiation to 0.010 per cent of inci¬

dent radiation. But it has not been thought quite safe to do

so for radiation with zenith sun. The quantity of radiation is

very small; the color of the water varies as also does the tur¬

bidity. For these reasons the depth stated for 0.010 per cent

Birge & Juday — Solar Radiation and Inland Lakes. 557

with zenith sun is bracketed ; but the position can not be in er¬

ror more than a small fraction of a meter. (See Table I).

The Minimum Observations.

A special interest attaches to the value of the smallest read¬

ings made, those that are near the limit of sensitivity of the

instrument. These usually come at the end of the series of

readings, since observations are made in descending order. In

many cases the pyrlimnometer was brought to the surface and

an air reading taken immediately after that at the greatest

depth. In other cases work was done at various depths before

coming to the surface. If the minimum reading is computed

as the last of a series which begins at one meter its value will

differ slightly from that obtained from computing it as a per¬

centage of the total radiation at the moment of observation.

But such differences are inconsiderable.

The minimum scale-reading was 1.3 divisions of the 4 range

of the galvanometer, and represented about 0.0035 per cent of

the reading in air. This last was 61 divisions of the 20 mv.

scale, equivalent to 37,200 of the 4 galv. scale. This observa¬

tion was made near the bottom of Midge Lake in 9 meters of

water. The last three discs of the pyrlimnometer were read

there twice with concordant results. In Silver Lake (15.7 m.)

and in Turtle Lake (6m.) minimum readings were obtained of

2 divisions of the 4 galv. range. In both cases the reading in

the air was low — 52-54 divisions of the 20 mv. range; and

the reading shows, at the depth named, about 0.006 per cent

of radiation incident on the surface. In 9 other lakes readings

of 4-7 divisions gave values of 0.011-0.018 per cent of the ra¬

diation in air.

No minute accuracy can be claimed for such small scale-

readings, though they were always repeated. There is often a

variation of reading, as in the case of White Sand Lake, where

readings equally good gave from 5 to 7 divisions of the scale.

Probably no great confidence would be placed in readings so

small if the pyrlimnometer had been lowered from the air di¬

rectly to the maximum depth; there is a great difference be¬

tween 4-7 divisions of a scale which is read directly, and the

30,000 or 40,000 divisions of the same scale, which are needed

to express the reading in the air. But the minimum is gradu-

558 Wisconsin Academy of Sciences , Arts, and Letters .

ally approached, in the series of observations, by reasonable

stages ; and Figures 1, 2, and 3 show that it plats into the curve

just about where extrapolation from the larger readings would

place it.

From these minimum readings may be computed the ex¬

treme limit for the present combination of pyrlimnometer and

galvanometer. Readings in air near noon in June and July

have given more than 100 divisions of the 20 mv. scale. This

would be equivalent to more than 60,000 divisions on the 4 galv.

scale ; a reading of two divisions of that scale would represent

about 0.0030 per cent of the radiation delivered to the surface

of the lake. Under such conditions, curves on semi-logarithmic

paper could be carried through five cycles, without danger of

serious error.

These actual and possible results may be compared with those

obtained by other types of instruments. Shelford and Gail

(’22 : 156), working in Puget Sound, found at 120 m. light to

the amount of 0.108 per cent of that present at the “wet” pho¬

tometer close to the surface. This would correspond to about

0.08-0.09 per cent of the light in the air. Poole and Atkins

(’29), working in the ocean near Plymouth, found 0.016 per

cent at 70 m., Jan. 3, 1928; 0.019 per cent at 60 m., July 23;

0.026 per cent at 40 m., Oct. 12. All of these observers used

photo-electric cells.

Oberdorfer (’28) used a photographic method in the Boden¬

see. He reports 0.02 per cent of incident light at 30 m. on one

occasion (’28 : 475). Apart from this case his minimum read¬

ing was 0.11 per cent at 25 m.

In all of these cases the base from which the percentage is

computed is the light, not the total radiation which we employ.

The percentages should therefore be divided by two in order

to make them roughly comparable with ours. All of the re¬

ported percentages seem to be those observed, not those ad¬

justed for zenith sun. So far, therefore, as the present rec¬

ord goes, all of these methods follow radiation to about the

same minimum value. It can be followed to about 0.01 per cent

of its surface value, and under favorable conditions readings

can be obtained which enable the observer to carry his curves

well into the next logarithmic cycle, which closes at 0.001 per

cent.

Birge & Juday — Solar Radiation and Inland Lakes . 559

Relation of Color, Turbidity, and Transmission

The number of observations is now large enough to permit

a provisional classification of lakes according to color and

transmission. There are not enough cases, especially in the

Table IV. Classification of Lakes by Color-group and Transmission.

groups with higher color, to give wholly satisfactory mean val¬

ues. But taking the facts as they stand in the table several

matters appear in them. (1) Transmission decreases as color

rises and with fair regularity, as is shown by Fig. 7. (2) The

range of transmission greatly overlaps in the several color

groups. (3) There is a wide range of transmission in each

color-group. (4) In spite of small numbers the maximum and

minimum in the successive color-groups show much the same

relation as do the means.

Fig. 7. Relation of color to transmission of radiation. The figure shows

the facts of Table IV. At the center of each color-group is platted the

mean transmission for that group, and with it are given the maximum and

minimum transmission found in that group.

560 Wisconsin Academy of Sciences, Arts, and Letters.

We seem warranted therefore in concluding that in all lakes

the characteristic transmission is determined by factors of col¬

or and turbidity ; that color is the main factor determining the

place of the lake in the general scale of transmission ; that tur¬

bidity causes much variation in transmission, causes overlap¬

ping of the several color groups, and wide range of transmis¬

sion in each group.

If the water of a lake were colorless and optically pure trans¬

mission near the surface would be 90 or more, increasing with

depth until it approached 98. In optically pure colored waters

transmission would be lower than in colorless water and would

follow the general order of the groups ; but in any group trans¬

mission would be higher than any recorded for that group in

the accompanying table.

Table V. Per Cent at One Meter and Characteristic Transmission with Zenith Sun.

Per Cent at One Meter and Characteristic Transmission

with Zenith Sun.

Table V brings together the observations on radiation in the

lakes of the list of 1931. Only one series is reported for each

year, and reference is made to earlier reports for the record

of lakes like Trout or Crystal which in some years were visited

several times.

Birge & Juday — Solar Radiation and Inland Lakes . 561

The table shows that in general the lakes retain their char¬

acters from summer to summer. Variations are often surpris¬

ingly small, as in Plum Lake. The greatest variability is shown

by the high colored lakes in which transmission is ordinarily

low. This variability is associated with the absence of a real

epilimnion during most of the summer. Radiation penetrates

into the water for a small distance only; wind has little effect

on these small lakelets ; and the temperature and density of the

water fall off rapidly from the surface. The result is a cor¬

responding change of color and turbidity in thin surface strata

of water, with accompanying alterations of transmission. In

these lakes transmission is given for the 1 — 2 m. strata only.

For transmission in deeper strata, as found in 1931, see p. 554.

Papers Cited

Anderson, J. S., 1930. Photo-electric cells and their applications; a dis¬

cussion at a joint meeting of the Physical and Optical Societies,

236 pp. London. See Poole and Poole, below.

Birge, E. A., 1922. A second report on limnological apparatus. Trans.

Wis. Acad. Sci. Arts and Lett. Madison. 20 : 512-551.

Birge, E. A., and Juday, C., 1929. Transmission of solar radiation by the

waters of inland lakes. Trans. Wis. Acad. Madison. 24 : 509-580.

Cited in this paper as ’29.

Birge, E. A., and Juday, C., 1930. A second report on solar radiation and

inland lakes. Trans. Wis. Academy. Madison. 25 : 285-335. Cited in

this paper as ’30.

Birge, E. A., and Juday, C., 1931. A third report on solar radiation and

inland lakes. Trans. Wis. Acad. Madison. 26 : 383-425. Cited in this

paper as ’31.

Chapman, R. N., 1931. Animal Ecology. McGraw-Hill Book Co. New

York. pp. i-x; 1-464.

Harvey, H. W., 1928. Biological Chemistry and Physics of Sea Water.

University Press. Cambridge, pp. i-x; 1-194.

Oberdorfer, E., 1928. Lichtverhaltnisse und Algenbesiedlung im Bodensee.

Zeitschrift filr Botanik. 20 : 464-568. G. Fisher, Jena.

Pietenpol, W. B., 1918. Selective absorption in the visible spectrum of

Wisconsin lake waters. Trans. Wis. Acad. Madison. 19 : 563-593.

Poole, H. H., and Atkins, W. R. G., 1928. Further photo-electric measure¬

ments of the penetration of light into sea water. Jour. Marine Biol.

Ass’n of United Kingdom. 15 : 455-483. Plymouth.

Poole, H. H., and Atkins, W. R. G., 1929. Photo-electric measurements of

submarine illumination throughout the year. Jour. Marine Biol.

Assyn of United Kingdom. 16 : 297-324. Plymouth.

562 Wisconsin Academy of Sciences , Arts , and Letters .

Poole, J. H. J., and Poole, H. H., 1930. The neon discharge tube photo¬

meter. In Anderson, above, pp. 142-149.

Shelford, V. E., and Gail, F. W., 1922. A study of light penetration into

sea water, etc. Pub. Puget Sound Biol. Sta. 3 : 141-175.

Table VI. General characters of lakes

Notes on Table VI

Column 4 shows the maximum depth, i. e., the depth as measured by

lead and line. It is ordinarily greater than would be found by a large flat

instrument like the pyrlimnometer, since the weight of the sounding line

sinks into the organic mud of the bottom. In lakes like Pickerel, Midge,

and Pauto there is no exact depth which can be called the bottom, since the

water passes almost insensibly into the soft organic ooze below it.

Column 5, Type. This classifies lakes as seepage (S) and drainage (D).

Muskellunge Lake, classified as seepage, once had a regular outlet and in

some years has now a very small overflow. Silver Lake ordinarily over¬

flows, but has no outlet in years with little rainfall. Little Tomahawk

Lake is at the head of a chain of lakes; it has an outlet but no affluent.

Column 6, Plankton. This column states in milligrams per liter the av¬

erage amount of organic matter in the centrifuge plankton from the sur¬

face water. The number of observations ranges from one (Helmet Lake)

to twelve (Crystal and Weber Lakes).

Column 7. Total organic. This states in milligrams per liter the aver¬

age amount of the total organic matter in the surface water, including

centrifuge plankton and dissolved material. It has been computed as

crude protein and as carbohydrate. Number of observations as in Column 6.

PROCEEDINGS OF THE ACADEMY

Sixtieth Annual Meeting, 1930

The sixtieth annual meeting of the Wisconsin Academy of Sciences,

Arts and Letters, in joint session with the Wisconsin Archeological So¬

ciety and the Midwest Museums Conference, was held in the Biology Au¬

ditorium of the University of Wisconsin, Madison, on Friday and Satur¬

day, April 11 and 12, 1930. The following papers were presented:

Friday morning, April 11: (1) Louise P. Kellogg: The Treaty at the

Cedars, 1836; (2) Albert B. Reagan: Some myths of the Hoh and

Quileute Indians; (3) May L. Bauchle: Wisconsin’s hidden jewel — Nasho-

ta; (4) Milton F. Hulburt: Sauk County Indian trails; (5) E. R. McIn¬

tyre: John Wesley Hoyt — western pioneer; (6) George Overton: Old

beach camp sites of the Winnebago County lake region; (7) Will F.

Bauchle: Trailing civilization by footprints; (8) Theodore Brown: Joseph

Jourdain, early French blacksmith; (9) John G. Gregory: Indians as

neighbors; (10) M. E. Hathaway: Barbed stone axes of Michigan.

Friday afternoon, April 11: (11) Geo. R. Fox: The archeology of the

Bahamas; (12) Paul B. Jenkins: The battle of Kings Mountain; (13) John

B. McHarg: Lincoln— things new and old; (14) Harry H. Clark: Lowell

and American political thought; (15) Ernst Voss: An edict of Philip,

Landgrave of Hessia, issued in 1539; (16) Rufus M. Bagg: The economic

and industrial development of South Africa; (17) John S. Bordner:

Evaluation of land in terms of utilization; (18) William W. Morris: Re¬

forestation problems in northern Wisconsin; (19) J. J. Davis: Notes on

parasitic fungi in Wisconsin, XVIII; (20) James A. Lounsbury: A

strange fungus from Transvaal soil; (21) Waldo E. Steidtmann: Studies

of growth rings of some tropical woods; (22) W. E. Tottingham and J. G.

Moore: Some effects of Vita glass upon plant growth; (23) A. H. Wiebe:

Pond culture.

Saturday morning, April 12: (24) Eric R. Miller: A century of pro¬

gressive changes of rainfall in Wisconsin; (25) Jeanette Jones: Notes on

the late Ordovician strata of the Green Bay-Lake Winnebago region; (26)

T. E. B. Pope: Wisconsin herpetoiogical notes; (27) A. A. Granovsky:

The use of the airplane in the control of forest insects; (28) J. B. Over-

ton : Seasonal variations between the hydrostatic and pneumatic systems in

trees; (29) W. T. McLaughlin: The origin of swamps and bogs; (30) L.

R. Wilson: Some plant remains in a peat bog; (31) Bernice I. Quandt:

Notes on some Washington County bogs; (32) N. C. Fassett: The ranges

of some Wisconsin plants; (33) H. A. Schuette: Early accounts of the

making of maple sugar; (34) Willoughby M. Babcock: Wing screen meth¬

ods of displaying coins; (35) Nile Behncke and Mrs. E. E. Rogers: Re¬

sults of special exhibits; (36) Edward R. Tyrrell: Materials used in

museum model construction (celluloid and Balsa wood).

564 Wisconsin Academy of Sciences, Arts, and Letters.

Business sessions were held immediately preceding each of the three

sessions for the reading of papers. The secretary presented the following

report, covering the period April 1, 1929 to March 31, 1930: Membership

as of March 31, 1930: honorary members, 5; life members, 14; corre¬

sponding members, 15 ; active members, 329 ; total, 363. Membership losses

during the year: deceased, 2; resigned, 1; dropped for non-payment of

dues, 15; total lost, 18. The deaths of the following members were an¬

nounced: Emil Godfrey Arzberger, January 29, 1930; William Christian

Sieker, December 1, 1929. The secretary presented the following applica¬

tions for membership, and on motion was unanimously instructed to east

the ballot of the Academy in their favor: Harry Hayden Clark, Madison;

Samuel Eddy, Minneapolis, Minn.; Waldo E. Steidtmann, Milwaukee;

Arthur N. Bragg, Milwaukee; W. T. McLaughlin, Madison; Leonard R.

Wilson, Madison; Jeanette Jones, Evanston, Ill.; Alphonse L. Heun, Mil¬

waukee; Alvin L. Throne, Milwaukee; Gilbert Raasch, Madison; Silas M,

Evans, Ripon. The nominating committee, consisting of George Wagner,

J. J. Davis and Charles E. Brown, reported nominations for the various

offices, and on motion the officers nominated were unanimously elected for

a term of three years: President , Charles E. Allen, Madison; Vice-

President in the Sciences , Rufus M. Bagg, Appleton; Vice-President in the

Arts , Otto L. Kowalke, Madison; Vice-President in Letters , William E.

Alderman, Beloit; Secretary-Treasurer , Lowell E. Noland, Madison;

Curator , Charles E. Brown, Madison; Librarian , Walter M. Smith, Madi¬

son; Committee on Publication: the president and secretary, ex officio;

Arthur Beatty, Madison; Committee on Library: the librarian, ex officio;

Howard Greene, Milwaukee; Mrs. Angie K. Main, Fort Atkinson; George

Van Biesbroeck, Williams Bay; R. C. Mullenix, Appleton; Committee on

Membership : the secretary, ex officio; Ralph N. Buckstaff, Oshkosh; E. F.

Bean, Madison; A. M. Keefe, West De Pere; W. N. Steil, Milwaukee. The

report of the Treasurer for the period, April 1, 1929 to March 31, 1930,

was presented as follows:

Receipts

Balance in State Treasury, April 1, 1929 . . . . .$2,023.75

Received from dues . . . . 288.50

Received from sales of Transactions, etc. . . . 129.62

Deposited from permanent fund . . . . .......... 909.45

State appropriation, July 1, 1929 . . . . 1,500.00

Total receipts . . . . .

Disbursements

To State Printing Board for printing _ _ _ _ _

Allowance for secretary . . .

Postage for mailing Volume 24 of Transactions

Other expenditures . . .

11 i

Total disbursements . . . . . . .

Balance in State Treasury, April 1, 1930 . . . . .

$4,851.32

$2,562.87

200.00

50.00

31.50

$2,844.37

2,006.95

Total

$4,851.32

Proceedings of the Academy.

565

Securities and Cash on Hand

City of Madison bonds . $1,100.00

Chapman Block bonds . 400.00

Commonwealth Telephone Company bonds . 200.00

Trust agreement, Centr. Wis. Trust Co . 1,000.00

4 certificates of deposit . 227.98

Cash . 13.58

Total . . . $2,941.56

The auditing committee, consisting of Lowell E. Noland and Ralph N.

Buckstaff, reported that it had examined the accounts of the treasurer

and had found them correct.

The annual dinner was held at the Memorial Union on Friday evening,

April 11. It was attended by 52 individuals. After the dinner S. A.

Barrett, President of the Academy, delivered an address on the subject,

“Tamest Africa”, which was illustrated with lantern slides and moving

pictures.

A committee appointed by President Barrett and consisting of three

members, Rufus M. Bagg, George A. West and T. E. B. Pope, presented

the following report: “Resolution of thanks to the University of Wiscon¬

sin and the members and officers of the Wisconsin Academy of Sciences,

Arts and Letters for their courteous reception and entertainment during

the convention held at Madison, April 11 and 12, 1930. We feel greatly

indebted to all the officers and directors for the very interesting educa¬

tional program presented and for their faithful services during the past

year.”

Chancey Juday,

S e ere tary- Tre asurer.

Sixty-first Annual Meeting, 1931

The sixty-first annual meeting of the Wisconsin Academy of Sciences,

Arts and Letters, in joint session with the Wisconsin Archeological Socie¬

ty and the Midwest Museums Conference, was held at Rip on College,

Ripon, on Friday and Saturday, April 10 and 11, 1931. The following

papers were presented:

Friday morning, Section A, in the Little Theater: Silas M. Evans. Ad¬

dress of Welcome; (1) Alton K. Fischer: Vertebral pathology of prehis¬

toric Wisconsin Indians; (2) George Overton: Silver ornaments from

Grand Butte; (3) Charles E. Brown: An effigy pipe from Pepin, Wiscon¬

sin; (4) John B. McHarg: Indian mound photography; (5) Robert R.

Jones: Central Pennsylvanian archeology; (6) Kermit Freckman: Pleas¬

ant Lake mound groups; (7) Alonzo W. Pond: A lower paleolithic site in

the Sahara desert.

Friday morning, Section B, in the Biology Lecture Room: (8) H. V.

Truman: The investigation of a post-glacial peat deposit near Lodi, Wis-

566 Wisconsin Academy of Sciences , Arts , and Letters .

consin; (9) L. R. Wilson: The interglacial forest bed at Two Creeks,

Wisconsin; (10) N. C. Fassett: An aquatic desert; (11) W. T. McLaugh¬

lin: Some noteworthy plants of the Fox River valley; (12) J. J. Davis:

Notes on parasitic fungi in Wisconsin, XIX (by title) ; (13) H. C. Greene:

Some studies on the myxomycetes of Wisconsin (by title) ; (14) William

W. Morris: Planting trees for profit in Wisconsin; (15) J. F. Groves:

Studies on hybrid barberry; (16) R. A. Brink and D. C. Cooper: The as¬

sociation of semisterile-1 in maize with two linkage groups.

Friday afternoon , Section A, in the Little Theater: (17) Elmer W.

Ellsworth: Varved clays of Wisconsin; (18) Ira Edwards: Observations

on the pleistocene of the Black River Falls quadrangle, Wisconsin; (19)

Albert B. Reagan: The Brush Creek region in northeastern Utah; (20)

Theodore T. Brown: The Albert H. Mill collection; (21) Will F. Bauchle:

Where the West begins; (22) Ernst Voss: Vom Schlauraffen Landt

(About the Land of Plenty) (by title); (23) Albert H. Griffith: Lincoln

literature, collectors and collections; (24) John B. McHarg: Early homes

of the Lincolns.

Friday afternoon , Section B, in the Biology Lecture Room: (25) D. C.

Cooper and R. A. Brink: Cytological evidence of segmental interchange

between non-homologous chromosomes in Zea mays; (26) L. J. Cole:

Crossing over in pigeon hybrids; (27) Agnes L. Zeimet: Embryo mor¬

tality in birds, particularly hybrids; (28) Alan Deakin: Pigmentation in

the mammary gland of swine; (29) S. X. Cross: The development of im¬

munity to the pathogenic effects of Trypanosoma lewisi in the rat; (30)

C. A. Herrick: Some relationships between the rat and the parasite Try¬

panosoma equiperdum; (31) Eric R. Miller: Sunshine and cloudiness in

Wisconsin.

Friday afternoon , General session in the Little Theater: (32) E. A.

Birge: Limnological investigations in the Trout Lake region of Wisconsin.

Illustrated by moving pictures.

Friday evening , General session in the College Chapel: (33) Alonzo

W. Pond: Reliving the past. Illustrated by lantern slides and moving pic¬

tures of North African explorations.

Saturday morning , Section A, in the Little Theater: (34) G. M. W.

Teyen: The relation of the museum’s library to the museum; (35) Ruth

Shuttleworth : Illustrating a museum special exhibit; (36) Frances S.

Dayton: A Winnebago camp site; (37) R. C. Corwin: Features of Great

Salt Lake; (38) W. D. Kline: Florida’s tung oil industry.

Saturday morning, Section B, in the Biology Lecture Room: (39) H. W.

Cornell: The mutual eclipses of Jupiter’s satellites to occur in the autumn

of 1931; (40) Edwin P. Greaser: The decapods of Wisconsin; (41) Lowell

E. Noland: Recognizing the species of bell-animalcules (Vortieellae) ; (42)

J. P. E. Morrison: Hydrogen-ion relations of certain molluscs in north¬

eastern Wisconsin lakes; (43) T. E. B. Pope: Wisconsin herpetological

notes; (44) Albert M. Fuller: Modern field methods in botany.

Saturday morning, General session in the Little Theater: (45) Huron

H. Smith : The ethnobotany of the Oneida Indians.

Proceedings of the Academy.

567

The annual business meeting of the Academy was held in the Little

Theater on Friday afternoon, April 10. The secretary presented the fol¬

lowing report on membership: Membership as of April 1, 1931: honorary

members, 5; life members, 15; corresponding members, 19; active mem¬

bers, 318; total, 357. Members lost during the year: deceased, 0; resigned,

15; dropped for non-payment of dues, 11; dropped for loss of address, 1;

total lost, 27. The following names were presented by the secretary for

election to membership: Milivoye Trifounovitch, Brussels, Belgium; S.

Kabboor, Bangalore, India; Morris A. Gilbert, Milwaukee; Edwin P.

Greaser, Ann Arbor, Michigan; Lellen Sterling Cheney, Barron, Wiscon¬

sin; H. C. Greene, Madison; Elmer W. Ellsworth, Stanford University,

California; Joe E. Morrison, Madison; G. M. W. Teyen, Milwaukee; Eliz¬

abeth Mac Donald, Oshkosh. It was moved and seconded and carried that

the secretary cast the unanimous ballot of the Academy for the new

members.

The treasurer’s report, as of April 9, 1931, was presented as follows:

Receipts

Balance in State Treasury, April 1, 1930 . $2006.95

State appropriation, July 1, 1930 . . . 1500.00

Dues received from members . 467.80

Wis. Geol. & Nat. Hist. Survey for plates . 429.90

Annual allowance from A. A. A. S . 116.00

From sale of Academy publications . 84.67

From members for extra reprints . . . 20.73

Total . $4626.05

Disbursements

Printing of Vol. 25 of Transactions . $2198.01

Other printing (programs, stationery, etc.) . 73.00

Zinc etchings for Vol. 26 of Transactions . 119.28

Secretary’s salary for the year . . . 200.00

Postage for correspondence and mailing Vol. 25 . 72.00

Supplies for secretary’s office . 8.68

Balance in State Treasury, April 9, 1931 . 1955.08

Total . . . . . $4626.05

Permanent Fund

Trust agreement, Centr. Wis. Trust Co . $1000.00

City of Madison bonds . 1000.00

Bonds, Chapman block, Madison . 400.00

Commonwealth Telephone Co. bonds . 400.00

Capitol Square Realty Co., Madison, bonds . 200.00

Cash on hand . 77.15

Total

$3077.15

568 Wisconsin Academy of Sciences, Arts, and Letters .

The auditing committee (Ira Edwards and C. A. Herrick) reported that

it had examined the treasurer’s accounts and securities and had found

them correct as stated in the report. The treasurer’s report and the au¬

diting committee’s report were approved.

It was moved, seconded and carried that the former secretary-treasurer,

Chancey Juday, be elected to life membership in the Academy in recogni¬

tion of his extended and valuable services during the nine years of his

secretaryship.

The following amendments to the constitution were adopted: (1)

MOVED, that Article III, Section 4, of the constitution be amended to

read as follows: “4. Active members shall be elected by the Academy or

by the council and shall enter upon membership on payment of the first

annual dues.” (2) MOVED, that the following sentence be stricken from

Article VIII of the constitution: “All members of the Academy shall re¬

ceive gratis the current issues of the Transactions.”

The codified by-laws of the Academy, as recommended by the council,

were then presented for action. L. J. Cole moved that the word “presi¬

dent” be stricken from Section 2. Carried. The remaining by-laws were

adopted as presented. These will be found printed in full elsewhere in this

issue of the Transactions.

The annual dinner of the Academy was held at the Grand View Hotel

on Friday evening, April 10, with 49 persons in attendance.

Lowell E. Noland,

Secretary-Treasurer.

THE CONSTITUTION OF THE WISCONSIN ACADEMY

OF SCIENCES, ARTS AND LETTERS

(January 1, 1932)

Article I — Name and Location

This association shall be known as the Wisconsin Academy of Sciences,

Arts and Letters, and shall be located at the city of Madison.

Article II — Object

The object of the Academy shall be the promotion of sciences, arts and

letters in the state of Wisconsin. Among the special objects shall be the

publication of the results of investigation and the formation of a library.

Article III — Membership

The Academy shall include four classes of members, viz.: life members,

honorary members, corresponding members and active members, to be

elected by ballot.

1. Life members shall be elected on account of special services rendered

the Academy. Life membership may also be obtained by the payment of

one hundred dollars and election by the Academy. Life members shall be

allowed to vote and to hold office.

2. Honorary members shall be elected by the Academy and shall be men

who have rendered conspicuous services to science, arts or letters.

3. Corresponding members shall be elected from those who have been

active members of the Academy, but who have removed from the state.

By special vote of the Academy men of attainments in science or letters

may be elected corresponding members. They shall have no vote in the

meetings of the Academy.

4. Active members shall be elected by the Academy or by the council,

and shall enter upon membership on payment of the first annual dues.

Article IV — Officers

The officers of the Academy shall be a president, a vice-president for

each of the three departments, sciences, arts and letters, a secretary, a li¬

brarian, a treasurer, and a custodian. These officers shall be chosen by

ballot, on recommendation of the committee on nomination of officers, by

the Academy at an annual meeting and shall hold office for three years.

Their duties shall be those usually performed by officers thus named in

scientific societies. It shall be one of the duties of the president to pre¬

pare an address which shall be delivered before the Academy at the an¬

nual meeting at which his term of office expires.

570 Wisconsin Academy of Sciences , Arts , and Letters .

Article V — Council

The council of the Academy shall be entrusted with the management of

its affairs during the intervals between regular meetings, and shall con¬

sist of the president, the three vice-presidents, the secretary, the treasur¬

er, the librarian, and the past presidents who retain their residence in

Wisconsin. Three members of the council shall constitute a quorum for

the transaction of business, provided the secretary and one of the presid¬

ing officers be included in the number.

Article VI — Committees

The standing committees of the Academy shall be a committee on publi¬

cation, a library committee, and a committee on the nomination of mem¬

bers. These committees shall be elected at the annual meeting of the

Academy in the same manner as the other officers of the Academy, and

shall hold office for the same term.

1. The committee on publication shall consist of the president and secre¬

tary and a third member elected by the Academy. They shall determine

the matter which shall be printed in the publications of the Academy.

They may at their discretion refer papers of a doubtful character to

specialists for their opinion as to scientific value and relevancy.

2. The library committee shall consist of five members, of which the li¬

brarian shall be ex officio chairman, and of which a majority shall not be

from the same city.

3. The committee on nomination of members shall consist of five mem¬

bers, one of whom shall be the secretary of the Academy.

Article VIII — Meetings

The annual meeting of the Academy shall be held at such time and place

as the council may designate; but all regular meetings for the election of

the board of officers shall be held at Madison. Summer field meetings shall

be held at such times and places as the Academy or the council may de¬

cide. Special meetings may be called by the council.

Article VIII — Publications

The regular publication of the Academy shall be known as its Transac¬

tions, and shall include suitable papers, a record of its proceedings, and

any other matter pertaining to the Academy. This shall be printed by the

state as provided in the statutes of Wisconsin.

Article IX — Amendments

Amendments to this constitution may be made at any annual meeting

by a vote of three-fourths of all the members present; provided , that the

amendment has been proposed by five members, and that notice has been

sent to all the members at least one month before the meeting.

Constitution and By-Laws .

571

BY-LAWS OF THE WISCONSIN ACADEMY OF

SCIENCES, ARTS AND LETTERS

1. The annual dues shall be two dollars for each active member, to be

charged to his account on the first day of January of each year. Five

dollars, paid in advance, shall constitute full payment for three years’

annual dues.

2. The annual dues shall be remitted for the secretary-treasurer and

librarian during their term of office.

3. As soon as possible after January first of each year the secretary-

treasurer shall send to members statements of dues payable, and in case

of non-payment shall, within the succeeding four months, send a second

and, if necessary, a third notice.

4. The secretary-treasurer shall strike from the list of members the

names of those who are one year or more in arrears in the payment of

their dues, and shall notify such members of this action, offering at the

same time to reinstate them upon receipt of the dues in arrears plus the

dues for the current year.

5. Each member of the Academy shall receive the current issue of the

Transactions, provided that his dues are paid. Any member in arrears at

the time the Transactions are published shall receive his copy as soon as

his dues are paid.

6. The fees received from life members shall be set apart as a per¬

manent endowment fund, to be invested exclusively in securities which are

legal as investments for Wisconsin trust companies or savings banks. The

income alone from such fund may be used for the general purposes of the

Academy.

7. The secretary-treasurer shall receive annually an allowance of two

hundred dollars for services.

8. The secretary-treasurer shall be charged with the special duty of

editing and overseeing the publication of the Transactions. In the per¬

formance of this duty he shall be advised by the committee on publication.

9. The Transactions shall contain in each volume: (a) a list of the offi¬

cers of the Academy, (b) the minutes of the annual meeting, and (c) such

papers as are accepted under the provisions of Section 10 of these By-

Laws and no others.

10. Papers to be published in the Transactions must be approved as to

content and form by the committee on publication. They must represent

genuine, original contributions to the knowledge of the subject discussed.

Preference shall be given to papers of special interest to the State of Wis¬

consin, and to papers presented at a regular meeting of the Academy. The

privilege of publishing in the Transactions shall be reserved for the mem¬

bers of the Academy.

572 Wisconsin Academy of Sciences , Arts , and Letters .

11. The Constitution and By-Laws and the names and addresses of the

members of the Academy shall be published every third year in the Tran¬

sactions. The Constitution and By-Laws shall also be available in reprint

form from the secretary-treasurer at any time.

12. Amendments to these By-Laws may be made at any annual meeting

by vote of three-fourths of all the members present.

SUBJECT AND AUTHOR INDEX TO THE PAPERS

PUBLISHED BY THE ACADEMY,

1870-1932

Compiled by

Lowell E. Noland,

Secretary

Subject Index

The figures refer to the number of the article in the alphabetized author index

which follows the subject index.

Addresses of a general nature: 135, 339, 587.

American Indians: 20, 67, 79, 175, 209, 290, 294, 295, 342-344, 484, 525,

528, 664, 679.

Archeology: 627 (see also American Indians).

Art: 511, 612.

Astronomy: 141, 216, 437, 553, 554, 597.

Bacteriology: 26, 225, 226, 232, 315, 601.

Biography: 18, 25, 60, 77a, 82, 86-88, 97, 98, 110, 114, 119, 158, 192, 202,

204, 217, 222, 228, 236, 245, 252, 264, 2 66, 289, 299, 371, 375, 376, 393,

401, 404, 443, 444, 458, 507, 508, 534, 537, 583, 584, 589, 605, 622, 633,

665, 681.

Biology: (see Botany and Zoology).

Botany: 664; Algae: 492, 568, 591-595, 623; Angiosperms: 19, 68, 115,

148, 179, 205, 206, 237, 241, 258, 387, 392, 397, 398, 402, 496, 544, 567,

609, 652, 655, 667; Bacteria (see under Bacteriology) ; Bryophytes: 4,

34, 130, 132, 143, 144; Cytology: 6, 28, 145, 146, 176, 178, 224, 251,

269, 278, 396, 397, 480, 493, 496, 557; Ecology: 231, 352, 361, 609;

Embryology : 369, 397; Flora of Wisconsin: 19, 65, 68, 128-132, 143,

144, 148, 167, 170, 172, 177, 179, 181, 182, 194, 205, 206, 229, 241,

246-250, 256, 387, 391, 398, 402, 421, 593, 594, 652, 674; Flora of

North America: 168, 171, 355, 595, 607; Fungi: 7, 28, 131, 133, 134,

145, 146, 167, 168, 170-172, 174, 177, 181, 182, 224, 229, 230, 232, 233,

246-251, 256, 269, 278, 316, 341, 389, 390, 394, 396, 436, 480, 493, 540,

541, 557, 628, 629, 659, 667, 697; Gymnosperms: 206, 350, 355, 563;

Palaeobotany : 675; Physiology : 147, 176, 178, 224, 237, 258, 278, 563,

655; Geography and Distribution (see Flora); Pteridophytes : 6, 65,

194, 406, 652, 674; Taxonomy in general: 366, 367.

Chemistry: 260, 658; Analysis: 58, 547-549, 590, 624; Inorganic: 188,

311, 312, 337, 364, 377, 378, 458, 509, 549, 579; Organic: 151, 255, 260,

312, 357, 358, 547, 614; Physical: 338, 347, 386, 579.

Economics: 35, 69, 100, 102, 123, 124, 185, 239, 287, 318, 319, 383, 448, 449,

506, 556, 572, 573, 582, 585, 598, 606, 618, 635, 691.

Education: 4, 221, 272, 284, 309, 395, 495, 504, 511, 539, 552, 664.

Engineering: Civil: 482, 599; Hydraulic: 475, 483; Materials: 24, 301,

472, 476; Methods and instruments: 472, 116; Surveying: 473;

Stresses: 279, 346, 476, 488; Waterways: 474, 475, 481.

574 Wisconsin Academy of Sciences, Arts, and Letters.

English (see Etymology, Linguistics, Literature).

Etymology: 33, 89, 90, 211, 372, 626.

Forestry and Wood Technology: 24, 200, 350.

Geodesy and Earth Magnetism: 388, 473.

Geography: Africa: 27, 127; North America: 152; Wisconsin: 333, 870,

465.

Geology: 64, 111, 265, 639; Africa: 27; Europe: 112; North America:

139, 152, 153, 154, 261, 262, 304, 380, 466; Wisconsin: 64, 74-76, 104,

107-109, 160, 189-191, 193, 195, 196, 262, 301-306, 320, 353, 356, 365,

379, 438, 562, 602, 613, 615, 620, 634, 638, 684, 686-689.

History: 13, 17; American: 11, 42, 71, 78, 80, 85, 118, 215, 382, 384, 478,

479, 510, 535, 572, 573; English: 8, 9, 14, 16, 319; European: 10, 12,

15, 240, 504, 543, 643-651; Wisconsin: 80, 127, 138, 203, 239, 342, 343,

344, 349, 596, 608, 672.

Home Economics: 26, 85, 570.

Languages: American Indian: 210, 372; German: 643-651; Gothic: 33;

Greek: 39, 242; Italian: 561; Latin: 242; Old English : 33.

Law: 101, 611.

Limnology: Apparatus: 51, 329, 331; Chemical studies: 58, 324, 825, 334,

335, 547, 548, 549, 590, 624, 631 ; Fauna of Lakes : 282, 286, 287, 324, 830,

407, 411, 461, 469, 653, 696; Flora of Lakes: 179, 492, 544, 567, 568;

Hydrography and General Studies of Lakes: 127, 323, 328, 368, 408,

515, 676; Light penetration: 53-56, 533, 578; Plankton: 45, 45a, 322,

411, 413, 492, 566, 593-595, 601, 630, 631, 676, 683; Stations: 326,

333; Temperature Studies: 47-50, 52, 408, 515, 631, 676; Tides: 475.

Linguistics, Comparative Grammar, Syntax: 212, 497-503.

Literature: American: 137, 608; American Indian: 20; English: 2-4, 38,

81, 91, 136, 156, 359, 360, 530, 532, 558, 656, 661, 662, 698-701;

French: 662; German: 373, 551; Greek: 504, 636; Italian: 84;

Latin: 504.

Mathematics : 66, 116, 140, 308, 529, 552, 586, 702.

Medical science: 159, 456, 539, 604.

Meteorology: 36, 50, 72, 73, 351, 450-454.

Mineralogy: 63, 153, 214, 262, 305, 306, 354, 555, 562, 637, 684.

Mining and Metallurgy: 27, 214, 354, 383, 467, 468.

Music: 213.

Obituary (see Biography) .

Pharmacy: 186, 357.

Philosophy: 37, 61, 95, 96, 117, 197, 198, 199, 270, 274, 300, 446, 559, 577,

581, 690, 702.

Physics: 59, 116, 162-166, 400, 682.

Physiology and Biochemistry : 159, 456, 463, 569, 604.

Plant Pathology : 321, 540, 541, 657.

Political Science: 11, 15, 16, 35, 69, 70, 99, 101, 180, 234, 235, 273, 296, 297,

298, 318, 349, 382.

Psychology: 62, 309, 310.

Religion : 20, 22, 543, 528.

Sociology: 8, 9, 10, 12, 22, 99, 185, 212, 240, 257, 271, 298, 348, 383, 439,

600, 618, 678, 692, 693.

Index to Academy Publications, 1870-1932 .

575

Travel notes: 152, 263, 326.

Zoology: 664; General: Ecology: 29, 231, 293 (see also Limnology);

Embryology : 369, 433, 434; Evolution: 96, 313, 491; Longevity: 120,

121, 122; Neurology: 44; Parasitology : 40, 440-442, 513, 514, 575;

Systematic: Protozoa: 174, 233, 324, 440, 441, 442, 493; Rotifera: 254,

470; Parasitic Worms: 513, 514; Annelida: 41, 324; Mollusca : 29, 31,

32, 324, 460, 461, 462; Crustacea: 43, 45a, 77, 150, 275, 276, 322, 407,

411, 429, Copepoda: 40, 46, 327, 374, 409, 410, 412, 414-418; Hydra-

carina (water mites) : 419, 420, 422-428; Araneida (spiders) : 477,

516-523; Fishes: 40, 286, 314, 368, 440-442, 463, 513, 514, 653, 696;

Amphibia: 44, 259, 292, 536; Reptilia: 23, 259, 313, 536; Birds: 142,

238, 287, 307, 403, 489, 564, 671; Mammals: 281, 291, 293, 456, 457,

569, 663; Zoogeographic: Africa: 521; Asia: 425, 522; North Ameri¬

ca: 105, 142, 201, 414-416, 418, 424, 517, 523; Wisconsin: 29, 30, 32,

105, 150, 218, 238, 254, 259, 280-282, 285, 307, 403, 407, 409, 411, 427,

428, 460, 461, 469, 536, 564, 671.

List of Articles

Published in Bulletins 1-5 (1870-1871) and in the

Transactions , Vols. 1-27 (1872-1932)

1. Alder, Hugo, joint author; see H. A. Schuette.

2. Alderman, William E. 1924. Bibliograpical evidence of the vogue of

Shaftesbury in the eighteenth century. Trans. 21 : 57-70.

3. Alderman, William E. 1931. Shaftesbury and the doctrine of be¬

nevolence in the eighteenth century. Trans. 26 : 137-159.

4. Ainsworth, Oliver M. 1924. Milton as a writer on education. Trans.

21 : 41-50.

5. Allen, Charles Elmer. 1904. Some Hepaticae of the Apostle Islands.

Trans. 14 (2) : 485-486.

6. Allen, Ruth Florence. 1914. Studies in spermatogenesis and apo-

gamy in ferns. Trans. 17 (1) : 1-56.

7. Allen, Ruth Florence, and Hally D. M. Jolivette. 1914. A study of

the light reactions of Pilobolus. Trans. 17 (1) : 533-598. Figs.

8. Allen, William Francis. 1872. The rural population of England, as

classified in the Domesday book. Trans. 1 : 167-177.

9. Allen, William Francis. 1874. The rural classes of England in the

thirteenth century. Trans. 2 : 220-233.

10. Allen, William Francis. 1874. Ranks and classes among the Anglo-

Saxons. Trans. 2 : 234-240.

11. Allen, William Francis. 1876. United States sovereignty — whence

derived, and where vested. Trans. 3 : 125-132.

12. Allen, William Francis. 1879. Peasant communities in France.

Trans. 4 : 1-6.

13. Allen, William Francis. 1879. The origin of the freeholders.

Trans. 4 : 19-24.

14. Allen, William Francis. 1882. The English cottagers of the middle

ages. Trans. 5 : 1-11.

576 Wisconsin Academy of Sciences , Arts , and Letters .

15. Allen, William Francis. 1885. The primitive democracy of the Ger¬

mans. Trans. 6 : 28-42.

16. Allen, William Francis. 1889. The village community and serfdom

in England. Trans. 7 : 130-140.

17. Allen, William Francis. 1889. Town, township and tithing. Trans.

7 : 141-154.

18. Allen, William Francis. 1889. 0. M. Conover. Trans. 7 : 257-258.

19. Almon, Lois. 1930. Preliminary reports on the flora of Wisconsin.

XI. Ranunculaceae. Trans. 25 : 205-214.

20. Andrews, Edmund. 1879. Discoveries illustrating the literature

and religion of the mound builders. Trans. 4 : 126-131.

21. Andrews, Joy Ella. 1921. Some experiments with the larva of the

bee-moth, Galleria mellonella. Trans. 20 : 255-261.

22. Armitage, William Edmond. 1872. The German Sunday. Trans.

1 : 62-71.

23. Atwood, William Henry. 1918. The visceral anatomy of the garter

snake. Trans. 19 (1) : 531-552. With figs.

24. Austin, Louis Winslow, and C. W. Eastman. 1902. On the relation

between heat conductivity and density in some of the common woods.

Trans. 13 (2) : 539-542.

25. Axtell, Luta. 1900. Wayland Samuel Axtell. Trans. 12 (2) :

560-561.

26. Bachmann, Freda Marie. 1927. A bacteriological study of salad

dressings. Trans. 23 : 529-537.

27. Bagg, Rufus Mather. 1930. The diamond mining industry of South

Africa. Trans. 25 : 79-87.

28. Baird, Edgar Alan. 1924. The structure and behavior of the nu¬

cleus in the life history of Phy corny ces nitens (Agardh) Kunze and Rhizo-

pus nigricans Ehrbg. Trans. 21 : 357-380. 2 pis.

29. Baker, Frank Collins. 1914. The molluscan fauna of Tomahawk

Lake, Wisconsin. With special reference to its ecology. Trans. 17 (1) :

200-246. 7 pis.

30. Baker, Frank Collins. 1924. The fauna of the Lake Winnebago re¬

gion; a quantitative and qualitative survey with special reference to the

Mollusca. Trans. 21 : 109-146.

31. Baker, Frank Collins. 1926. Nomenclatorial notes on American

fresh water Mollusca. Trans. 22 : 193-205.

32. Baker, Frank Collins. 1928. The fresh water Mollusca of Wiscon¬

sin. (This is a monograph published in two separate volumes. Part I

Gastropoda. 507 pp. 28 plates. 202 text figures. Part II. Pelecypoda. 495

pp. 77 plates. 299 text figures.) Not published in the regular Transac¬

tions, but for sale by the Academy as a separate monograph.

33. Balg, Gerhard Hubert. 1892. The relation of Old English ‘reomig’

to Gothic ‘rimis\ Trans. 8 : 167-168.

34. Barnes, Charles Reid. 1892. Artificial keys to the genera and spe¬

cies of mosses recognized in Lesquereux and James’s Manual of the

Mosses of North America. Trans. 8 : 11-81. Additions and corrections.

Trans. 8 : 163-166.

Index to Academy Publications , 1870-1932 .

577

35. Barnett, James Duff. 1905. The state administration of taxation

in Wisconsin. Trans. 15 (1) : 163-177.

36. Bartlett, James Lowell. 1909. The cold-waves of south-central Wis¬

consin. Trans. 16 (1) : 289-306. 7 pis.

37. Bascom, John. 1885. Freedom of will empirically considered.

Trans. 6 : 2-20.

38. Beatty, Arthur. 1907. The St. George, or Mummers’, plays : a study

in the protology of the drama. Trans. 15 (2) : 273-324.

39. Bennett, Charles Edwin. 1892. Some new theories of the Greek

ka-perfect. Trans. 8 : 141-155.

40. Bere, Ruby. 1931. Copepods parasitic on fish of the Trout Lake

region, with descriptions of two new species. Trans. 26 : 427-436.

41. Bere, Ruby. 1931. Leeches from the lakes of northeastern Wiscon¬

sin. Trans. 26 : 437-440. 2 pis.

42. Bille, John H. 1898. A history of the Danes in America. Trans.

11 : 1-48. 1 pi.

43. Birge, Edward Asahel. 1879. Notes on Cladocera. I. Trans. 4 : 77-

112. II. Trans. 8 : 379-398 (1892). III. Trans. 9 : 275-317 (1893). IV.

Trans. 16 (2) : 1017-1066. Plates.

44. Birge, Edward Asahel, 1885. On the motor ganglion cells of the

frog’s spinal cord. Trans. 6 : 51-81. 1 pi.

45. Birge, Edward Asahel, O. A. Olson and H. P. Harder. 1895. Plank¬

ton studies on Lake Mendota. Trans. 10 : 421-484. Plates.

45a. Birge, Edward Asahel. 1898. Plankton studies on Lake Mendota.

II. The Crustacea of the plankton from July, 1894, to December, 1896.

Trans. 11 : 274-448.

46. Birge, Edward Asahel, and C. Juday. 1909. A summer resting

stage in the development of Cyclops bicuspidatus Claus. Trans. 16 (1) :

1-9.

47. Birge, Edward Asahel. 1910. An unregarded factor in lake tem¬

peratures. Trans. 16 (2) : 989-1004. 2 pis.

48. Birge, Edward Asahel. 1910. On the evidence for temperature

seiches. Trans. 16 (2) : 1005-1016. 1 pi.

49. Birge, Edward Asahel. 1916. The heat budgets of American and

European lakes. Trans. 18 (1) : 166-213. Figs.

50. Birge, Edward Asahel. 1916. The work of the wind in warming a

lake. Trans. 18 (2) : 341-391. Figs.

51. Birge, Edward Asahel. 1921. A second report on limnological ap¬

paratus. Trans. 20 : 533-552. Figs., 2 pis. (First report, Juday 1916)

52. Birge, Edward Asahel, C. Juday ,and H. W. March. 1927. The

temperature of the bottom deposits of Lake Mendota; a chapter in the

heat exchanges of the lake. Trans. 23 : 187-231. Figs.

53. Birge, Edward Asahel, and C. Juday. 1929. Transmission of solar

radiation by the waters of inland lakes. Trans. 24 : 509-580. Figs.

54. Birge, Edward Asahel, and C. Juday. 1930. A second report on

solar radiation and inland lakes. Trans. 25 : 285-335. Figs.

55. Birge, Edward Asahel, and C. Juday. 1931. A third report of solar

radiation and inland lakes. Trans. 26 : 383-425. Figs.

578 Wisconsin Academy of Sciences, Arts, and Letters .

56. Birge, Edward Asahel, and C. Juday. 1932. Solar radiation and

inland lakes. Fourth report. Observations of 1931. Trans. 27 : 000-000.

Figs.

57. Birge, E. A., joint author; see Chancey Juday.

58. Black, Charles Spurgeon. 1929. Chemical analyses of lake de¬

posits. Trans. 24 : 127-133.

59. Blackstone, Dodge Pierce. 1885. The variation in attraction due to

the figure of the attracting bodies. Trans. 6 : 197-254.

60. Blaisdell, James Joshua. 1893. Aaron Lucius Chapin. Trans. 9 :

lxxi-lxxiv. Portrait.

61. Blaisdell, James Joshua. 1898. The methods of science, as being in

the domain of logic. Trans. 11 : 49-65.

62. Blaisdell, James Joshua. 1893. Some suggestions concerning meth¬

ods of psychological study. Trans. 9 : 33-43.

63. Blake, William Phipps. 1903. Arizona diatomite. Trans. 14 (1) :

107-111. 2 pis.

64. Blake, William Phipps. 1893. The process of geological surveys

in the state of Wisconsin — a review and bibliography. Trans. 9 : 225-231.

65. Breakey, Edith W., and Ruth I. Walker. 1931. Preliminary reports

on the flora of Wisconsin. XII. Polypodiaceae. Trans. 26 : 263-273. Figs.

66. Bremiker, Karl. 1902. On the errors with which logarithmic com¬

putations are affected. (Translated by Pearl Eugene Doudna and Elwyn

Francis Chandler). Trans. 13 (2) : 427-474.

67. Brown, Charles Edward. 1907. Wisconsin quartzite implements.

Trans. 15 (2) : 656-663. 2 pis.

68. Brues, Charles Thomas, and Beirne B. Brues. 1914. The grasses of

Milwaukee County, Wisconsin. Trans. 17 (1) : 57-76. 3 pis.

69. Bruncken, Ernest. 1898. On some differences between private and

public business. Trans. 12 (1) : 325-336.

70. Bruncken, Ernest. 1898. The use of parties in municipal govern¬

ment. Trans. 11 : 225-235.

71. Buck, Solon Justus. 1907. The settlement of Oklahoma. Trans.

15 (2) : 325-380. 6 pis.

72. Buckley, Ernest Robertson. 1901. Ice ramparts. Trans. 13 (1) :

141-162. 18 pis.

73. Buckley, Ernest Robertson. 1909. Sleet storm in the Ozark region

of Missouri. Trans. 16 (1) : 307-310. 7 pis.

74. Buell, Ira Maynard. 1882. The corals of Delafield. Trans. 5 :

185-193.

75. Buell, Ira Maynard. 1893. Geology of the Waterloo quartzite area.

Trans. 9 : 255-274. 3 pis.

76. Buell, Ira Maynard. 1895. Bowlder trains from the outcrops of

the Waterloo quartzite areas. Trans. 10 : 485-509. 5 pis.

77. Bundy, Will F. 1882. A list of the Crustacea of Wisconsin. With

notes on some new or little known species. Trans. 5 : 177-184.

77a. Burd, Henry A. 1918. Eight unedited letters of Joseph Ritson.

Trans. 19 (1) : 1-21.

78. Butler, James Davie. 1874. The naming of America. Trans. 2 :

203-219.

Index to Academy Publications , 1870-1932 .

579

79. Butler, James Davie. 1876. Copper tools found in the state of

Wisconsin. Trans. 3 : 99-104.

80. Butler, James Davie. 1882. First French foot-prints beyond the

Lakes; or, What brought the French so early into the Northwest?

Trans. 5 : 85-145.

81. Butler, James Davie. 1882. The Xeya/ieva in Shakespere.

Trans. 5 : 161-174.

82. Butler, James Davie. 1892. William Francis Allen. Trans. 8 :

439-441. Portrait.

83. Butler, James Davie. 1895. Phases of witticism. Trans. 10 : 41-60.

84. Butler, James Davie. 1898. Dante. His quotations and his orig¬

inality; the greatest imitator and the greatest original. Trans. 11 : 149-

164.

85. Butler, James Davie. 1898. Codfish. Its place in American history.

Trans. 11 : 261-273.

86. Butler, James Davie. 1898. George P. Delaplaine. Trans. 11 :

522-524.

87. Butler, James Davie. 1898. Simeon Mills. Trans. 11 : 524-525.

88. Butler, James Davie. 1900. Harlow S. Orton, 1817-1895. Trans.

12 (2) : 554-555.

89. Butler, James Davie. 1901. Household words: their etymology.

Trans. 13 (1) : 366-383.

90. Butler, James Davie. 1902. Personal names: their significance and

historical origin. Trans. 13 (2) : 475-485.

91. Butler, James Davie. 1903. The vocabulary of Shakespeare. Trans.

14 (1) : 40-55.

92. Cahn, Alvin Robert. 1918. Notes on the vertebrate fauna of

Houghton and Iron counties, Michigan. Trans. 19 (1) : 483-510. 5 pis.

93. Campbell, Mrs. Katharine; see Katharine Bernice Stewart.

94. Carlton, E. P., joint author; see under W. S. Miller.

95. Carpenter, Stephen Haskins. 1874. The metaphysical basis of sci¬

ence. Trans. 2 : 23-34.

96. Carpenter, Stephen Haskins. 1874. The philosophy of evolution.

Trans. 2 : 39-58.

97. Carpenter, Stephen Haskins. 1879. John Baptist Feuling. Trans.

4 : 316-318.

98. Caverno, Charles. 1871. Abstract of a paper on the relations be¬

tween social and moral science. Bull. 4 : 59-61.

99. Caverno, Charles. 1872. Social science and woman suffrage.

Trans. 1 : 72-89.

100. Caverno, Charles. 1876. The people and the railroads. Trans. 3 :

143-150.

101. Caverno, Charles. 1879. The abolition of the jury system. Trans.

4 : 7-18.

102. Caverno, Charles. 1882. Life insurance, savings banks, and the

industrial situation. Trans. 5 : 21-37.

103. Chamberlin, Thomas C. 1871. Suggestions as to the basis for the

gradation of the vertebrata. (An abstract). Bull. 5 : 76-80.

580 Wisconsin Academy of Sciences, Arts, and Letters .

104. Chamberlin, Thomas C. 1871. Facts relating to the local geology

of the Whitewater region. (An abstract). Bull. 5 : 81.

105. Chamberlin, Thomas C. 1871. On the 17-year cicada; its geo¬

graphical distribution and time of appearance in this state. (Title only).

Bull. 5 : 81.

106. Chamberlin, Thomas Crowder. 1872. Suggestions as to a basis for

the gradation of the Vertebrata. Trans. 1 : 138-150.

107. Chamberlin, Thomas Crowder. 1874. Some evidences bearing upon

the method of the upheaval of the quartzites of Sauk and Columbia coun¬

ties. Trans. 2 : 129-132. Figs.

108. Chamberlin, Thomas Crowder. 1874. On fluctuations in level of

the quartzites of Sauk and Columbia counties. Trans. 2 : 133-138. Figs.

109. Chamberlin, Thomas Crowder. 1879. On the extent and signifi¬

cance of the Wisconsin kettle moraine. Trans. 4 : 201-234. 2 pis.

110. Chamberlin, Thomas Crowder. 1879. In memoriam. Prof. James

H. Eaton. Trans. 4 : 314-316.

111. Chamberlin, Thomas Crowder. 1882. On a proposed system of

lithological nomenclature. Trans. 5 : 234-247.

112. Chamberlin, Thomas Crowder. 1882. Observations on the recent

glacial drift of the Alps. Trans. 5 : 258-270.

113. Chamberlin, Thomas Crowder. 1892. Some additional evidences

bearing on the interval between the glacial epochs. Trans. 8 : 82-86.

114. Chamberlin, Thomas Crowder. 1892. Roland Duer Irving. Trans.

8 : 433-437. Portrait.

114a. Chamberlin, Thomas C. 1921. The founding of the Wisconsin

Academy of Sciences, Arts and Letters. Trans. 20 : 693-701.

115. Chandler, Charles Henry. 1892. Notes and a query concerning

the Ericaceae. Trans. 8 : 161-162.

116. Chandler, Charles Henry. 1895. An improved harmonograph.

Trans. 10 : 61-63.

117. Chandler, Charles Henry. 1898. Transcendental space. Trans. 11

: 239-248.

118. Chandler, Charles Henry. 1898. An historical note on early Ameri¬

can railways. Trans. 12 (1) : 317-324.

119. Chandler, Charles Henry. 1898. Newton Stone Fuller, Trans. 11 :

526.

120. Chandler, Charles Henry. 1900. The inter-generation period.

Trans. 12 (2) : 499-504. 2 pis.

121. Chandler, Charles Henry. 1901. A problem of longevity. Trans.

13 (1) : 384-386.

122. Chandler, Charles Henry. 1903. A study in longevity. Trans.

14 (1) : 56-62.

122a. Chandler, E. F., translator, see Karl Bremiker.

123. Chapin, Aaron Lucius. 1872. The relations of labor and capital.

Trans. 1 : 45-61.

124. Chapin, Aaron Lucius. 1882. The nature and functions of credit.

Trans. 5 : 57-65.

Index to Academy Publications , 1870-1932.

581

125. Chase, Ruth Wayland. 1921. The length of the life of the larva

of the wax moth, Galleria mellonella L., in its different stadia. Trans. 20 :

263-267.

126. Chase, Ruth W.; see also under Ruth Chase Noland.

127. Chase, Wayland Johnson, and Lowell E. Noland. 1927. The his¬

tory and hydrography of Lake Ripley (Jefferson County, Wisconsin).

Trans. 23 : 179-186. Map.

128. Cheney, Lellen Sterling, and R. H. True. 1893. On the flora of

Madison and vicinity, a preliminary paper on the flora of Dane County,

Wisconsin. Trans. 9 : 45-135. 1 pi.

129. Cheney, Lellen Sterling. 1893. A contribution to the flora of the

Lake Superior region. Trans. 9 : 233-254.

130. Cheney, Lellen Sterling. 1895. Sphagna of the upper Wisconsin

Valley. Trans. 10 : 66-68.

131. Cheney, Lellen Sterling. 1895. Parasitic Fungi of the Wisconsin

Valley. Trans. 10 : 69.

132. Cheney, Lellen Sterling. 1895. Hepaticae of the Wisconsin Val¬

ley. Trans. 10 : 70-72.

133. Christman, Arthur Henry. 1905. Observations on the wintering

of grain rusts. Trans. 15 (1) : 98-107. 1 pi.

134. Christman, Arthur Henry. 1907. The nature and development of

the primary uredospore. Trans. 15 (2) : 517-526.

135. Clancy, George C. 1924. Floundering in modernity. Trans. 21 :

51-55.

136. Clark, Harry Hayden. 1929. The romanticism of Edward Young.

Trans. 24 : 1-45.

137. Clark, Harry Hayden. 1930. What made Freneau the father of

American prose? Trans. 25 : 39-50.

138. Cole, Harry Ellsworth. 1926. Stagecoach and tavern days in the

Baraboo region. Trans. 22 : 1-8.

139. Collie, George Lucius. 1895. The geology of Conanicut Island, R.

I. Trans. 10 : 199-230. 1 pi.

140. Comstock, Eltin Houghtaling. 1898. The real singularities of har¬

monic curves of three frequencies. Trans. 11 : 452-464.

141. Comstock, George Carey. 1892. The present condition of the lati¬

tude problem. Trans. 8 : 229-232.

142. Congdon, Russel Thompson. 1904. Saskatchewan birds. Trans. 14

(2) : 569-620. 8 pis.

143. Conklin, George H. 1914. Preliminary report on a collection of

Hepaticae from the Duluth-Superior district. States of Minnesota and

Wisconsin. Trans. 17 (2) : 985-1010.

144. Conklin, George Hall. 1929. The Hepaticae of Wisconsin. Trans.

24 : 197-247. Map.

145. Cooper, George Olds. 1929. A cytological study of fertilization in

Achlya hypogyna Coker and Pemberton. Trans. 24 : 303-308. 1 pi.

146. Cooper, George Olds. 1929. Cytological studies on the sporange

development and gametogenesis in Brevilegnia declina Harvey. Trans.

24 : 309-322. 3 pis.

582 Wisconsin Academy of Sciences, Arts, and Letters .

147. Copeland, Edwin Bingham and Louis Kahlenberg. 1900. The in¬

fluence of the presence of pure metals upon plants. Trans. 12 (2) : 454-

474.

148. Costello, David P. 1931. Preliminary reports on the flora of Wis¬

consin. XIII. Fagaceae. Trans. 26 : 275-279. Figs.

149. Crathorne, Mrs. Charlotte E.; see Charlotte Elvira Pengra.

150. Creaser, Edwin P. 1932. The Decapod Crustaceans of Wisconsin.

Trans. 27 : 000-000. Figs.

151. Crooker, Orin Edson. 1898. Aluminum alcoholates. Trans. 11 :

255-260.

152. Culver, Garry Eugene. 1892. Notes on a little known region in

northwestern Montana. Trans. 8 : 187-205. Figs., map.

153. Culver, Garry Eugene and Wm. H. Hobbs. 1892. On a new occur¬

rence of olivine diabase in Minnehaha County, South Dakota. Trans. 8 :

206-210.

154. Culver, Garry Eugene. 1895. Some New Jersey eskers. Trans. 10 :

19-23.

155. Culver, Garry Eugene. 1895. The erosive action of ice. Trans.

10 : 339-366.

156. Cunliffe, John William. 1914. Browning’s idealism. Trans. 17

(1) : 661-681.

157. Curran, C. Howard. 1926. Revision of the Nearctic species of

Helophilus and allied genera. Trans. 22 : 207-281. 3 pis.

158. Curtis, Wardon Allan. 1907. David Bower Frankenburger. Trans.

15 (2) : 912-915. Portrait.

159. Daniells, William Willard. 1874. Note on the rapidity of the ab¬

sorption of arsenic by the human liver. Trans. 2 : 128.

160. Daniells, William Willard. 1898. On the analysis of the water of

a flowing artesian well at Marinette, Wisconsin. Trans. 11 : 112-113.

161. Davies, John Eugene. 1871. Abstract of a paper on the impor¬

tance and practicability of finding a unit of force in physics that shall be

of universal application. Bull. 2 : 34.

162. Davies, John Eugene. 1871. On the kinetic measures of force.

(Title only). Bull. 4 : 67.

163. Davies, John Eugene. 1872. On potentials and their application

in physical science. Trans. 1 : 155-164.

164. Davies, John Eugene. 1876. Recent progress in theoretical phys¬

ics. Trans. 3 : 205-221.

165. Davies, John Eugene. 1879. Recent progress in theoretical phys¬

ics. Part II. Magnetic rotatory polarization of light. Trans. 4 : 241-264.

166. Davies, John Eugene. 1893. On some analogies between the equa¬

tions of elasticity and electro-magnetism. Trans. 9 : 1-20.

167. Davis, John Jefferson. 1893-1910. A supplementary list of para¬

sitic Fungi of Wisconsin. I. Trans. 9 : 153-188 (1893). II. Trans. 11 :

165-178 (1898). III. Trans. 14 : 83-106 (1903). IV. Trans. 16 : 739-776

(1910).

168. Davis, John Jefferson. 1907. Mycological narrative of a brief

journey through the Pacific northwest. Trans. 15 (2) : 775-780.

Index to Academy Publications, 1870-1932.

583

169. Davis, John Jefferson. 1907. The Academy: its past and future.

Trans. 15 (2) : 887-896.

170. Davis, John Jefferson. 1914. A provisional list of parasitic Fungi

in Wisconsin. Trans. 17 (2) : 846-984.

171. Davis, John Jefferson. 1919. North American Ascochytae, a de¬

scriptive list of species. Trans. 19 (2) : 655-670.

172. Davis, John Jefferson. 1916-1932. Notes on parasitic Fungi in

Wisconsin. I. Trans. 18 (1) : 78-92 (1916). II. Trans. 18 (1) : 93-109

(1916). III. Trans. 18 (1) : 251-271 (1916). IV. Trans. 19 (2) : 671-689

(1919). V. Trans. 19 (2) : 690-704 (1919). VI. Trans. 19 (2) : 705-727

(1919). VII. Trans. 20 : 399-411 (1921), figs., 2 pis. VIII. Trans. 20 :

413-431e (1921), figs. IX. Trans. 21 : 251-269 (1924), figs. X. Trans.

21 : 271-286 (1924), figs. XI. Trans. 21 : 287-302e (1924), figs. XII.

Trans. 22 : 155-163 (1926). XIII. Trans. 22 : 165-179 (1926), figs.

XIV. Trans. 22 : 181-192 (1926). XV. Trans. 24 : 269-277 (1929). XVI.

Trans. 24 : 279-293 (1929), fig. XVII. Trans. 24 : 295-302c (1929).

XVIII. Trans. 26 : 253-261 (1931). XIX. Trans. 27 : 000-000 (1932).

173. Day, Fish Holbrook. 1879. On the fauna of the Niagara and upper

Silurian rocks as exhibited in Milwaukee County, Wisconsin, and in coun¬

ties contiguous thereto. Trans. 4 : 113-125.

174. Dean, Alletta Friscone. 1914. The Myxomycetes of Wisconsin.

Trans. 17 (2) : 1221-1299.

175. De Hart, J. N. 1879. The antiquities and platycnemism of the

mound builders of Wisconsin. Trans. 4 : 188-200. Figs., 1 pi.

176. Denniston, Rollin Henry. 1904. The structure of the starch grain.

Trans. 14 (2) : 527-533.

177. Denniston, Rollin Henry. 1905. The Russulas of Madison and vi¬

cinity. Trans. 15 (1) : 71-88.

178. Denniston, Rollin Henry. 1907. The growth and organization of

the starch grain. Trans. 15 (2) : 664-708. 3 pis.

179. Denniston, Rollin Henry. 1921. A survey of the larger aquatic

plants of Lake Mendota. Trans. 20 : 495-500.

180. Desmond, Humphrey Joseph. 1892. The sectional feature in

American politics. Trans. 8 : 1-10.

181. Dodge, Bernard Ogilvie. 1914. A list of Fungi, chiefly sapro¬

phytes, from the region of Kewaunee County, Wisconsin. Trans. 17 (2) :

806-845.

182. Dodge, Bernard Ogilvie. 1914. Wisconsin Discomycetes. Trans.

17 (2) : 1027-1056.

183. Domogalla, Bernard P., joint author; see W. L. Tressler.

184. Doudna, Pearl Eugene, translator; see Karl Bremiker.

185. Downes, Robert Hugh, and Katherine Patricia Regan. 1902. Eco¬

nomic and social development of Kenosha and La Fayette counties. (With

an introduction by Orin Grant Libby). Trans. 13 (2) : 543-609.

186. Du Mez, Andrew Grover. 1919. A century of the United States

pharmacopoeia, 1820-1920. I. The galenical oleoresins. Trans. 19 (2) ;

907-1194.

584 Wisconsin Academy of Sciences, Arts, and Letters .

187. Eastman, C. W., joint author; see L. W. Austin.

188. Eaton, James H. 1871. On the formation of certain new com¬

pounds of manganese. (An abstract). Bull. 2 : 35-36.

189. Eaton, James H. 1871. On the geology of the region about Devil’s

Lake, Sauk County, Wisconsin; being a report of observations made at the

request of the Academy. (An abstract). Bull. 4 : 55-58.

190. Eaton, James Howard. 1872. Report on the geology of the region

about Devil’s Lake. Trans. 1 : 124-128.

191. Eaton, James Howard. 1874. On the relation of the sandstone,

conglomerates and limestone of the Baraboo Valley to each other and to

the azoic quartzites. Trans. 2 : 123-127. Figs.

192. Eaton, Edward Dwight. 1898. James J. Blaisdell. Trans. 11 :

517-522.

193. Ekern, George L., and F. T. Thwaites. 1930. The Glover Bluff

structure, a disturbed area in the Palaeozoics of Wisconsin. Trans. 25 :

89-97. Figs., 1 pi.

194. Ellen, Sister Mary. 1924. Some ferns of southwestern Wisconsin.

Trans. 21 : 249-250.

195. Ellsworth, Elmer W., and W. L. Wilgus. 1930. The varved clay

deposit at Waupaca, Wisconsin. Trans. 25 : 99-111. Figs., 1 pi.

196. Ellsworth, Elmer W. 1932. Varved clays of Wisconsin. Trans.

27 : 00-00. Figs.

197. Elmendorf, John James. 1879. Nature and freedom. Trans. 4 :

62-76.

198. Elmendorf, John James. 1882. Nature and the supernatural.

Trans. 5 : 66-84.

199. Elmendorf, John James. 1892. Aristotle’s Physics (Physike akro-

asis) reviewed. Trans. 8 : 169-175.

200. Engelmann, P. 1871. On the importance of more attention to the

preservation and culture of forest trees in Wisconsin. (An abstract).

Bull. 2 : 30.

201. Engelmenn, P. 1871. Some observations upon the fauna of Mam¬

moth Cave. (An abstract). Bull. 3 : 39-40.

202. Evening Wisconsin, The. 1900. Alice Marian (Aikens) Bremer.

Trans. 12 (2) : 559. (Reprint of newspaper article).

203. Everest, Kate A. 1892. Early Lutheran immigration to Wiscon¬

sin. Trans. 8 : 289-298.

204. Fallows, Samuel. 1907. Stephen Vaughn Shipman. Trans. 15

(2) : 927-931. Portrait.

205. Fassett, Norman Carter. 1930. The plants of some northeastern

Wisconsin lakes. Trans. 25 : 157-168. Figs.

206. Fassett, Norman Carter. 1929-1932. Preliminary reports on the

flora of Wisconsin. (All accompanied by distribution maps.) I. Juncagi-

naceae, Alismaceae. Trans. 24 : 249-256 (1929). II. Ericaceae. Trans.

24 : 257-268 (1929). III. (See K. L. Mahony 1929). IV. (See L. R. Wil¬

son 1930). V. Coniferales. Trans. 25 : 177-182 (1930). VI. Pandanales.

Trans. 25 : 183-187 (1930). VII. Betulaceae. Trans. 25 : 189-194 (1930).

VIII. Aceraceae. Trans. 25 : 195-197 (1930). IX. Elatinaceae — water-

Index to Academy Publications, 1870-1932.

585

wort family. Trans. 25 : 199-200 (1930). X. Haloragidaceae — water mil¬

foil family. Trans. 25 : 201-203. (1930). XI. (See Lois Almon 1930).

XII. (See Edith W. Breakey 1931). XIII. (See D. F. Costello 1931). XIV.

(See W. T. McLaughlin 1931). XV. (See K. L. Mahony 1932). XVI.

Xyridales. Trans. 27 : 000-000 (1932). XVII. Myricaceae, Juglandaceae.

Trans. 27 : 000-0 (1932). XVIII. (See Florence B. Livergood 1932).

XIX. Saxifragaceae. Trans. 27 : 000-000 (1932). XX. (See Alice M.

Hagen 1932).

207. Faxon, Charles E., translator; see F. L. Grundtvig.

208. Feinberg, S. M., joint author; see J. W. Mavor.

209. Feuling, John B. 1871. On the place which the study of the In¬

dian languages should hold in ethnology. (A brief editor’s abstract).

Bull. 4 : 68.

210. Feuling, John B. 1872. Of the place of the Indian languages in the

study of ethnology. Trans. 1 : 178.

211. Feuling, John B. 1874. The etymology of “church”. Trans. 2 :

182-192.

212. Feuling, John Baptist. 1876. Studies in comparative grammar.

Trans. 3 : 117-121.

213. Fillmore, John Comfort. 1898. The forms spontaneously assumed

by folk-songs. Trans. 11 : 119-126. With music.

214. Finch, A. J. 1871. On metallic veins and the deposition of miner¬

als. (An abstract). Bull. 4: 50-53.

215. Fish, Carl Russell. 1907. Table illustrating the progress of rota¬

tion in office to 1835. Trans. 15 (2) : 709-712.

216. Flint, Albert S to well. 1895. Note on the progress of meridian

transit observations for stellar parallax at the Washburn observatory.

Trans. 10 : 64-65.

217. Flint, Albert Stowell. 1900. Christian Preusser. Trans. 12 (2) :

557-558.

218. Fluke, Charles Lewis, jr. 1921. Syrphidae of Wisconsin. Trans.

20 : 215-253. 2 pis.

219. Fluke, Charles Lewis, jr. 1931. Notes on certain syrphus flies re¬

lated to Xanthogramma (Diptera, Syrphidae) with descriptions of two

new species. Trans. 26 : 289-309. 2 pis.

220. Fluke, C. L. , joint author; see C. H. Curran.

221. Folkmar, Daniel. 1898. The duration of school attendance in Chi¬

cago and Milwaukee. Trans. 12 (1) : 253-305.

222. Foye, Janette W. 1900. James Clark Foye. Trans. 12 (2) : 560.

223. Fred, E. B., joint author; see Laetitia M. Snow.

224. Frey, Charles N. 1924. The cytology and physiology of Venturia

inequalis (Cooke) Winter. Trans. 21 : 303-343. 2 pis.

225. Frost, William Dodge. 1914. The bacteriological control of public

milk supplies. Trans. 17 (2) : 1305-1365. Figs., 2 pis.

226. Frost, William Dodge, and Ruth Chase Noland. 1924. New and

corrected names of certain milk bacteria. Trans. 21 : 219-222.

227. Frost, W. D., joint author; see Ola E. Johnston.

586 Wisconsin Academy of Sciences, Arts, and Letters .

228. Giese, William Frederic. 1907. Amos Arnold Knowlton. Trans.

15 (2) : 915-917. Portrait.

229. Gilbert, Edward Martinius. 1910. Studies on the Tremellineae of

Wisconsin. Trans. 16 (2) : 1137-1170. 3 pis.

230. Gilbert, Edward Martinius. 1921. Cytological studies of the lower

Basidiomycetes. I. Dacrymyces. Trans. 20 : 387-397. Fig., 1 pi.

231. Graenicher, Sigmund. 1905. The relations of the andrenine bees

to the entomophilous flora of Milwaukee County. Trans. 15 (1) : 89-97.

232. Granovsky, Alexander Anastacievitch. 1929. Preliminary studies

of the intracellular symbionts of Saissetia oleae (Bernard). Trans. 24 :

445-456. 3 pis.

233. Greene, Henry C. 1932. Wisconsin Myxomycetes. Trans. 27 :

000-000. 6 pis.

234. Gregory, Charles Noble. 1895. Political corruption and English

and American laws for its prevention. Trans. 10 : 262-298.

235. Gregory, John Goadby. 1898. Negro suffrage in Wisconsin.

Trans. 11 : 94-101.

236. Gregory, John Goadby. 1907. John Lendrum Mitchell. Trans. 15

(2) : 920-923. Portrait.

237. Grossenbacher, John Gasser. 1875. The periodicity and distribu¬

tion of radial growth in trees and their relation to the development of “an¬

nual” rings. Trans. 18 (1) : 1-77.

238. Grundtvig, Frederik Lange. 1895. On the birds of Shiocton in

Bovina, Outagamie County, Wisconsin. (Translated by Charles Edward

Faxon.) Trans. 10 : 73-158. 1 pi.

239. Hadden, Clarence Bernard. 1895. History of early banking in

Wisconsin. Trans. 10 : 159-198.

240. Haertel, Martin Henry. 1914. Social conditions in southern Ba¬

varia in the thirteenth century, as shown in Meier Helmbrecht. Trans. 17

(2) : 1057-1072.

241. Hagen, Alice M. 1932. Preliminary reports on the flora of Wis¬

consin. XX. Malvales. Trans. 27 : 000-000.

242. Haldeman, Samuel Stehman. 1874. On several points in the pro¬

nunciation of Latin and Greek. Trans. 2 : 178-181.

243. Hardenberg, Christian Bernhardus. 1907. Comparative studies

in the trophi of the Scarabaeidae. Trans. 15 (2) : 548-602. 5 pis.

244. Harder, H. P., joint author; see E. A. Birge.

245. Harper, Charles Lewis. 1902. Willard Harris Chandler. Trans.

13 (2) : 617-620. Portrait.

246. Harper, Edward Thompson. 1914. Species of Pholiota of the region

of the Great Lakes. Trans. 17 (1) : 470-502. 32 pis.

247. Harper, Edward Thompson. 1914. Species of Pholiota and Stro-

pharia in the region of the Great Lakes. Trans. 17 (2) : 1011-1026. 9 pis.

248. Harper, Edward Thompson. 1914. Species of Hypholoma in the

region of the Great Lakes. Trans. 17 (2) : 1142-1164. 13 pis.

249. Harper, Edward Thompson. 1916. Additional species of Pholiota,

Stropharia and Hypholoma in the region of the Great Lakes. Trans. 18

(2) : 392-421. 14 pis.

Index to Academy Publications , 1870-1932 .

587

250. Harper, Edward Thompson. 1921. Species of Lentinus in the re¬

gion of the Great Lakes. Trans. 20 : 365-385. 15 pis.

251. Harper, Robert Aimer. 1900. Nuclear phenomena in certain

stages in the development of the smuts. Trans. 12 (2) : 475-498. 2 pis.

252. Harper, Robert Aimer. 1904. Hamilton Greenwood Timberlake.

Trans. 14 (2) : 690-693. Portrait.

253. Harper, Robert Aimer, joint author; see R. J. Holden.

254. Harring, Harry K., and Frank J. Myers. 1921-1930. The rotifer

fauna of Wisconsin. I. Trans. 20 : 553-662 (1921). 21 pis. II. Trans.

21 : 415-549 (1924). 28 pis. III. Trans. 22 : 315-423 (1926). 40 pis. IV.

Trans. 23 : 667-808 (1927). 27 pis. V. Trans. 25 : 353-413 (by Myers

alone, 1930). 17 pis.

255. Harvey, Ellery Hale, and Henry August Schuette. 1930-1931. The

sulfur monochloride reaction of the fatty oils. I. (Published in Jr. Ind.

Engr. Chem. 1930). II. On the nature of the reaction product. Trans. 26 :

225-230 (1931). III. A note on the thermal behavior of their fatty acids.

Trans. 26 : 231-232 (1931). IV. On the evolution of hydrogen chloride.

Trans. 26 : 233-239 (1931). Fig.

256. Harvey, James Vernon. 1927. A survey of water molds occurring

in the soils of Wisconsin, as studied during the summer of 1926. Trans.

23 : 551-565. 4 pis.

257. Hastings, Samuel Dexter. 1872. The present condition of the com¬

mon jails of the country. Trans. 1 : 90-97.

258. Helgeson, Earl A. 1932. Impermeability in mature and immature

sweet clover seeds as affected by conditions of storage. Trans. 27 : 000-000.

259. Higley, William Kerr. 1889. Reptilia and Batrachia of Wisconsin.

Trans. 7 : 155-176.

260. Hillyer, Homer Winthrop. 1895. On the action of aluminum chlo¬

ride on saturated hydrocarbons. Trans. 10 : 367-369.

261. Hobbs, William Herbert. 1892. On some metamorphosed eruptives

in the crystalline rocks of Maryland. Trans. 8 : 156-160. Figs., 1 pi.

262. Hobbs, William Herbert. 1892. Note on cerussite from Illinois

and Wisconsin. Trans. 8 : 399-400.

263. Hobbs, William Herbert. 1893. Notes on a trip to the Lipari

Islands in 1889. Trans. 9 : 21-32. 1 pi.

264. Hobbs, William Herbert. 1902. Edward Orton. Trans. 13 (2) :

610-613. Portrait.

265. Hobbs, William Herbert. 1905. The correlation of fracture sys¬

tems and the evidences of planetary dislocations within the earth’s crust.

Trans. 15 (1) : 15-29. Map.

266. Hobbs, William Herbert. 1907. Nathaniel Southgate Shaler.

Trans. 15 (2) : 924-927. Portrait.

267. Hobbs, William Herbert, joint author; see G. E. Culver.

268. Hoffman, Alice E., joint author; see H. A. Schuette.

269. Holden, Roy Jay, and R. A. Harper. 1903. Nuclear divisions and

nuclear fusion in Coleosporium sonchi-arvensis Lev. Trans. 14 (1) :

63-82. 2 pis.

270. Holland, Frederic May. 1874. Vexed questions in ethics. Trans.

2 : 35-38.

588 Wisconsin Academy of Sciences, Arts, and Letters .

271. Holland, Frederic May. 1874. Records of marriages. Trans. 2:

73-76.

272. Holland, Frederic May. 1876. Industrial education. Trans. 3 :

136-142.

273. Holland, Frederic May. 1876. The boa constrictor of politics.

Trans. 3 : 151-160.

274. Holland, Frederic May. 1876. Were the Stoics utilitarians?

Trans. 3 : 179-195.

275. Holmes, Samuel Jackson. 1909. Description of a new subterrane¬

an amphipod from Wisconsin. Trans. 16 (1) : 77-80. 2 pis.

276. Holmes, Samuel Jackson. 1910. Description of a new species of

Eubranchipus from Wisconsin with observations on its reaction to light.

Trans. 16 (2) : 1252-1255. 1 pi.

277. Homberger, A. W., joint author; see Victor Lenher.

278. Hopkins, Ervin William. 1929. Microchemical tests on the cell

walls of certain Fungi. Cellulose and chitin. Trans. 24 : 187-196.

279. Hoskins, Leander Miller. 1895. Maximum stresses in bridge mem¬

bers. Trans. 10 : 24-40. 1 pi.

280. Hoy, Philo Romayne. 1871. Abstract of a paper on the fauna of

Lake Michigan off Racine. Bull. 2 : 34-35.

281. Hoy, Philo Romayne. 1871. The Mammalia of Wisconsin. (An

abstract.) Bull. 4 : 62.

282. Hoy, Philo Romayne. 1872. Deep-water fauna of Lake Michigan.

Trans. 1 : 98-101.

283. Hoy, Philo Romayne. 1872. Insects injurious to agriculture.

Aphides (plant lice). Trans. 1 : 110-116.

284. Hoy, Philo Romayne. 1874. Natural history as a branch of ele¬

mentary education. Trans. 2 : 105-106.

285. Hoy, Philo Romayne. 1874. Some peculiarities of the fauna near

Racine. Trans. 2 : 120-122.

286. Hoy, Philo Romayne. 1876. Fish-culture. Trans. 3 : 37-39.

287. Hoy, Philo Romayne. 1876. On the extent of the Wisconsin fish¬

eries. (An abstract of notes sent by Dr. Hoy). Trans. 3 : 65-67.

288. Hoy, Philo Romayne. 1876. On the Catocalae of Racine County.

Trans. 3 : 96-98.

289. Hoy, Philo Romayne. 1876. Increase A. Lapham. Trans. 3 :

264-267.

290. Hoy, Philo Romayne. 1879. How did the aborigines of this coun¬

try fabricate copper implements? Trans. 4 : 132-137.

291. Hoy, Philo Romayne. 1879. Why are there no upper incisors in

the Ruminantia? Trans. 4 : 147-150.

292. Hoy, Philo Romayne. 1882. Water puppy. (Menobranchus later¬

alis Say.) Trans. 5 : 248-250.

293. Hoy, Philo Romayne. 1882. The larger wild animals that have be¬

come extinct in Wisconsin. Trans. 5 : 255-257.

294. Hoy, Philo Romayne. 1885. Who built the mounds? Trans. 6 :

84-100.

295. Hoy, Philo Romayne. 1885. Who made the ancient copper imple¬

ments? Trans. 6 : 101-106.

Index to Academy Publications, 1870-1932.

589

296. Hoyt, John Wesley. 1874. Requisites to a reform of the civil

service. Trans. 2 : 89-104.

297. Hoyt, John Wesley. 1876. On the formal commendation of gov¬

ernment officials. Trans. 3 : 133-135.

298. Hoyt, John Wesley. 1876. On the revolutionary movement among

women. Trans. 3 : 161-176.

299. Hoyt, John Wesley. 1910. Some personal recollections of Abra¬

ham Lincoln. Trans. 16 (2) : 1305-1309.

300. Hubbell, Herbert Porter. 1876. An examination of Prof. S. H.

Carpenter’s position in regard to evolution. Trans. 3 : 196-202.

301. Huntington, Ellsworth. 1898. Experiments with available road¬

making materials of southern Wisconsin. Trans. 11 : 249-254.

302. Irving, Roland Duer. 1872. The age of the quartzite, schists and

conglomerates of Sauk Co., Wis. Trans. 1 : 129-137. Figs., map.

303. Irving, Roland Duer. 1874. On some points in the geology of

northern Wisconsin. Trans. 2 : 107-119. Map.

304. Irving, Roland Duer. 1874. On a hand specimen, showing the ex¬

act junction of the primordial sandstones, and Huronian schists. Trans.

2 : 139.

305. Irving, Roland Duer. 1874. On the occurrence of gold and silver in

minute quantities in quartz from Clark County. Trans. 2 : 140-141.

306. Irving, Roland Duer. 1876. On kaolin in Wisconsin. Trans. 3 :

3-30. Fig.

307. Jackson, Hartley Harrad Thompson. 1927. Notes on the summer

birds of Door Peninsula, Wisconsin, and adjacent islands. Trans. 23 :

639-665. Map.

308. Jansky, Cyril Methodius. 1931. Note on the evaluation of a se¬

ries. Trans. 26 : 223-224.

309. Jegi, John I. 1904. Auditory memory span for numbers in school

children. Trans. 14 (2) : 509-513.

310. Jewell, James Stewart. 1879. Mind in the lower animals. Trans.

4 : 164-187.

311. Johnson, Arden Richard. 1909. Electrolytic production of iodo¬

form. Trans. 16 (1) : 253-257. 4 pis.

312. Johnson, Arden Richard. 1914. The chemistry of boron and some

new organic-boron compounds. Trans. 17 (1) : 528-532.

313. Johnson, Roswell Hill. 1902. Axial bifurcation in snakes. Trans.

13 (2) : 523-538. 8 pis.

314. Johnson, Roswell Hill. 1907. The individuality and variation of

the pyloric caeca of the Centrarchidae. Trans. 15 (2) : 713-732. 5 pis.

315. Johnston, Ola E., and William Dodge Frost. 1924. The charac¬

teristics of certain fecal bacteria as shown by the little plate method.

Trans. 21 : 223-224. 1 pi.

316. Jolivette, Hally Delilia Mary. 1910. Spore formation in Geoglos-

sum glabrum Pers. Trans. 16 (2) : 1171-1190. 3 pis.

317. Jolivette, Hally D. M., joint author; see Ruth F. Allen.

590 Wisconsin Academy of Sciences , Arts, and Letters .

318. Jones, Edward David. 1895. The relation of economic crises to

erroneous and defective legislation, with especial reference to banking

legislation. Trans. 10 : 370-420.

319. Jones, Edward David. 1900. Chartism — a chapter in English

industrial history. Trans. 12 (2) : 509-529.

320. Jones, Jeanette. 1931. Note on the late Ordovician strata of the

Green Bay — Lake Winnebago region. Trans. 26 : 121-126.

321. Jones, Lewis Ralph. 1921. Experimental work on the relation of

soil temperature to disease in plants. Trans. 20 : 433-459. 5 pis.

322. Juday, Chancey. 1904. The diurnal movement of plankton Crus¬

tacea. Trans. 14 (2) : 534-568.

323. Juday, Chancey. 1907. Studies on some lakes in the Rocky and

Sierra Nevada mountains. Trans. 15 (2) : 781-793. 3 pis.

324. Juday, Chancey. 1909. Some aquatic invertebrates that live un¬

der anaerobic conditions. Trans. 16 (1) : 10-16.

325. Juday, Chancey, and George Wagner. 1909. Dissolved oxygen as

a factor in the distribution of fishes. Trans. 16 (1) : 17-22.

326. Juday, Chancey. 1910. Some European biological stations. Trans.

16 (2) : 1257-1277. 4 pis.

327. Juday, Chancey. 1914. A new species of Diaptomus. Trans. 17

(2) : 803-805. Fig.

328. Juday, Chancey. 1916. Limnological studies on some lakes in

Central America. Trans. 18 (1) : 214-250. Figs.

329. Juday, Chancey. 1916. Limnological apparatus. Trans. 18 (2) :

566-592. Figs., 5 pis.

330. Juday, Chancey. 1921. Quantitative studies of the bottom fauna

of Lake Mendota. Trans. 20 : 461-493.

331. Juday, Chancey. 1926. A third report on limnological apparatus.

Trans. 22 : 299-314. Figs.

332. Juday, Chancey, E. A. Birge, G. I. Kemmerer and R. J. Robinson.

1927. Phosphorus content of lake waters of northeastern Wisconsin.

Trans. 23 : 233-248.

333. Juday, Chancey, and E. A. Birge. 1930. The Highland Lake district

of northeastern Wisconsin and the Trout Lake limnological laboratory.

Trans. 25 : 337-352. Fig., map.

334. Juday, Chancey, and E. A. Birge. 1931. A second report on the

phosphorus content of Wisconsin lake waters. Trans. 26 : 353-382. Figs.

335. Juday, Chancey, and E. A. Birge. 1932. Dissolved oxygen and

oxygen consumed in the lake waters of northeastern Wisconsin. Trans.

27 : 000-000.

336. Juday, Chancey, joint author; see E. A. Birge.

337. Kahlenberg, Louis. 1903. Action of metallic magnesium upon

aqueous solutions. Trans. 14 (1) : 299-312.

338. Kahlenberg, Louis. 1905. On the nature of the process of osmosis

and osmotic pressure, with observations concerning dialysis. Trans. 15

(1) : 209-272. Figs.

339. Kahlenberg, Louis. 1910. President’s address, 1909. Some factors

in the progress of scientific research. Trans. 16 (2) : 1289-1304.

Index to Academy Publications , 1870-1932.

591

340. Kahlenberg, Louis, joint author; see E. C. Copeland.

341. Keene, Mary Lucille. 1919. Studies of zygospore formation in

Phycomyces nitens Kunze. Trans. 19 (2) : 1195-1220. 3 pis.

342. Kellogg, Louise Phelps. 1924. The removal of the Winnebago.

Trans. 21 : 23-29.

343. Kellogg, Louise Phelps. 1929. Wisconsin Indians during the

American revolution. Trans. 24 : 47-51.

344. Kellogg, Louise Phelps. 1931. The Menominee treaty at the Ce¬

dars, 1836. Trans. 26 : 127-135.

345. Kemmerer, George, joint author; see Rex J. Robinson; see also C.

J uday.

346. King, Charles Isaac. 1879. Boiler explosions. Trans. 4 : 151-163.

347. King, Franklin Hiram. 1909. On the suspension of solids in fluids

and the nature of colloids and solutions. Trans. 16 (1) : 275-288.

348. Kinley, David. 1893. The direction of social reform. Trans. 9 :

137-151.

349. Knaplund, Paul. 1924. The unification of South Africa: a study

in British colonial policy. Trans. 21 : 1-21.

350. Knapp, Joseph Gillett. 1871. Abstract of a paper on the coniferae

of the Rocky Mountains and their adaptation to the soil and climate of

Wisconsin. Bull. 2 : 30-31.

351. Knapp, Joseph Gillett. 1871. Abstract of a paper on the isother¬

mal lines of the northwest. Bull. 3 : 40-44.

352. Knapp, Joseph Gillett. 1871. On the climatic relations of the flora

of Wisconsin. (Title only.) Bull. 4 : 62.

353. Knapp, Joseph Gillett. 1871. On the ancient lakes of Wisconsin.

(Title only.) Bull. 5 : 76.

354. Knapp, Joseph Gillett. 1871. On the rocks and mines of the upper

Wisconsin river. (Title only.) Bull. 5 : 81.

355. Knapp, Joseph Gillett. 1872. Coniferae of the Rocky Mountains.

Trans. 1 : 117-123.

356. Knapp, Joseph Gillett. 1872. Ancient lakes of Wisconsin. Trans.

1 : 151-153.

357. Kremers, Edward. 1892. The limonene group of terpenes. Trans.

8 : 312-362.

358. Kremers, Edward. 1895. On the classification of carbon-com¬

pounds. Trans. 10 : 310-326.

359. Krey, August Charles. 1910. John of Salisbury’s knowledge of the

classics. Trans. 16 (2) : 948-987.

360. Kuhl, Ernest Peter. 1916. Chaucer’s burgesses. Trans. 18 (2) :

652-675.

361. Kumlein, Thure. 1876. On the rapid disappearance of Wisconsin

wild flowers; a contrast of the present time with thirty years ago. Trans.

3 : 56-57.

362. Kunz, Jakob, joint author; see V. E. Shelford.

363. Landsberg, Johann; see translation by Ernst Voss.

364. Langenhan, Henry August. 1921. The arsenical solutions. No. 1.

Liquor potassii arsenitis (Fowler’s solution). Trans. 20 : 141-197.

592 Wisconsin Academy of Sciences, Arts, and Letters .

365. Lapham, Increase Allen. 1871. On the age of the quartzite of

Baraboo. (A very brief abstract only.) Bull. 2 : 35.

366. Lapham, Increase Allen. 1871. On the classification of plants.

(An abstract.) Bull. 3 : 47-48.

367. Lapham, Increase Allen. 1872. On the classification of plants.

Trans. 1 : 102-109. Figs.

368. Lapham, Increase Allen. 1872. Oconomowoc Lake, and other

small lakes of Wisconsin, considered with reference to their capacity for

fish-production. Trans. 3 : 31-36. Figs., map.

369. Lapham, Increase Allen. 1876. The law of embryonic develop¬

ment — the same in plants as in animals. Trans. 3 : 110-113. Figs.

370. Lathrop, H. O. 1932. The geography of the upper Rock River

valley. Trans. 27 : 1-00.

371. Lawson, Publius Virgilius. 1921. Thure Kumlein. Trans. 20 :

663-686. 3 pis.

372. Legler, Henry Eduard. 1903. Origin and meaning of Wisconsin

place-names; with special reference to Indian nomenclature. Trans. 14

(1) : 16-39.

373. Legler, Henry Eduard. 1904. A Wisconsin group of German

poets. Trans. 14 (2) : 471-484.

374. Lehmann, Harriet. 1903. Variations in form and size of Cyclops

brevispinosus Herrick and Cyclops americanus Marsh. Trans. 14 (1) :

279-298. 4 pis.

375. Leland, E. R. 1876. In memoriam. Prof. Peter Engelmann. Trans.

3 : 258-263.

376. Leland, E. R. 1876. Additional tribute to the memory of Dr. Lap¬

ham. Trans. 3 : 268-269.

377. Lenher, Victor. 1903. Fluoride of gold. Trans. 14 (1) : 313-315.

378. Lenher, Victor, and Alfred Wilhelm Homberger. 1910. The gravi¬

metric determination of tellurium. Trans. 16 (2) : 1279-1285.

379. Leverett, Frank. 1889. Raised beaches of Lake Michigan. Trans.

7 : 177-192.

380. Leverett, Frank. 1892. On the correlation of moraines with raised

beaches of Lake Erie. Trans. 8 : 233-240. Map.

381. Levi, Mrs. Kate; see Kate A. Everest.

382. Libby, Orin Grant. 1900. A study of the Greenback movement.

Trans. 12 (2) : 530-543.

383. Libby, Orin Grant, F. Belle Stanton, Bernard M. Palmer, and Al¬

lard J. Smith. 1901. An economic and social study of the lead region in

Iowa, Illinois and Wisconsin. Trans. 13 (1) : 188-281. 4 pis.

384. Libby, Orin Grant. 1901. Some pseudo-histories of the American

revolution. Trans. 13 (1) : 419-425.

385. Libby, Orin Grant, joint author; see R. H. Downes.

386. Lincoln, Azariah Thomas. 1900. The electrical conductivity of

non-aqueous solutions. Trans. 12 (2) : 395-453. 6 pis.

387. Livergood, Florence B. 1932. Preliminary reports on the flora of

Wisconsin. XVIII. Sarraceniales. Trans. 27 : 000-000. Maps.

388. Loomis, Hiram Benjamin. 1892. The effects of changes in tem¬

perature on the distribution of magnetism. Trans. 8 : 273-287. 1 pi.

Index to Academy Publications, 1870-1982 .

593

389. Lounsbury, James Anderson. 1927. A case of phenomenal zoo¬

spore behavior of an apparently sterile Isoachlya and a description of the

plant. Trans. 23 : 539-549. 3 pis.

390. Lounsbury, James Anderson. 1930. Investigations on the nature

of Protoachlya paradoxa Coker. Trans. 25 : 215-225. Figs.

391. Lueders, Herman Frederick. 1895. The vegetation of the town

Prairie du Sac. Trans. 10 : 510-524. 1 pi.

392. Lueders, Herman Frederick. 1898. Floral structure of some

Gramineae. Trans. 11 : 109-111. 1 pi.

393. Lueders, Mrs. Edith (Silverfriend). 1907. Herman Frederick

Lueders. Trans. 15 (2) : 917-920. Portrait.

394. Lugg, Joseph Henry. 1929. Some notes on Allomyces arbuscula

Butler. Trans. 24 : 343-355. 1 pi.

395. Lurton, Freeman Ellsworth. 1914. A study of retarded children

in a group of northwestern school systems. Trans. 17 (1) : 275-298.

396. Lutman, Benjamin Franklin. 1910. Some contributions to the life

history and cytology of the smuts. Trans. 16 (2) : 1191-1244. 8 pis.

397. McAllister, Frederick. 1914. On the cytology and embryology of

Smilacina racemosa. Trans. 17 (1) : 599-660. 3 pis.

398. McLaughlin, Willard T. 1931. Preliminary reports on the flora

of Wisconsin. XIV. Hypericaceae. Trans. 26 : 281-288. Fig., maps.

399. McLeod, Andrew Finley. 1914. The Walden inversion— a critical

review. Trans. 17 (1) : 503-527.

400. McMurphy, J. G. 1879. Rotation as a factor of motion. Trans.

4 : 235-240. Figs.

401. McMynn, John Gibson. 1893. Philo Romayne Hoy (late president

of the Wisconsin academy of sciences, arts and letters.) Trans. 9 : lxxv-

lxxvii. Portrait.

402. Mahony, Kenneth L. 1929-1932. Preliminary reports on the flora

of Wisconsin. III. Lobeliaceae, Campanulaceae, Cucurbitaceae. Trans.

24 : 343-361 (1929). Maps. XV. Polygonaceae. Trans. 27 : 000-000

(1932.) Maps.

403. Main, Mrs. Angelia Kumlein. 1927. Yellow-headed blackbirds at

Lake Koshkonong and vicinity. Trans. 23 : 631-638.

404. Main, Mrs. Angelia Kumlein. 1929. Life and letters of Edward

Lee Greene. Trans. 24 : 147-185.

405. March, H. W., joint author; see E. A. Birge.

406. Marquette, William George. 1909. Concerning the organization of

the spore mother-cells of Marsilia quadrifolia. Trans. 16 (1) : 81-106.

2 pis.

407. Marsh, Charles Dwight. 1892. On the deep water Crustacea of

Green Lake. Trans. 8 : 211-213.

408. Marsh, Charles Dwight. 1892. Notes on depth and temperature of

Green Lake. Trans. 8 : 214-218. 1 pi.

409. Marsh, Charles Dwight. 1892. On the Cyclopidae and Calanidae

of central Wisconsin. Trans. 9 : 189-224. 4 pis.

410. Marsh, Charles Dwight. 1895. On two new species of Diaptomus.

Trans. 10 : 15-18. 1 pi.

594 Wisconsin Academy of Sciences , Arts, and Letters.

411. Marsh, Charles Dwight. 1898. On the limnetic Crustacea of Green

Lake. Trans. 11 : 179-224. 10 pis.

412. Marsh, Charles Dwight. 1900. On some points in the structure

of the larva of Epischura lacustris Forbes. Trans. 12 (2) 544-548. 2 pis.

413. Marsh, Charles Dwight. 1901. The plankton of fresh water lakes.

(Address of the retiring president). Trans. 13 (1) : 163-187.

414. Marsh, Charles Dwight. 1903. On a new species of Canthocamp-

tus from Idaho. Trans. 14 (1) : 112-116. 1 pi.

415. Marsh, Charles Dwight. 1907. A revision of the North American

species of Diaptomus. Trans. 15 (2) : 381-516. 14 pis.

416. Marsh, Charles Dwight. 1910. A revision of the North American

species of Cyclops. Trans. 16 (2) : 1067-1134. 10 pis.

417. Marsh, Charles Dwight. 1914. Structural abnormalities in Cope-

poda. Trans. 17 (1) : 195-196. 1 pi.

418. Marsh, Charles Dwight. 1914. On a new species of Diaptomus

from Colorado. Trans. 17 (1) : 197-199. 1 pi.

419. Marshall, Ruth. 1903. Ten species of Arrenuri belonging to the

subgenus Megalurus Thon. Trans. 14 (1) : 145-172. 5 pis.

420. Marshall, Ruth. 1904. A new Arrenurus and notes on collections

made in 1903. Trans. 14 (2) : 520-526. 1 pi.

421. Marshall, Ruth. 1910. The vegetation of Twin Island. Trans. 16

(2) : 773-797. 2 pis.

422. Marshall, Ruth. 1914. Some new American water mites. Trans. 17

(2) : 1300-1304. 2 pis.

423. Marshall, Ruth. 1921. New American water mites of the genus

Neumania. Trans. 20 : 205-213. 3 pis.

424. Marshall, Ruth. 1924. Arrhenuri from Washington and Alaska.

Trans. 21 : 213-218. 2 pis.

425. Marshall, Ruth. 1927. Water mites from China. Trans. 23 :

601-609. 3 pis.

426. Marshall, Ruth. 1929. The morphology and developmental stages

of a new species of Piona. Trans. 24 : 401-404. 1 pi.

427. Marshall, Ruth. 1930. The water mites of the Jordan Lake re¬

gion. Trans. 25 : 245-253. 2 pis.

428. Marshall, Ruth. 1931-1932. Preliminary list of the Hydracarina

of Wisconsin. I. The red mites. Trans. 26 : 311-319 (1931). 2 pis. II.

Trans. 27 : 000-000 (1932). 4 pis.

429. Marshall, William Stanley. 1903. Entocythere cambaria (nov.

gen, et nov. spec.), a parasitic ostracod. Trans. 14 (1) : 117-144. 4 pis.

430. Marshall, William Stanley, and Henry Severin. 1904. Some points

in the anatomy of Ranatra fused P. Beauv. Trans. 14 (2) : 487-508. 3 pis.

431. Marshall, William Stanley. 1905. The reproductive organs of the

female maia moth, Hemileuea maid (Drury). Trans. 15 (1) : 1-14. 2 pis.

432. Marshall, William Stanley. 1914. On the anatomy of the dragon¬

fly, Libellula quadrimaculata Linne. Trans. 17 (2) : 755-790. 4 pis.

433. Marshall, William Stanley. 1921. The development of the frenu¬

lum of the wax moth, Galleria mellonella Linn. Trans. 20 : 199-204. 1 pi.

Index to Academy Publications, 1870-1932.

595

484. Marshall, William Stanley. 1927. The development of the com¬

pound eye of the confused flour beetle, Tribolium confusum Jacq. Trans.

23 : 611-630. 4 pis.

485. Marshall, William Stanley. 1929-1930. The hypodermal glands of

the black scale, Saissetia oleae (Bernard). Trans. 24 : 427-443 (1929).

3 pis. II. The ventral glands. Trans. 25 : 255-272 (1930). 3 pis.

436. Martin, Ella May. 1924. Cytological studies of Taphrina coryli

Nishida on Corylus americana. Trans. 21 : 345-356. 2 pis.

437. Mason, Russell Z. 1871. Abstract of a paper on the nebular hy¬

pothesis in astronomy. Bull. 3 : 44-46.

438. Mason, Russell Z. 1871. On the clay deposits and the fossils found

therein, in the region about Appleton. (An abstract). Bull. 5 : 73-76.

439. Mason, Russell Z. 1879. The duty of the state in its treatment of

the deaf and dumb, the idiotic, the crippled and deformed, and the insane.

Trans. 4 : 25-30.

440. Mavor, James Watt, and William Strasser. 1916. On a new myx-

osporidian, Henneguya wisconsinensis, n. sp., from the urinary bladder of

the yellow perch, Perea flavescens. Trans. 18 (2) : 676-682. Fig.

441. Mavor, James Watt, and William Strasser. 1918. Studies of Myx-

osporidia from the urinary bladders of Wisconsin fishes. Trans. 19 (1) :

553-558. 3 pis.

442. Mavor, James Watt, and Samuel Maurice Feinberg. 1918. Lymph-

ocystis vitrei , a new sporozoan from the pike-perch, Stizostedion vitreum

Mitchell. Trans. 19 (1) : 559-561. 1 pi.

443. Maxson, Henry Doty. 1892. Lucius Heritage. Trans. 8 : 442-444.

444. Meachem, John Goldesborough, jr. 1900. John Goldesborough

Meachem. Trans. 12 (2) : 555-557.

445. Meloche, Villiers W., joint author; see Leslie Titus.

446. Merrell, Edward Huntington. 1898. The relation of motives to

freedom. Trans. 12 (1) : 389-393.

447. Merrill, Harriet Bell. 1893. The structure and affinities of

Bunops scutifrons Birge. Trans. 9 : 319-342. 2 pis.

448. Meyer, Balthasar Henry. 1898. The adjustment of railroad rates

in Prussia. Trans. 11 : 78-93.

449. Meyer, Balthasar Henry. 1898. Early general railway legislation

in Wisconsin, 1853-1874. Trans. 12 (1) : 337-388.

450. Miller, Eric Rexford. 1927. A century of temperatures in Wis¬

consin. Trans. 23 : 165-177. Figs.

451. Miller, Eric Rexford. 1929. Rainfall maps of Wisconsin and ad¬

joining states. Trans. 24 : 501-508. Maps.

452. Miller, Eric Rexford. 1930. Monthly rainfall maps of Wisconsin

and adjoining states. Trans. 25 : 135-156. Maps.

453. Miller, Eric Rexford. 1931. Extremes of temperature in Wiscon¬

sin. Trans. 26 : 61-68. Maps.

454. Miller, Eric Rexford. 1932. The distribution of cloudiness in Wis¬

consin. Trans. 27 : 00-00. Maps.

455. Miller, William Snow. 1895. The anatomy of the heart of Cam-

barus. Trans. 10 : 327-338. 2 pis.

596 Wisconsin Academy of Sciences , Arts , and Letters .

456. Miller, William Snow, and E. P. Carlton. 1895. The relation of

the cortex of the cat’s kidney to the volume of the kidney, and an estima¬

tion of the number of glomeruli. Trans. 10 : 525-538.

457. Miller, William Snow. 1904. Variations in the distribution of the

bile duct of the cat (Felis domesticus). Trans. 14 (2) : 621-629. Fig., 1 pi.

458. Mitchell, Irving Nelson. 1904. John I. Jegi. Trans. 14 (2) :

695-696.

459. Morris, Harold Hulett. 1918. The preparation of selenic acid,

Trans. 19 (1) : 369-373.

460. Morrison, Joseph Paul Eldred. 1929. A preliminary list of the

Mollusca of Dane County, Wisconsin. Trans. 24 : 405-425.

461. Morrison, Joseph Paul Eldred. 1932. A report on the Mollusca of

the northeastern Wisconsin lake district. Trans. 27 : 000-000. Figs.

462. Morrison, Joseph Paul Eldred. 1932. Studies on the life history

of Acella haldemani (“Desh.” Binney). Trans. 27 : 000-000. Figs., 2 pis.

463. Munro, Caroline Walker. 1921. A preliminary study of the diges¬

tive secretions of pickerel and perch. Trans. 20 : 269-273.

464. Munro, Dana Carlton. 1916. President’s address, 1915: Some tend¬

encies in history. Trans. 18 (2) : 695-712.

465. Murphy, Raymond Edward. 1931. The geography of the north¬

western pine barrens. Trans. 26 : 69-120. Figs., maps .

466. Murrish, John. 1871. Abstract of a paper on the origin of the

Potsdam sandstone. Bull. 3 : 32-33.

467. Murrish, John. 1871. Abstract of a paper on the results of recent

investigations in the lead regions of Wisconsin. Bull. 4 : 62-64.

468. Murrish, John. 1871. Further results of the examination of the

lead region of Wisconsin. (An abstract). Bull. 5 : 80.

469. Muttkowski, Richard Anthony. 1918. The fauna of Lake Men-

dota: a qualitative and quantitative survey with special reference to the

insects. Trans. 19 (1) : 374-482.

470. Myers, Frank J. 1930. The rotifer fauna of Wisconsin. V. The

genera Euchlanis and Monommata. Trans. 25 : 353-413. 17 pis.

471. Myers, Frank J., joint author; see H. K. Harring.

472. Nader, John. 1874. The strength of materials as applied to en¬

gineering. Trans. 2 : 153-160.

473. Nader, John. 1876. Leveling and the use of the barometer. Trans.

3 : 68-76.

474. Nader, John. 1876. Improvement of the mouth of the Mississippi

River. Trans. 3 : 84-95.

475. Nader, John. 1882. The tides. Trans. 5 : 207-233. Map.

476. Nader, John. 1882. A chapter on foundations. Trans. 5 : 282-289.

477. Nebel, Catherine Elizabeth. 1918. The amount of food eaten by

the spider, Aranea serieata. Trans. 19 (1) : 524-530.

478. Nettels, Curtis. 1927. The beginnings of money in Connecticut.

Trans. 23 : 1-28.

479. Nettels, Curtis. 1930. The national cost of the inland frontier,

1830. Trans. 25 : 1-37.

Index to Academy Publications , 1870-1932 .

597

480. Nichols, Susie Percival. 1905. The nature and origin of the bi-

nucleated cells in some Basidiomycetes. Trans. 15 (1) : 30-70. 3 pis.

481. Nicodemus, William Joseph Leonard. 1874. On the Wisconsin

River improvement. Trans. 2 : 142-152.

482. Nicodemus, William Joseph Leonard. 1874. Railway gauges.

Trans. 2 : 161-177.

483. Nicodemus, William Joseph Leonard. 1874. History of the science

of hydraulics. Trans. 2 : 193-202.

484. Nicodemus, William Joseph Leonard. 1876. On the ancient civili¬

zation of America. Trans. 3 : 58-64.

485. Noland, Lowell Evan, joint author; see W. J. Chase.

486. Noland, Mrs. Ruth (Chase). 1924. The anatomy of Troctes

divinatorius Muell. Trans. 21 : 195-211. 3 pis.

487. Noland, Ruth Chase, joint author; see W. D. Frost; see also Ruth

W. Chase.

488. Norton, Richard Greenleaf. 1907. An investigation into the

breaking of watch mainsprings in greater numbers in the warm months of

the year than in the cold months. Trans. 15 (2) : 654-655. 1 pi.

489. Oberholser, Harry Church. 1918. A review of the plover genus

Ochthodromus Reichenbach and its nearest allies. Trans. 19 (1) : 511-523.

490. Ockerman, John W. 1926. Fauna of the Galena limestone near

Appleton. Trans. 22 : 99-142. 3 pis.

491. Oldenhage, Henry F. 1879. Remarks on the descent of animals.

Trans. 4 : 138-146.

492. Olive, Edgar William. 1905. Notes on the occurrence of Oscilla-

toria prolifica (Greville) Gomont in the ice of Pine Lake, Waukesha Coun¬

ty, Wisconsin. Trans. 15 (1) : 124-134.

493. Olive, Edgar William. 1907. Cytological studies on Ceratiomyxa.

Trans. 15 (2) : 753-774. 1 pi.

494. Olson, O. A., joint author; see E. A. Birge.

495. O’Shea, Michael Vincent. 1905. The parts of speech in the child’s

linguistic development. Trans. 15 (1) : 178-208.

496. Overton, James Bertram. 1921. The organization of the nuclei in

the root tips of Podophyllum peltatum. Trans. 20 : 275-322. 1 pi.

497. Owen, Edward Thomas. 1898. The meaning and function of

thought-connectives. Trans. 12 (1) : 1-48.

498. Owen, Edward Thomas. 1901. A revision of the pronouns, with

special examination of relatives and relative clauses. Trans. 13 (1) :

1-140.

499. Owen, Edward Thomas. 1904. Interrogative thought and the

means of its expression. Trans. 14 (2) : 353-470.

500. Owen, Edward Thomas. 1909. Hybrid parts of speech, a develop¬

ment of this proposition : In a single sentence a word may operate, though

unrepeated, as different parts of speech. Trans. 16 (1) : 108-252.

501. Owen, Edward Thomas. 1914. The relations expressed by the

passive voice. Trans. 17 (1) : 77-148.

502. Owen, Edward Thomas. 1927. Linguistic aberrations. Trans.

23 : 255-528.

598 Wisconsin Academy of Sciences, Arts, and Letters .

503. Owen, Edward Thomas. 1931. Syntax of the adverb, preposition

and conjunction. Trans. 26 : 167-221.

504. Paetow, Louis John. 1909. The neglect of the ancient classics at

the early medieval universities. Trans. 16 (1) : 311-319.

505. Palmer, Bernard M., joint author; see 0. G. Libby.

506. Parkinson, John Barber. 1882. Wealth, capital and credit. 5 :

46-56.

507. Parkinson, John Barber. 1902. John Eugene Davies. Trans. 13

(2) : 614-617. Portrait.

508. Parkinson, John Barber. 1904. Samuel Dexter Hastings. Trans.

14 (2) : 686-690. Portrait.

509. Patten, Harrison Eastman. 1903. Action upon metals of solutions

of hydrochloric acid in various solvents. Trans. 14 (1) : 316-352. Figs.

510. Paxson, Frederic Logan. 1914. The railroads of the “Old North¬

west” before the civil war. Trans. 17 (1) : 243-274. Maps.

511. Payne, Alford. 1879. Art as education. Trans. 4 : 31-43.

512. Pearse, Arthur Sperry. 1914. On the habits of Uca yugnax

(Smith) and TJ. pugilator (Bose). Trans. 17 (2) : 791-802. Figs.

513. Pearse, Arthur Sperry. 1924. Observations on parasitic worms

from Wisconsin fishes. Trans. 21 : 147-160. 3 pis.

514. Pearse, Arthur Sperry. 1924. The parasites of lake fishes Trans.

21 : 161-194.

515. Peckham, Mrs. Elizabeth (Gifford), and George W. Peckham. 1882.

Temperature of Pine, Beaver and Okanchee lakes, Waukesha County,

Wisconsin, at different depths, extending from May to December, 1879;

also particulars of depths of Pine Lake. Trans. 5 : 271-275. Map.

516. Peckham, George Williams, and Elizabeth G. Peckham. 1885.

Genera of the family Attidae: with a partial synonymy. Trans. 6 :

255-342.

517. Peckham, George Williams, and Elizabeth G. Peckham. 1889. At¬

tidae of North America. Trans. 7 : 1-104. 6 pis.

518. Peckham, George Williams, and William H. Wheeler. 1889. Spi¬

ders of the sub-family Lyssomanae. Trans. 7 : 221-256.

519. Peckham, George Williams, and Elizabeth G. Peckham. 1895. The

sense of sight in spiders with some observations on the color sense. Trans.

10 : 231-261.

520. Peckham, George Williams, and Elizabeth G. Peckham. 1901.

Spiders of the Phidippus group of the family Attidae. Trans. 13 (1) :

282-358. 6 pis.

521. Peckham, George Williams, and Elizabeth G. Peckham. 1903. New

species of the family Attidae from South Africa, with notes on the dis¬

tribution of the genera found in the Ethiopian region. Trans. 14 (1) :

173-278. 11 pis.

522. Peckham, George Williams, and Elizabeth G. Peckham. 1907. The

Attidae of Borneo. Trans. 15 (2) : 603-653.

523. Peckham, George Williams, and Elizabeth G. Peckham. 1909. Re¬

vision of the Attidae of North America. Trans. 16 (1) : 355-646. 23 pis.

Index to Academy Publications, 1870-1932.

599

524. Peet, Stephen Denison. 1882. Primitive architecture in America.

Trans. 5 : 290-320. Figs.

525. Peet, Stephen Denison. 1885. Ancient villages among emblematic

mounds. 6 : 154-176. Plates.

526. Peet, Stephen Denison. 1889. The socalled elephant mound in

Grant County, and the effigies in the region surrounding it. Trans. 7 :

205-220. Figs.

527. Peet, Stephen Denison. 1892. The clan centers and clan habitat

of the effigy builders. Trans. 8 : 299-311. Figs., 1 pi.

528. Peet, Stephen Denison. 1909. The animal effigies of Wisconsin

and the totem system. Trans. 16 (1) : 320-324.

529. Pengra, Charlotte Elvira. 1904. On the conformal representation

of plane curves, particularly for the cases p = 4, 5 and 6. Trans. 14

(2) : 655-669.

530. Perrow, Eber Carle. 1914. The last will and testament as a form

of literature. Trans. 17 (1) : 682-753.

531. Piehl, Addie Emma. 1929. The cytology and morphology of Sor -

daria fimicola Ces. and De Not. Trans. 24 : 323-341. 2 pis.

532. Pierson, Merle. 1916. The relation of the Corpus Christi proces¬

sion to the Corpus Christi play in England. Trans. 18 (1) : 110-165.

533. Pietenpol, William Brasser. 1918. Selective absorption in the

visible spectrum of Wisconsin lake waters. Trans. 19 (1) : 562-593. Figs.

534. Plantz, Samuel. 1904. Dr. George McKendree Steele. Trans. 14

(2) : 678-682. Portrait.

535. Plumb, Ralph Gordon. 1914. Early harbor history of Wisconsin.

Trans. 17 (1) : 187-194.

536. Pope, Thomas Edmund Burt. 1930-1931. Wisconsin herpetologi-

cal notes. Trans. 25 : 273-284; 26 : 321-329.

537. Pradt, John B. 1874. In memoriam. William Edmond Armitage.

Trans. 2 : 253-254.

538. Preliminary reports on the flora of Wisconsin; see N. C. Fassett.

539. Puls, Arthur John. 1898. The need of a medical faculty in con¬

nection with the state university. Trans. 11 : 236-238.

540. Reed, George Matthew. 1905. Infection experiments with Ery-

siphe graminis DC. Trans. 15 (1) : 135-162.

541. Reed, George Matthew. 1907. Infection experiments with the mil¬

dew on cucurbits, Erysiphe cichor ace arum DC. Trans. 15 (2) : 527-547.

542. Regan, Katherine Patricia, joint author; see R. H. Downes.

543. Richardson, Robert Kimball. 1924. Augustine of Hippo qua pa¬

triot. Trans. 21 : 31-39.

544. Rickett, Harold William. 1921. A quantitative study of the larg¬

er aquatic plants of Lake Mendota. Trans. 20 : 501-527.

545. Ritson, Joseph; see Henry A. Burd.

546. Robinson, Francis J., joint author; see H. A. Schuette.

547. Robinson, Rex J., and George Kemmerer. 1930. Determination of

organic phosphorus in lake waters. Trans. 25 : 117-121.

548. Robinson, Rex J., and George Kemmerer. 1930. The determina¬

tion of Kjeldahl nitrogen in natural waters. Trans. 25 : 123-128. Figs.

600 Wisconsin Academy of Sciences , Arts , and Letters .

549. Robinson, Rex J., and George Kemmerer. 1930. Determination of

silica in mineral waters. Trans. : 129-134. Figs.

550. Robinson, Rex J., joint author; see Chancey Juday.

551. Roedder, Edwin Carl. 1921. Richard Wagner’s “Die Meistersing-

er von Nurnberg” and its literary precursors. Trans. 20 : 83-129.

552. Safford, Truman Henry. 1900. Combinations of Pythagorean tri¬

angles as giving exercises in computation. Trans. 12 (2) : 505-508.

553. Safford, Truman Henry. 1885. On the present state of our knowl¬

edge of stellar motion. Trans. 6 : 145-152.

554. Safford, Truman Henry. 1889. On the employment of the method

of least squares in the reduction of transit observations. Trans. 7 : 193-204.

555. Salisbury, Rollin D. 1885. Notes on the dispersion of drift copper.

Trans. 6 : 42-50. Map.

556. Sanborn, John Bell. 1898. Railroad land grants. Trans. 12 (1) :

306-316.

557. Sands, Mary Christina. 1907. Nuclear structure and spore forma¬

tion in Microsphaera alni. Trans. 15 (2) : 733-752. 1 pi.

558. Sawyer, Wesley Caleb. 1879. Letters an embarrassment to liter¬

ature. Trans. 4 : 50-55.

559. Sawyer, Wesley Caleb. 1882. The philosophy of F. H. Jacobi.

Trans. 5 : 146-160.

560. Sax, Mrs. H. D. M., see Hally D. M. Jolivette.

561. Schlatter, Edward Bunker. 1914. The development of the vowel

of the unaccented initial syllable in Italian. Trans. 17 (2) : 1073-1141.

562. Schlundt, Herman. 1910. The radioactivity of some spring waters

at Madison, Wisconsin. Trans. 16 (2) : 1245-1251.

563. Schorger, Arlie William. 1919. Contribution to the chemistry of

American conifers. Trans. 19 (2) : 728-766.

564. Schorger, Arlie William. 1929-1931. The birds of Dane County.

I. Trans. 24 : 457-499 (1929). 2 pis. II. Trans. 26 : 1-60 (1931). 2 figs.

565. Schubring, Selma Langenhan. 1926. A statistical study of lead

and zinc mining in Wisconsin. Trans. 22 : 9-98. Figs., maps.

566. Schuette, Henry August. 1918. A biochemical study of the plank¬

ton of Lake Mendota. Trans. 19 (1) : 594-613.

567. Schuette, Henry August, and Alice E. Hoffman. 1921, 1929. Notes

on the chemical composition of some of the larger aquatic plants of Lake

Mendota. Trans. 20 : 529-531. (1921). II. Vallisneria and Potamogeton.

(Hugo Alder, joint author). Trans. 24 : 249-254 (1929). Ill Castalia

odorata and Najas flexilis. (Hugo Alder, joint author). Trans. 24 : 135-

139 (1929).

568. Schuette, Henry August, and Hugo Alder. 1929. A note on the

chemical composition of Chara from Green Lake, Wisconsin. Trans. 24 :

141-145.

569. Schuette, Henry August, and Ralph W. Thomas. 1930. The com¬

position of the fat of the silver black fox. Trans. 25 : 113-116.

570. Schuette, Henry August, and Francis J. Robinson. 1932. The de¬

velopment of the ice cream freezer. Trans. 27 : 00-000. Figs.

571. Schuette, Henry August, joint author; see E. H. Harvey.

Index to Academy Publications , 1870-1932 .

601

572. Scott, Jonathan French. 1914. An investigation in regard to the

condition of labor and manufactures in Massachusetts, 1860-1870. Trans.

17 (1) : 167-186.

573. Secrist, Horace. 1914. The Anti-auction movement and the New

York Workingmen’s party of 1829. Trans. 17 (1) : 149-166.

574. Severin, Henry Herman Paul and Harry C. M. Severin. 1909.

Anatomical and histological studies of the digestive canal of Cimbex

americana Leach. Trans. 16 (1) : 38-60. 4 pis.

575. Severin, Henry Herman Paul, and Harry C. M. Severin. 1909.

Habits of the American saw-fly, Cimbex americana Leach, with observa¬

tions on its egg parasite, Trichogramma pretiosa Riley. Trans. 16 (1) :

61-76. 1 pi.

576. Severin, Henry, joint author; see W. S. Marshall.

577. Sharp, Frank Chapman. 1895. The personal equation in ethics.

Trans. 10 : 299-309.

578. Shelf ord, Victor Ernest, and Jakob Kunz. 1926. The use of photo¬

electric cells of different alkali metals and color screens in the measure¬

ment of light penetration into water. Trans. 22 : 283-298. Figs.

579. Sherman, William H. 1872. Vermilion by a new process — its pho¬

tographic properties. Trans. 1 : 165-166.

580. Shrosbree, George. 1909. The scientific development of taxidermy

and its effect upon museums. Trans. 16 (1) : 343-346.

581. Simmons, Henry Martyn. 1879. Mr. Spencer’s social anatomy.

Trans. 4 : 56-61.

582. Simons, Algie Martin. 1898. Railroad pools. Trans. 11 : 66-77.

2 pis.

583. Skinner, Ernest Brown. 1902. Truman Henry Safford. Trans.

13 (2) : 620-625.

584. Skinner, Ernest Brown. 1904. Frederick Pabst. Trans. 14 (2) :

693-694.

585. Skinner, Ernest Brown. 1907. The determination of the value of

the right of way of Wisconsin railroads as made in the appraisal of 1903.

Trans. 15 (2) : 794-822.

586. Slichter, Charles Sumner. 1898. Harmonic curves of three fre¬

quencies. Trans. 11 : 449-451.

587. Slichter, Charles Sumner. 1903. Recent criticism of American

scholarship. Address of the retiring president. Trans. 14 (1) : 1-15.

588. Smith, Allard J., joint author; see O. G. Libby.

589. Smith, Charles Forster. 1904. President Charles Kendall Adams.

Trans. 14 (2) : 670-678. Portrait.

590. Smith, Erastus Gilbert. 1901. On the determination of chlorine

in natural waters, its accuracy and significance. Trans. 13 (1) : 359-365.

591. Smith, Gilbert Morgan. 1914. The organization of the colony in

certain four-celled algae. Trans. 17 (2) : 1165-1220. 7 pis.

592. Smith, Gilbert Morgan. 1916. A monograph of the algal genus

Scenedesmus based upon pure culture studies. Trans. 18 (2) : 422-530.

9 pis.

593. Smith, Gilbert Morgan. 1916. A preliminary list of algae found

in Wisconsin lakes. Trans. 18 (2) : 531-565.

602 Wisconsin Academy of Sciences, Arts, and Letters .

594. Smith, Gilbert Morgan. 1918. A second list of algae found in

Wisconsin lakes. Trans. 19 (1) : 614-654. 6 pis.

595. Smith, Gilbert Morgan. 1921. The phytoplankton of the Muskoka

region, Ontario, Canada. Trans. 20 : 323-364. 6 pis.

596. Smith, Guy Harold. 1929. The settlement and the distribution of

the population in Wisconsin. Trans. 24 : 53-107. Maps.

597. Smith, John Y. 1871. Abstract of a paper on the laws which gov¬

ern the configuration of comets. Bull. 4 : 64-67.

598. Smith, John Y. 1874. The effect of duties on imports upon the

value of gold. Trans. 2 : 77-88.

599. Smith, Leonard Sewall. 1895. An experimental study of field meth¬

ods which will insure to stadia measurements greatly increased accuracy.

Trans. 10 : 539-555. Figs., 1 pi.

600. Smith, O. R. 1871. On the duty of the state to its idiotic chil¬

dren. (A brief abstract). Bull. 4 : 67.

601. Snow, Laetitia Morris and Edwin Brown Fred. 1926. Some charac¬

teristics of the bacteria of Lake Mendota. Trans. 22 : 143-154. 1 pi.

602. Squire, George Hall. 1909. Peculiar local deposits on bluffs adja¬

cent to the Mississippi. Trans. 16 (1) : 258-274. Figs., 1 pi.

603. Stanton, F. Belle, joint author; see O. G. Libby.

604. Stark, Marian E. 1932. Standards for predicting basal meta¬

bolism: I. Critical study of available standards for girls under 21, with a

proposed standard for predicting for girls from 17 to 21. Trans. 27 : 000-

000. Figs.

605. Stearns, John William. 1892. H. D. Maxson. Trans. 8 : 445-446.

606. Steele, George McKendree. 1874. Population and sustenance.

Trans. 2 : 59-72.

607. Stewart, Alban. 1916. Some observations concerning the botanical

conditions on the Galapagos Islands. Trans. 18 (1) : 272-340.

608. Stewart, Katharine Bernice, and Homer Andrew Watt. 1916.

Legends of Paul Bunyan, lumberjack. Trans. 18 (2) : 639-651.

609. Stout, Arlow Burdette. 1914. A biological and statistical analy¬

sis of the vegetation of a typical wild hay meadow. Trans. 17 (1) : 405-

469. 6 pis.

610. Strasser, William, joint author; see J. W. Mavor.

611. Strong, Edgar Freeman. 1898. The legal status of trusts. Trans.

11 : 127-148.

612. Stuart, James Reese. 1879. The harmonic method in Greek art.

Trans. 4 : 44-49.

613. Sweet, Edmund Theodore. 1876. Notes on the geology of northern

Wisconsin. Trans. 3 : 40-55.

614. Sweet, Edmund Theodore. 1876. On kerosene oil. Trans. 3 : 77-83.

615. Swezey, Goodwin De Loss. 1882. On some points in the geology of

the region about Beloit. Trans. 5 : 194-204. Plates.

616. Teller, Edgar Eugene. 1910. An operculated gastropod from the

Niagara formation of Wisconsin. Trans. 16 (2) : 1286-1288. 1 pi.

617. Thomas, Ralph W., joint author; see H. A. Schuette.

Index to Academy Publications, 1870-1932 .

603

618. Thompson, George. 1901. The Gothenburg method of regulating

the liquor traffic, 1892-1898. Trans. 13 (1) : 387-418.

619. Thwaites, Fredrik Turville. 1927. The development of the theory

of multiple glaciation in North America. Trans. 23 : 41-164.

620. Thwaites, Fredrik Turville. 1929. Glacial geology of part of

Vilas County, Wisconsin. Trans. 24 : 109-125. Figs.

621. Thwaites, Fredrik Turville, joint author; see George L. Ekern.

622. Thwaites, Reuben Gold. 1907. James Davie Butler. Trans. 15

(2) : 897-911. Portrait.

623. Timberlake, Hamilton Greenwood. 1902. Development and struc¬

ture of the swarm-spores of Hydrodictyon. Trans. 13 (2) : 486-522. 2 pis.

624. Titus, Leslie, and Villiers Willson Meloche. 1931. Notes on the

determination of total phosphorus in lake water residues. Trans. 26 :

441-444.

625. Todd, James Edward. 1882. A description of some fossil tracks

from the Potsdam sandstone. Trans. 5 : 276-281. Figs.

626. Tolman, Albert Harris. 1895. English surnames. Trans. 10 :

1-14.

627. Tolman, Herbert Cushing. 1892. The cuneiform inscriptions on

the monuments of the Achaemenides; translated. Trans. 8: 241-272. 2 pis.

628. Trelease, William. 1885. Preliminary list of Wisconsin parasitic

Fungi. Trans. 6 : 106-144.

629. Trelease, William. 1889. The morels and puff-balls of Madison.

Trans. 7 : 105-120. 3 pis.

630. Trelease, William. 1889. The “working” of the Madison lakes.

Trans. 7 : 121-129. 1 pi.

631. Tressler, Willis Lattanner and Bernard P. Domogalla. 1931. Lim¬

nological studies of Lake Wingra. Trans. 26 : 331-351. Figs., map.

632. True, R. H., joint author; see L. S. Cheney.

633. Turneaure, Frederick Eugene. 1904. John Butler Johnson. Trans.

14 (2) : 683-686. Portrait.

634. Ulrich, Edward Oscar. 1924. Notes on new names in table of

formations and on physical evidence of breaks between paleozoic systems

in Wisconsin. Trans. 21 : 71-107. Figs.

635. Urdahl, Thomas Klingenberg. 1898. The fee system in the Unit¬

ed States. Trans. 12 (1) : 49-252.

636. Van Cleef, Frank Louis. 1892. The pseudo-Gregorian drama

Xpiaros n Taoxuv in its relation to the text of Euripides. Trans. 8 : 363-378.

637. Van Hise, Charles Richard. 1892. The iron ores of the Lake Su¬

perior region. Trans. 8 : 219-228. 1 pi.

638. Van Hise, Charles Richard. 1895. The origin of the Dells of the

Wisconsin. Trans. 10 : 556-560.

639. Van Hise, Charles Richard. 1898. Earth movements. Address

of the retiring president. Trans. 11 : 465-516.

640. Vickery, R. A., joint author; see H. F. Wilson.

641. Vorhies, Charles Taylor. 1905. Habits and anatomy of the larva

of the caddis-fly, Platyphylax designatus, Walker. Trans. 15 (1) : 108-

123. 2 pis.

604 Wisconsin Academy of Sciences , Arts, and Letters .

642. Vorhies, Charles Taylor. 1909. Studies on the Trichoptera of

Wisconsin. Trans. 16 (1) : 647-738. Figs., 10 pis.

643. Voss, Ernst. 1907. Jakob Wimpheling’s “Tutschland”. Trans. 15

(2) : 823-873.

644. Voss, Ernst. 1907. An ordinance of the city of Nuremberg, adopt¬

ed in the year 1562. Trans. 15 (2) : 874-886.

645. Voss, Ernst. 1909. Decree of the honorable and wise council of

Nuremberg concerning the prohibition of the great vices of blasphemy,

swearing, carousing and treating. Trans. 16 (1) : 347-354.

646. Voss, Ernst. 1914. The regulations of the University of Witten¬

berg, issued in the year 1546, regarding the dress of the professors, their

wives and the student body. Trans. 17 (1) : 397-404.

647. Voss, Ernst. 1916. A true bit of instruction showing why we are

under obligations to pay taxes and tithes for the preservation of Christian

peace and the avoidance of trouble; written by Johannes Landtsperger, an

humble servant of Christ, 1528. Trans. 18 (2) : 683-694.

648. Voss, Ernst. 1927. Thomas Murner’s attitude in the controversy

between Reuchlin and the theologians of Cologne, based on his translations

from the Hebrew and the Epistolae obscurorum virorum, Vol. II, Nos. 3,

9, 59. Trans. 23 : 29-40.

649. Voss, Ernst. 1930. Karnoeffelspiel, a German card game of the

sixteenth century. Trans. 25 : 51-56.

650. Voss, Ernst. 1930. An ordinance of the city of Frankfort of the

year 1597 regulating dresses, marriages, baptisms, etc. Trans. 25 : 57-77.

651. Voss, Ernst. 1931. An edict of Philipp, by the grace of God, land¬

grave of Hessia, count of Catzenelenbogen, Dietz, Ziegenhain and Nidda —

How and in which form the Jews from now on shall be tolerated and

treated in our principality and our counties and dominions. 1539. Trans.

26 : 161-166.

652. Wadmond, Samuel C. 1910. Flora of Racine and Kenosha coun¬

ties, Wisconsin: a list of the fern and seed plants growing without cultiva¬

tion. Trans. 16 (2) : 798-888.

653. Wagner, George. 1909. Notes on the fish fauna of Lake Pepin.

Trans. 16 (1) : 23-37.

654. Wagner, George, joint author; see Chancey Juday.

655. Wakeman, Nellie Antoinette. 1919. Pigments of flowering plants.

Trans. 19 (2) : 767-906.

656. Wales, Julia Grace. 1932. Shakespeare’s use of English and for¬

eign elements in the setting of The Two Gentlemen of Verona. Trans.

27 : 000-000.

657. Walker, John Charles. 1924. On the nature of disease resistance

in plants. Trans. 21 : 225-247.

658. Walker, Milo Scott. 1898. Some uses of the low potential alter¬

nating current in a chemical laboratory. Trans. 11 : 114-118.

659. Walker, Ruth Irene. 1927. Cytological studies of some of the

short-cycled rusts. Trans. 23 : 567-582. 3 pis.

660. Walker, Ruth Irene, joint author; see Edith Breakey.

Index to Academy Publications, 1870-1932 .

605

661. Wallerstein, Ruth. 1932. Cowley as a man of letters. Trans. 27 :

000-000.

662. Wann, Louis. 1918. The influence of French farce on the Towne-

ley cycle of mystery plays. Trans. 19 (1) : 356-368.

663. Ward, Henry Levi. 1904. A study in the variations of propor¬

tions in bats, with brief notes on some of the species mentioned. Trans. 14

(2) : 630-654. 6 pis.

664. Ward, Henry Levi. 1909. Modern exhibitional tendencies of mu¬

seums of natural history and ethnography designed for public use. Trans.

16 (1) : 325-342.

665. Watrous, Jerome Anthony. 1907. Charles Frederick A. Zimmer¬

man. Trans. 15 (2) : 931-933. Portrait.

666. Watt, Homer A., joint author; see Katherine B. Stewart.

667. Wentzel, Erna Augusta. 1931. Morphological studies of Erysiphe

aggregate/, on Alnus incana. Trans. 26 : 241-252. 2 pis.

668. Wheeler, William Morton. 1892. On the appendages of the first

abdominal segment of embryo insects. Trans. 8 : 87-140. 3 pis.

669. Wheeler, William H., joint author; see George W. Peckham.

670. Wilgus, W. L., joint author; see E. W. Ellsworth.

671. Willard, S. W. 1885. Migration and distribution of North Ameri¬

can birds in Brown and Outagamie counties. Trans. 6 : 177-196.

672. Williams, Frank Ernest. 1921. The passing of an historic water¬

way. Trans. 20 : 131-140.

673. Wilson, Harley Frost, and R. A. Vickery. 1918. A species list of

the Aphididae of the world and their recorded food plants. Trans. 19 (1) :

22-355.

674. Wilson, Leonard R. 1930. Preliminary reports on the flora of

Wisconsin. IV. Lycopodiaceae, Selaginellaceae. Trans. 25 : 169-175.

675. Wilson, Leonard R. 1932. The Two Creeks forest bed, Manitowoc

County, Wisconsin. Trans. 27 : 00-00. Figs.

676. Wimmer, Edward Joseph. 1929. A study of two limestone quarry

pools. Trans. 24 : 363-399. Figs.

677. Wimpheling, Jakob; see Ernst Voss.

678. Wingate, Uranus Owen Brackett. 1898. The scientific importance

of more complete vital statistics of the state of Wisconsin. Trans. 11 :

102-108.

679. Wisconsin Academy of sciences, arts and letters. 1876. Report of

Committee on Exploration of Indian Mounds. Trans. 3 : 105-109. Figs.

680. Wisconsin academy of sciences, arts and letters. 1893. Catalog of

the library of the Academy, compiled by W. H. Hobbs. Trans. 9 : 1-212.

681. Wisconsin State Journal. 1879. Death of Professor S. H. Carpen¬

ter. (Reprinted.) Trans. 4 : 318-320.

682. Wolff, Henry Charles. 1900. The unsteady motion of viscous

liquids in capillary tubes. Trans. 12 (2) : 550-553.

683. Woltereck, Richard. 1932. Races, associations and stratification

of pelagic Daphnids in some lakes of Wisconsin and other regions of the

United States. Trans. 27 : 000-000. 6 pis.

684. Woodman, Edwin Ellis. 1882. The pipestone of DeviTs Lake.

Trans. 5 : 251-254.

606 Wisconsin Academy of Sciences, Arts, and Letters .

685. Wright, Albert C. 1871. Abstract of a paper on the classification

of the sciences. Bull. 2 : 27-29.

686. Wright, Albert 0. 1871. The metamorphic rocks in the town of

Portland, Dodge Co. (An abstract). Bull. 3 : 38-39.

687. Wright, Albert 0. 1871. The metamorphic rocks at Devil’s Lake.

(An abstract). Bull. 3 : 39.

688. Wright, Albert 0. 1871. The mineral well at Waterloo, Wiscon¬

sin. (An abstract.) Bull. 3 : 46-47.

689. Wright, Albert Orville. 1872. On the mineral well at Waterloo,

Wis. Trans. 1 : 154.

690. Wright, Albert Orville. 1882. The philosophy of history. Trans.

5 : 12-20.

691. Wright, Albert Orville. 1882. Distribution of profits. A new ar¬

rangement of that subject. Trans. 5 : 39-45.

692. Wright, Albert Orville. 1885. The increase of insanity. Trans.

6 : 20-28. Map.

693. Wright, Albert Orville. 1892. The defective classes. Trans. 8 :

176-186.

694. Wright, Stillman. 1927. A new species of Diaptomus from the

Philippine Islands. Trans. 23 : 583-586. 1 pi.

695. Wright, Stillman. 1927. A contribution to the knowledge of the

genus Pseudodiaptomus. Trans. 23 : 587-600. Fig.

696. Wright, Stillman. 1929. A preliminary report on the growth of

the rock bass, Ambloplites rupestris (Rafinesque) in two lakes of northern

Wisconsin. Trans. 24 : 581-595. Figs.

697. Young, Elaine Margaret. 1930. Physiological studies in relation

to the taxonomy of Monascus spp. Trans. 25 : 227-244. 2 pis.

698. Young, Karl. 1910. The harrowing of hell in liturgical drama.

Trans. 16 (2) : 889-947.

699. Young, Karl. 1914. Officium pastorum: a study of the dramatic

developments within the liturgy of Christmas. Trans. 17 (1) : 299-396.

700. Young, Karl. 1916. William Gager’s defence of the academic

stage. Trans. 18 (2) : 593-638.

701. Young, Karl. 1921. Ordo prophetarum. Trans. 20 : 1-82.

702. Zimmerman, Oliver Brunner. 1904. A treatment of instant angu¬

lar and linear velocities in complex mechanisms. Trans. 14 (2) : 514-519.

3 pis.

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