TRANSACTIONS

OF THE

WISCONSIN ACADEMY

OF

SCIENCES, ARTS AND LETTERS

VOL. XXIX

NATURAE SPECIES RATIOQUE

MADISON, WISCONSIN

1935

TRANSACTIONS

OF THE

WISCONSIN ACADEMY

OF

SCIENCES, ARTS AND LETTERS

VOL. XXIX

NATURAE SPECIES RATIOQU E

MADISON, WISCONSIN

1935

OFFICERS OF THE WISCONSIN ACADEMY OF SCIENCES,

ARTS AND LETTERS

President

Rufus M. Bagg, Lawrence College

Vice-Presidents

In Science: Storrs B. Barrett, Yerkes Observatory

In the Arts: Anselm M. Keefe, St. Norbert College

In Letters: A. R. Hohlfeld, University of Wisconsin

Secretary -Treasurer

H. A. Schuette, 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

Henry L. Ward, past president

M. A. Brannon, past president

L. J. Cole, past president

S. A. Barrett, past president

Charles E. Allen, 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

A. W. Schorger, Madison

W. S. Marshall, University of Wisconsin

A. L. Barker, Ripon College

Committee on Membership

The secretary, ex officio

J. 0. Carbys, Milwaukee

Paul W. Boutwell, Beloit College

G. W. Keitt, University of Wisconsin

C , 73

'La> 7 0dt?3

CONTENTS

Page

The Carbon Dioxide and Hydrogen Ion Content of the Lake Waters of North¬

eastern Wisconsin (32 text figures.) C. Juday, E. A. Birge and V. W.

Meloche . . . . 1

Further Notes on the Occurrence of Parasitic Copepods on Fish of the Trout

Lake Region, with a Description of the Male of Argulus biramosus. (Plate I.)

Ruby Bere . 83

Two New Subspecies of Fishes from Wisconsin. (Plates II and III.) Carl

L. Hubbs and C. Willard Greene . . . . . . 89

Growth of the Yellow Perch ( Perea jiavescens Mitchill) in Nebish, Silver and

Weber Lakes, Vilas County, Wisconsin. (S text figures.) Edward

Schneberger . . . . . . . . 103

Fish Food Studies of a Number of Northeastern Wisconsin Lakes. (11 text

figures.) Faye M. Couey . . 131

Photosynthesis of Algae at Different Depths in Some Lakes of Northeastern

Wisconsin. I. Observations of 1933. (7 text figures.) Harold A. Schomer

and Chancey Juday . 173

The Course and Significance of Sexual Differentiation. Charles E. Allen . 195

Wild Life Research in Wisconsin. Aldo Leopold . 203

Maple Sugar: A Bibliography of Early Records. H. A. Schuette and Sybil

C. Schuette . 209

The Geography of the Central Sand Plain of Wisconsin. (Plate IV and 7 text

figures.) J. Riley Staats . 237

Insoluble Residues from Wisconsin Sedimentary Rocks. (Plate V and 3 text

figures.) R. R. Shrock . . . . . . . . . . . 257

Preliminary List of the Hydracarina of Wisconsin. Part IV. (Plates VI to

XI.) Ruth Marshall . 273

Preliminary Reports on the Flora of Wisconsin. XXIV. Salicaceae. (4 text

figures.) David F. Costello . . . . . 299

Elaboration of Setting in Othello and the Emphasis of the Tragedy. Julia

Grace Wales . 319

The Literary Language and its Relation to the German Dialect. Alfred Senn.... 341

The Winnebago Visit to Washington in 1828. Louise Phelps Kellogg . 347

The Determination of Organic Nitrogen: Past and Present. H. A. Schuette

and Frederick C. Oppen . 355

The Cupro- Alkali Metal Carbonate Solution in the Determination of Reducing

Sugars. II. A Modification of Pellet’s Solution. (2 text figures.) Chang

Y. Chang and H. A. Schuette . 381

A Study of Ligneous Substances in Lacustrine Materials. John F. Steiner

and V. W. Meloche . . . . 389

Proceedings of the Academy . . . . . 403

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

should be directed to the secretary, H. A. Schuette, Chemistry Pudding, University of Wisconsin,

Madison, Wis. Publications intended for the Library of the Academy should be sent directly to the

Librarian, Walter M. Smith, University of Wisconsin Library, Madison, Wis.

THE CARBON DIOXIDE AND HYDROGEN ION CONTENT

OF THE LAKE WATERS OF NORTHEASTERN

WISCONSIN

C. Juday, E. A. Birge and V. W. Meloche

From the Limnological Laboratory of the Wisconsin Geological and Natural

History Survey.* Notes and reports No. 55.

I. Carbon Dioxide

Introduction

Dissolved carbon dioxide plays a very important role in the

biological economy of natural waters. It is one of the raw

materials required by aquatic plants for the manufacture of or¬

ganic substances in the process of photosynthesis. Its scarcity

or abundance, therefore, is of vital importance not only to the

aquatic plants but also to aquatic animals because the former

serve as the source of the food of the latter, either directly or

indirectly. Its significance from the standpoint of photosyn¬

thesis as well as from that of the chemical status of the water

makes it necessary to study this gas quantitatively in order to

get an idea of its ecological relations.

Such a quantitative study is particularly valuable for the

lakes of northeastern Wisconsin because their waters show such

a wide range in carbon dioxide content. The largest quantity

of carbon dioxide, including free, half-bound and bound, found

in any of these lakes was a little more than 75.0 mg/1, while

the smallest amount was a little less than 1.0 mg/1; thus the

quantity in the former sample was approximately eighty times

as large as that in the latter. The waters of a large percentage

of the lakes are quite soft. The Ca content in many instances

is less than 1.0 mg/1.

The carbon dioxide supply of lake waters is derived from the

atmosphere, from rain, from springs and streams, from the res¬

piration of living organisms and from the decomposition of

organic matter. Carbon dioxide is usually present in the at-

*This investigation was made in co-operation with the U. S. Bureau of Fisheries and the

results are published with the permission of the Commissioner of Fisheries.

1

ftc 3 '* 1935

2 Wisconsin Academy of Sciences, Arts, and Letters.

mosphere in the ratio of 3 to 4 parts per 10,000 and pure water

absorbs 1. 0-2.0 mg/1 when freely exposed to the air. An ex¬

periment with distilled water in 1931 yielded an average of 1.7

mg/1 of free carbon dioxide after exposure to the air for several

hours. Also a sample of rain water obtained on August 8, 1931,

yielded 1.0 mg/1 of free carbon dioxide. In certain types of

lakes, more especially the bog type, decomposition is the chief

source of the carbon dioxide supply.

Natural waters usually contain dissolved substances which

increase their capacity for carbon dioxide very materially. Cal¬

cium and magnesium play the most important role in that re¬

spect, but sodium and potassium play a part in some cases. Pure

water that is free of carbon dioxide will dissolve about 13.0

mg/1 of CaCO at 20° C. and about 60.0 mg/1 of MgCCh; the

bi carbonates of these two substances, however, are much more

soluble.

Rain water contains only a small amount of free carbon

dioxide when it falls on the land, but it acquires a larger supply

as it passes through the soil and subsoil. When this water that

is charged with free carbon dioxide comes into contact with

calcium and magnesium carbonates in the earth, it converts them

into bicarbonates which readily pass into solution. The quantity

of bicarbonates present in the ground water will depend upon

the amount of free carbon dioxide present in the percolating

water and also upon the abundance of CaCOs and MgCO in the

geological strata through which the water passes.

Methods and Nomenclature

Detailed descriptions of the various methods used in making

quantitative determinations of the carbon dioxide content of

natural waters have been given by a number of authors, more

recently by Werestschagin (1931) and Maucha (1932), so that

they need not be included in this report. The carbon dioxide re¬

sults presented here were obtained by titrating 100 cc samples

of water with N/44 Na^CCb and N/44 HC1, using phenolphthalein

and methyl orange for! indicators. Both of these titration

methods have their shortcomings, however. The results obtained

for free carbon dioxide are a little too low, but the error is neg¬

ligible in waters that are as soft as most of these. Likewise the

Juday, Birge & Meloche — Lake Waters of N. E. Wisconsin 3

HCl-methyl orange titration is subject to certain inaccuracies due

chiefly to the personal error involved in reading the endpoint. On

the other hand, both methods are the best that have been devised

for field work and they give results that are accurate enough for

general limnological purposes.

In previous reports dealing with the carbon dioxide content

of Wisconsin lake waters, the terminology used by Seyler (1894)

was employed, namely free, half-bound and fixed carbon dioxide.

The term “bound” is now more generally used than “fixed” and it

seems advisable to use the former instead of the latter word.

Pia (1933) has recently proposed another type of classification;

he uses the terms free and bound carbon dioxide, but separates

each of these classes into two components. The two kinds of

free are (1) “zugehorige freie Kohlensaure” and (2) “angrei-

fende” (aggressive). The former is defined as “die notwendig ist,

damit die vorhandenen Bikarbonate bestehen bleiben ; this may

be called “attached” free carbon dioxide because it is closely asso¬

ciated with the bicarbonate. The aggressive includes the free

carbon dioxide that is present in excess of the “attached”

(zugehorige) ; it is available for changing the monocarbonates

to bicarbonates.

According to Pia, the bound carbon dioxide consists of (1)

the “firmly-bound” (fest gebundene) and (2) the half-bound

(halbgebundene) . The firmly-bound carbon dioxide is that which

changes CaO into CaCCh and the half -bound is that which changes

CaCOs into Ca(HC03)2 or that which converts the monocarbon¬

ates into bicarbonates. In the present report on Wisconsin lakes,

the terms bound and half -bound are used; the former term is

equivalent to the “firmly-bound” carbon dioxide, but the prefix

“firmly” is omitted in order to simplify the expression.

Four different methods have been used by investigators for

stating the results of the HCl-methyl orange titration, namely,

(1) in terms of normality of the bicarbonates, (2) in cubic cen¬

timeters of N/10 HC1 or N/HC1, (3) as CaCCh and (4) in terms

of bound (fixed) carbon dioxide. It appears to the authors that

the last form of statement (4) is most convenient for limnolo¬

gical purposes because the limnologist is interested primarily in

the biological rather than in the purely chemical significance of

the carbon dioxide content of natural waters. A knowledge of

4 Wisconsin Academy of Sciences , Arts , and Letters.

the amount of this gas that is available for the photosynthetic

activities of the aquatic plants is especially important in bodies

of water with such limited supplies as many of these north¬

eastern lakes possess. If desired, the results given for bound

(firmly bound) carbon dioxide in this report can be changed into

terms of acid by dividing them by the number five ; the respect¬

ive quotients will represent the cubic centimeters of N/44 HC1

used in titrating 100 cc samples of water with methyl orange as

an indicator.

In the process of assimilation, aquatic plants are able to

use all of the free (aggressive and attached) and all of the half¬

bound carbon dioxide. Also Schutow (1926), Maucha (1929),

Neresheimer and Ruttner (1929) and Wiebe (1931) have re¬

ported that the bound or monocarbonate carbon dioxide can be

utilized after the free and half -bound have been exhausted. In

general however, the free and half-bound are the main sources

upon which the plants depend for their supply of this gas.

In waters that give a neutral or acid reaction to phenolphthal-

ein, the quantity of the half-bound is usually regarded as equal

to that of the bound carbon dioxide; this is only approximately

true, however, but the difference is not very great in waters that

fall within the carbon dioxide range of these lakes. The carbon

dioxide that is readily available for the aquatic plants, namely

the free and the half -bound, is thus substantially equal to the

sum of the free and the bound, while the total quantity is the

sum of the free and twice jthe bound.

An alkaline reaction to phenolphthalein shows that a certain

amount of normal or monocarbonate carbon dioxide is present

and the quantity is determined by the titration with acid until

the pink color is discharged. The samples that gave an alkaline

reaction to phenolphthalein are indicated by a minus sign in the

free carbon dioxide column of the tables of this report and also

in the text; the number following the minus sign shows the

quantity of the normal or monocarbonate carbon dioxide present.

In the case of these lakes, it may also be regarded as represent¬

ing the amount of half-bound carbon dioxide that has been util¬

ized by the aquatic plants in the process of photosynthesis. The

quantity of half-bound carbon dioxide remaining in these sam¬

ples, therefore, is equal to the algebraic sum of the monocarbon-

Juday, Birge & Meloche—Lake Waters of N. E. Wisconsin 5

ate as indicated by the minus sign and the bound carbon dioxide,

and the total is the sum of the half-bound and the bound.

Including both phenolphthalein and methyl orange titrations,

approximately 5,600 carbon dioxide determinations were made

on 2,800 samples of water during this investigation. This num¬

ber represents 2,400 determinations on surface samples and

3,200 on samples taken at other depths. Readings were ob¬

tained from 537 lakes, but the results on 19 are omitted in the

present report because either the free carbon dioxide or the

hydrogen ion concentration was not determined on them. The

observations were limited to a single surface sample on 283

lakes, but two or more samples were obtained from all of the

others. Series of samples covering the entire depth were taken

in 82 lakes ; all of those known to have a maximum depth of 18 m.

or more were included in the series as well as a number of lakes

ranging from 4 m). to 18 m. in depth. The series taken on the

various lakes consisted of 2 to 13 samples each, depending upon

the depth of the lake and the status of the carbon dioxide at the

different depths. The largest number of determinations was

made on Trout Lake, namely 278.

Seepage and Drainage Lakes

A large number of the northeastern lakes do not have an in¬

let or an outlet; they receive water through direct precipitation

on their surfaces and from the surface drainage of limited basins.

Any gain from or loss to the ground water takes place through

the process of seepage ; hence they have been designated as “seep¬

age” lakes. In general the seepage lakes are characterized by

their very soft waters; results from 238 of them are included

in this report.

Those bodies of water which have temporary or permanent

outlets have been called “drainage” lakes. Some of them show

characteristics akin to those of the seepage lakes because they

have no inlets and their outlets possess water only for a brief

period each year, or sometimes only at intervals of several years,

depending upon the amount of precipitation. Observations on

280 drainage lakes are included in this report.

6 Wisconsin Academy of Sciences, Arts, and Letters.

FREE CARBON DIOXIDE

Surface samples . Phenolphthalein titrations were made on

1,170 surface samples from 518 lakes. The largest number was

made on Trout Lake, namely 26. In 121 samples the water gave

an alkaline reaction to phenolphthalein; the reaction was neutral

in 14 and acid in 1,035. Thus approximately 12 per cent of the

samples gave a neutral or alkaline reaction to this indicator and

88 per cent an acid reaction.

Samples from 74 lakes gave an alkaline reaction to phenol¬

phthalein ranging from a minimum of — 0.1 mg/1 in Little St.

Germain Lake to a maximum of — 10.7 mg/1 in Mann Lake,

when expressed in terms of carbon dioxide. This alkaline re¬

action to phenolphthalein was due to the photosynthetic activities

of the various aquatic plants, chiefly to phytoplankton forms;

these organisms removed some of the half-bound carbon dioxide

from the bicarbonates during the process of assimilation, thus

leaving a certain amount of normal carbonate in the water which

gave it an alkaline reaction to phenolphthalein. All of the lakes

on which this phenomenon was observed belong to the drainage

class and their specific conductances ranged from a minimum of

23 x lO 6 in Adelaide Lake to a maximum of 132 x Id6 in Wild

Cat Lake.

The sample which gave the maximum alkaline reaction to

phenolphthalein was obtained from Mann Lake on July 7, 1928;

this lake has an area of about 100 ha. and a maximum depth of

2.5 m. It is fed by springs and has an outlet; the specific con¬

ductance of the water averages about 100 x 10'6. The quantity

of bound carbon dioxide on the above date was 25.6 mg/1, so

that a little more than 41 per cent of the half-bound carbon diox¬

ide had been utilized by the aquatic plants. The phytoplankton

organisms numbered 3,200 cells and colonies per cubic centimeter

on this date and there was also a luxuriant growth of the large

aquatics. Six surface samples were obtained from Mann Lake

between 1926 and 1930, inclusive; 5 of them gave an alkaline

reaction to phenolphthalein ranging from —2-8 to — 10.7 mg/1

and one gave an acid reaction equivalent to 4.0 mg/1 of free

carbon dioxide.

Fourteen surface samples obtained from 11 different lakes

gave a neutral reaction to phenolphthalein; with one exception

Juday, Birge & Meloche — Lake Waters of N. E. Wisconsin 1

these samples came from drainage lakes. The exception was

Pallette Lake; the specific conductance of this seepage lake

averaged 19 x Id6 and the bound carbon dioxide was 3.8 mg/1.

In the other 10 lakes whose surface samples gave a neutral re¬

action to phenolphthalein, the specific conductance varied from

53 to 120 x 10'6 and the bound carbon dioxide content fell be¬

tween 11.8 and 30.5 mg/1. In 4 of these lakes, the surface water

gave an alkaline reaction to phenolphthalein on other dates.

Allen or Brazell Lake ranked second in respect to its alka¬

line reaction to phenolphthalein; it was equivalent to — 6.0 mg/1

of carbon dioxide on July 8, 1928. It is a drainage lake with a

specific conductance of 62 x Id6 and a bound carbon dioxide

content of 15.0 mg/1 ; it has an area of 14 ha. and a maximum

depth of 2 m. The algal population on the above date was

2,950 cells and colonies per cubic centimeter of water.

All of the other surface samples, namely 1,035, gave an acid

reaction to phenolphthalein, thus indicating the presence of

free carbon dioxide. The results obtained for free carbon diox¬

ide in the various lakes are summarized in Tables I and II and

they are shown graphically in Fig. 1. In compiling the tables,

the mean of the various observations on a given lake has been

used in assigning it to a particular group when more than one

determination was made. In computing the means for some of

the lakes in which the surface water gave an alkaline reaction

to phenolphthalein the samples that gave an acid reaction more

than balanced those that were alkaline in reaction ; in such cases

the lakes were placed in the acid groups. As a result, only 36

of the 74 lakes in which some of the surface samples gave an

alkaline reaction to phenolphthalein fell permanently into the

alkaline groups as shown in Table II. For purposes of compar¬

ison the 518 lakes have been separated into seepage and drain¬

age types.

Seepage lakes. The mean results for free carbon dioxide

in the 238 seepage lakes are given in Table I. The surface water

of all lakes in this group gave an acid reaction to phenolphthal¬

ein. One surface sample from Pallette Lake gave a neutral re¬

action to phenolphthalein, but the other 5 samples from this

lake gave an acid reaction to this indicator, so that the mean

falls in the acid group.

8 Wisconsin Academy of Sciences, Arts, and Letters .

Fig. 1. Free carbon dioxide in surface waters of seepage and drainage lakes.

The diagram is based on results from 238 seepage and 280 drainage lakes. The

horizontal spaces show the amount of free carbon dioxide by 0.5 mg/1 intervals and

the vertical scale indicates the percentage of lakes in each of the groups. The dia¬

gram is based on results given in Tables I and II.

The free carbon dioxide content of the surface waters of the

seepage lakes ranged from a minimum of 0.3 mg/1 in Clear Bass

Lake to a maximum mean of 7.0 mg/1 in the Cardinal Bog lake-

let. Table I shows that the maximum number of lakes falls in

the 1.5-1. 9 mg/1 carbon dioxide group, namely 79 or 33 per cent

of the 238 seepage lakes. The 1. 0-1.4 mg/1 group is second with

61 lakes and the 2.0-24 group is third with 45. These three

groups, therefore, include approximately 78 per cent of the total

number of seepage lakes. The large percentage of lakes in these

three groups is to be expected, however, since water with rela¬

tively small amounts of salts in solution usually contains 1.0 to

2.0 mg/1 of free carbon dioxide when freely exposed to the air;

Juday, Birge & Meloche — Lake Waters of N. E. Wisconsin 9

this quantity is subject to modification in lake waters, however,

through the processes of photosynthesis, respiration and decom¬

position. The percentage distribution of the lakes in the various

groups is shown in Fig. 1.

The two seepage lakes with the minimum amounts of free car¬

bon dioxide were populated with considerable numbers of phy¬

toplankton organisms; one yielded 1,470 and the other 4,200

cells and colonies per cubic centimeter of water. The photosyn¬

thetic activities of these organisms had used up the greater part

of the free carbon dioxide.

Table I shows 11 seepage lakes with 4.0 mg/1 or more of

free carbon dioxide and 6 of them have 5.0 mg/1 or more. All

of these lakes have more or less definite bog characteristics ; the

waters (of most of them are highly colored, which indicates the

presence of correspondingly large amounts of organic matter.

The two with maximum amounts of free carbon dioxide are

only small bodies of open water with wide margins of bog de¬

posits.

The results on bog waters that are presented in this report,

are to be regarded only as incidental to the general lake survey.

Many bog ponds and lakelets are found in this district, but no

careful study of them has been attempted because this would

have interfered with the work on lakes. In order to obtain some

idea of the chemico-biologi cal status of the waters of these bog

ponds and lakelets, however, a few of the more accessible ones

were visited and some of the results obtained on them are dis¬

cussed in this connection.

The largest amount of free carbon dioxide was found in the

Forestry Bog pond, namely 10.7 mg/1 and the second in rank was

the Cardinal Bog with 10.0 mg/1 at the surface. The open water

in the former has an area of 990 sq. m. and a maximum depth of

2 m., while that of the latter has an area of 430 sq. m. and a

maximum depth of 5 m. Decomposition is very active in these

bog ponds and, as a result, the water receives a large supply of

free carbon dioxide. On August 21, 1933, for example, a sample

of water from a depth of 4.5 m. in the Cardinal Bog yielded 51.8

mg/1 of free carbon dioxide, with 7.9 mg/1 at the surface.

Drainage lakes . The mean quantities of free carbon dioxide

in the 280 drainage lakes are given in Table II ; the percentage

10 Wisconsin Academy of Sciences , Arts, and Letters .

of these lakes that fall in the different groups is shown graph¬

ically in Figure 1. The waters of 36 of the drainage lakes gave

a mean alkaline reaction to phenolphthalein, so that the group is

distributed through a wider range than the seepage lakes; this

fact is well illustrated in Figure 1.

Since the drainage lakes are distributed through a larger

number of groups, the percentage in the maximum group is

smaller than that in the seepage lakes; a maximum of 21 per

cent is found in the 1. 0-1.4 group of drainage lakes, while a

maximum of 33 per cent of the seepage lakes falls in the 1.5-1. 9

mg/1 group. (Tables I and II). In the drainage lakes, the 1.5-

1.9 group ranks second with 15 per cent and the 2. 0-2-4 mg/1

group third with almost 13 per cent. These three groups contain

137 drainage lakes, or about 49 per cent of the total number.

Adding this number to the 30 lakes of the 0.5-0.9 mg/1 group

gives a total of 167 lakes in these four groups, which is almost

60 per cent of the total number of drainage lakes. In compar¬

ison with this, there are 185 seepage lakes, or 78 per cent of the

total number, in the three groups between 1.0 and 2-4 mg/1. The

two types of lakes are alike in that their maximum numbers fall

within approximately the same range; that is, between 0.5 and

2.4 mg/1 of free carbon dioxide, which is generally about the

quantity of free carbon dioxide absorbed by water when freely

exposed to the atmosphere.

Thirty-six of the drainage lakes, or almost 13 per cent of the

total number, gave a mean alkaline reaction to phenolphthalein.

The quantity of monocarbonate carbon dioxide in them ranged

from a minimum of — 0.1 mg/1 to a maximum mean of — 4.3

mg/1. Table III includes the 7 drainage lakes in which the

largest amounts of monocarbonate carbon dioxide were found ; in

other words, they represent the maximum deficiencies of half¬

bound carbon dioxide. The maximum for a single sample was

obtained at the surface of Mann Lake on July 7, 1928, namely

— 10.7 mg/1 ; the next in rank was — 6.0 mg/1 in Allen of Brazell

Lake. The hydrogen ion of these 7 samples ranged from pH 8.2

to 9.1 and the bound carbon dioxide content from 14.5 to 25.6

mg/1. In comparison with the drainage lakes, the surface waters

of all of the seepage lakes gave a mean acid reaction to phen¬

olphthalein.

Juday , Birge & Meloche — Lake Waters of N. E. Wisconsin 11

The other drainage lakes, 244 in number, gave a mean acid

reaction to phenolphthalein, ranging from a minimum of 0.1

mg/1 to a maximum of 7.0 mg/1 of free carbon dioxide. Of this

number, the mean carbon dioxide content of 186 did not exceed

2.4 mg/1, while 58, or 20 per cent of the total number of drain¬

age lakes, exceeded this amount. Only 37 seepage lakes, or 15

per cent of the total number, yielded means of 2.5 mg/1 or

more of free carbon dioxide. In the drainage type, 13 lakes

had 5.0 mg/1 or more as compared with 6 of the seepage type.

The drainage lakes possessing this large amount of free carbon

dioxide are similar to those of the seepage class in that they be¬

long chiefly to the bog type and most of them have highly col¬

ored waters. In 9 of 13 drainage lakes with high free carbon

dioxide, the color readings were well above 100 on the platinum-

cobalt scale. The last column in Table IV shows that the waters

of these 13 drainage lakes contained rather large amounts of

organic carbon, thus indicating that they possessed correspond¬

ing amounts of organic matter ; the decomposition of this organic

matter would tend to increase the free carbon dioxide content of

the water.

Variations in Free Carbon Dioxide

Observations were not made frequently enough on any one

lake to show the seasonal or annual variations in the free carbon

dioxide content of the surface water; in all lakes except Trout,

the number of determinations was limited to one to three each

year during the period of the general chemical survey; that is,

from 1925 to 1930, inclusive. Certain quantitative differences

were noted in some of the lakes in different years, however.

The maximum difference in free carbon dioxide content of

the surface water was found in Mann Lake where the quantity

ranged from —10.7 mg/1 on July 7, 1928 to 4.0 mg/1 on August

14, 1929, thus making a net difference of 14.7 mg/1 of carbon

dioxide. Carroll Lake showed the next highest difference which

ranged from —3.0 mg/1 to 2.7 mg/1, or a total of 5.7 mg/1.

Upper Gresham was third with a maximum difference of 5.1

mg/1 ; 7 other lakes showed annual differences of 4.0 to 5.0 mg/1.

The largest number of free carbon dioxide determinations

was made on the surface water of Trout Lake, namely 26; the

quantity of free carbon dioxide varied from — 0.9 mg/1 to 2.8,

12 Wisconsin Academy of Sciences, Arts, and Letters .

a difference of 3.7 mg/1. The mean quantity of free carbon

dioxide in the 26 samples was 1.4 mg/1; only two of them gave

an alkaline reaction to phenolphthalein.

In the lakes and bog lakelets whose surface waters always

gave an acid reaction to phenolphthalein, the maximum differ¬

ence in free carbon dioxide was found in the Forestry Bog where

the amount varied from 4.7 to 10.7 mg/1 in four sets of obser¬

vations. The Cardinal Bog was second; the quantity varied

from 5-8 to 10.0 mg/1 in the five surface samples obtained from

it.

In the larger bodies of water belonging to this group, and

especially in those without any bog characteristics, the varia¬

tions in the quantity of free carbon dioxide from year to year

were not so marked; the mean difference for 163 lakes which

were visited more than once was 1.2 mg/1. These differences

ranged from zero to 4.5 mg/1 ; it exceeded 2.0 mg/1 in only 24

of the 163 lakes.

Vertical Distribution of Free Carbon Dioxide

The vertical distribution of the free carbon dioxide is sub¬

stantially the same in the seepage and drainage lakes, so that

the two types need not be separated for this discussion. Both

classes are represented in the diagrams (Figs. 2-15 and 17-27)

and the type to which each belongs is indicated in the explana¬

tions of the figures.

In lakes that are shallow enough for the wind to keep the

water in complete circulation during the summer, the free carbon

dioxide, as well as the other dissolved substances, is uniformly

distributed from surface to bottom. This kind of distribution

is shown in Bear Lake (Fig. 2). While there was a slight de¬

crease in the temperature of the water at the bottom, there was

no change in the dissolved gases 1 m. above the bottom. A slight

stratification was noted in the deepest part of Dorothy Dunn

Lake (Fig. 3) and this was accompanied by an increase in the

quantity of free carbon dioxide near the bottom. Lake Laura

shows a similar situation (Fig. 4) and somewhat larger amounts

of free carbon dioxide were noted in the lower water of Finley

and Weber lakes (Figs- 5, 6) and still larger quantities in Alle-

quash, Blue and Bragonier (Figs. 7, 8, 9) ; the latter three had

9.0, 12.0 and 14.2 mg/1, respectively, in the deeper water. The

maximum depths in these 8 lakes range from 6 to 13 m.

Juday, Birge & Meloche—Lake Waters of N. E. Wisconsin 13

Fig. 2. Vertical distribution of the free (F) and bound (B) carbon dioxide, pH

and temperature (T) in Bear Lake, July 25, 1925. A seepage lake.

Fig. 3. Vertical distribution of the free (F) and bound (B) carbon dioxide, pH

and temperature (T) in Dorothy Dunn Lake, August 28, 1926. A drainage lake.

Fig. 4. Vertical distribution of the free (F) and bound (B) carbon dioxide, pH

and temperature (T) in Lake Laura, August 9, 1926. A seepage lake.

A rather large variation in the free carbon dioxide content

of the lower water was found in lakes ranging from 15 to 20 m.

in depth. Big Carr and Crystal lakes (Figs. 10, 11) belong to

the seepage type and they show comparatively small increases

in free carbon dioxide in the lower water; the amount was 5.6

mg/1 in the former and 5.0 mg/1 in the latter in the deepest

14 Wisconsin Academy of Sciences , Arts , and Letters.

samples. In Muskellunge, Papoose and Two Sisters lakes (Figs.

12, 13, 14), there was a more marked increase of free carbon

dioxide in the thermocline and hypolimnion ; a maximum of 21.5

mg/1 was found in the sample taken 1 m. above the bottom in

Muskellunge Lake on August 25, 1932, while the deepest sample

(18 m.) in Papoose Lake yielded 9.0 mg/1 on August 2, 1928-

Nebish Lake, with a maximum depth of 15.8 m., yielded 21.6

mg/1 of free carbon dioxide at 14 m. on August 23, 1932 (Fig.

15).

Fig. 5. Vertical distribution of the free (F) and bound (B) carbon dioxide, pH

and temperature (T) in Finley Lake, July 28, 1927. A seepage lake.

Fig. 6. Vertical distribution of the free (F) and bound (B) carbon dioxide,

pH and temperature (T) in Bragonier Lake, July 28, 1927. A seepage lake with

brown colored water. Compare with fig. 5.

Anderson, Pallette and Silver lakes (Figs. 17, 18, 19) showed

a different type of vertical distribution of free carbon dioxide;

there was a smaller amount in the thermocline than either above

or below this stratum. These decreases in the carbon dioxide

content of the water in the thermocline were due to the photo¬

synthetic activities of the phytoplankton of this stratum. The

maximum consumption of carbon dioxide in the thermocline

Juday, Birge & Meloche — Lake Waters of N. E. Wisconsin 15

was found in Pallette Lake on August 22, 1928 (Fig. 18) ; on

this date all of the free and 2.6 mg/1 of the half-bound carbon

dioxide at 8 m. had been used by the phytoplankton in the pro¬

cess of assimilation. Similar but smaller losses of half-bound

carbon dioxide were observed in the thermocline of Pallette

Lake on August 17, 1927 and on July 21, 1931. A maximum loss

of — 1.7 mg/1 of half-bound carbon dioxide was found at 7 m. in

Anderson Lake on August 8, 1929 and of — 1.2 mg/at 9 m. in

Silver Lake on July 25, 1931. These decreases in the carbon

dioxide content of the water of the thermocline were correlated

with increases in dissolved oxygen and with higher pH values

as shown in the diagrams. Attention may also be called to the

marked rise in the free carbon dioxide content of the lower

water in these three lakes ; a maximum of 27.5 mg/1 was found

at 18 m. in Silver Lake.

Figures 20 to 27 show the vertical distribution of the free

carbon dioxide in some of the lakes that are more than 20 m.

deep. In all of them except Presque Isle Lake, the water of the

epilimnion contained free carbon dioxide; the amount ranged

from 1.0 to 2.3 mg/1. In Presque Isle Lake, the upper water

was alkaline to phenolphthalein ; the maximum loss of half¬

bound carbon dioxide was equivalent to —1.0 mg/1 at a depth

of 5 m. In all of these lakes, including Presque Isle, there was a

more or less marked increase in the quantity of free carbon

dioxide in the lower water ; the amount ranged from 7.8 mg/1 in

the bottom water of Fence Lake (Fig. 24) to 42-5 mg/1 in that

of Lake Mary (Fig. 25).

BOUND CARBON DIOXIDE

Some 2,800 methyl orange titrations were made during this

investigation for the purpose of determining the bound carbon

dioxide content of the various waters. Of this number, 1,200

were surface samples and 1,600 came from other depths. These

titrations showed that the 518 lakes and lakelets covered a wide

range in bound carbon dioxide content; the quantity in the sur¬

face waters ranged from a minimum of 0.2 mg/1 in two lakes to

a maximum of 39.0 mg/1 in a small spring-fed lakelet.

The seepage and drainage lakes showed marked differences

in their bound carbon dioxide content in most cases and the

16 Wisconsin Academy of Sciences , Arts , and Letters.

■ ■ : \'>Vv -\\v\ V\ / ' xn\ WVx'n \ 'OV^^CVW' x

various bodies of water have been separated into these two

classes for purposes of comparison. Both types have been fur¬

ther separated into groups by one milligram intervals on the

basis of the bound carbon dioxide content of the surface waters.

The results of these groupings are given in Tables V and VII.

-5 T-COz 0 5 10 15 20 25

pH 7.0 7.5 8.0 8.5

Fig. 7. Vertical distribution of the free (F) and bound (B) carbon dioxide, pH

and temperature (T) in Allequash Lake, August 15, 1926. A drainage lake.

Seepage Lakes

Surface waters. The mean quantity of bound carbon dioxide

in the surface waters of the seepage lakes is given for the various

groups in Table V. This table includes results obtained from

238 lakes; of this number, 132 were single determinations on

each lake, while from 2 to 17 surface samples were secured from

each of the other lakes. The largest number (17) was taken in

Weber Lake during the period from 1925 to 1932 inclusive ; 14

surface determinations each were made on Crystal and Muskel-

lunge lakes during the same period of time. In compiling

Table V, the mean of the various analyses was used in grouping

the 106 lakes visited more than once.

In a previous report (Juday and Birge 1933), it was pointed

out that many of these seepage lakes have very low specific con¬

ductances, which indicates that their waters contain correspond¬

ingly small amounts of electrolytes. In view of this fact, it is

not surprising that 29 of them, or a little more than 12 per cent

of the number belonging to this class, yielded less than 1.0 mg/1

of bound carbon dioxide, while 106, or more than 44 per cent of

the total, yielded between 1.0 and 1.9 mg/1 ; that is, the surface

waters of 135 of these seepage lakes, which is more than 56 per

cent of the total number, did not contain more than 1.9 mg/1

of bound carbon dioxide. The table also shows that only 16 of

Juday, Birge & Meloche — Lake Waters of N. E. Wisconsin 17

the seepage lakes (6.7 per cent of the total) yielded 5.0 mg/1 or

m;ore and none of them exceeded a mean of 11.3 mg/1.

Their distribution into the various groups is shown graph¬

ically in Figure 16, where they are grouped by 2.0 mg/1 inter¬

vals. This diagram serves to bring out more clearly the fact

that only a very small percentage of the seepage lakes yielded as

much as 4.0 mg/1 of bound carbon dioxide or more; nearly 89

per cent of them had less than 4.0 mg/1 and more than 56 per

cent of them had less than 2.0 mg/1.

0 T-C02 5 ^ 10 15 20

pH 5.5 6.0

Fig. 8. Vertical distribution of the free (F) and bound (B) carbon dioxide, pH

and temperature (T) in Weber Lake, August 29, 1928. A seepage lake. Compare

with figs. S and 9.

The small amount of bound carbon dioxide found in the waters

of a large percentage of the seepage lakes is due in part to the

scarcity of calcium and magnesium in the glacial deposits of

the region and in part to the isolation of the lake waters from

the ground waters. While the glacial debris contained only a

relatively small amount of calcareous material at the time of its

deposition, the quantity of it has been reduced still further by

the process of leaching; this is true especially of the upper or

soil stratum. In some localities, for example, it would require

from one to more than 5 metric tons of lime per hectare to cor¬

rect the soil acidity for agricultural purposes. Thus the limited

amount of water that drains into these lakes from the adjacent

18 Wisconsin Academy of Sciences, Arts, and Letters .

land contains only small amounts of carbonate substances in solu¬

tion.

A maximum mean of 11.3 mg/1 of bound carbon dioxide was

found in Forest Lake, which indicates that it lies in an area that

is more abundantly supplied with calcareous material than most

of the other seepage lakes. The next highest mean was 10.8

mg/1 in Lake Laura, while a single sample from the surface of

Sandy Beach Lake also yielded 10.8 mg/1. None of the surface

samples from the other seepage lakes contained as much as 10.0

mg/1 of bound carbon dioxide and only 4 of them fell between

9.0 and 10.0 mg/1.

0 T-CO, 5 10 15 20 25

r«_ - i - - ~-rQ - - - - - - -

pH 6.5 7.0 7.5

Fig. 9. Vertical distribution of the free (F) and bound (B) carbon dioxide, pH

and temperature (T) in Blue Lake, August 6, 1929. A seepage lake. Compare with

fig. 8.

Variations. Annual variations were noted in the bound

carbon dioxide content of the surface waters of most of the

seepage lakes that were visited more than once. A fourfold dif¬

ference was found in Crystal Lake; the quantity ranged from

0.5 mg/1 on July 14, 1931 to 2.0 mg/1 on June 26, 1926, in 12

different samples. In 5 lakes the differences were three-fold

and in 20 others they were twofold or more, but less than three¬

fold. While these percentile differences seem large, the actual

quantitative differences were comparatively small. In all of

these high percentage cases, the quantitative difference exceeded

2.0 mg/1 of bound carbon dioxide in only one lake; it did not

exceed 1.0 mg/1 in 8 lakes, so that the differences fell between

1.0 and 2.0 mg/1 in 17 of the high percentage lakes. Samples ob-

Juday, Birge & Meloche — Lake Waters of N . E . Wisconsin 19

tained from 4 of the seepage lakes in different years yielded the

same quantity of this gas in both years.

In the 106 seepage lakes that were visited more than once,

the variation in the bound carbon dioxide content of the surface

water ranged from zero in these 4 lakes to a maximum of 8.5

mg/1 in Lake Laura; in the latter a minimum of 9.8 mg/1

was found on August 19, 1927 and a maximum of 12.8 mg/1 on

August 28, 1929, so that the percentile difference was only a

little more than 27 per cent of the maximum quantity. The

range of variation in the various lakes is given in Table VI. In

71 of the 106 lakes, the annual differences in bound carbon diox¬

ide content of the surface waters did not exceed 1.0 mg/1 and it

exceeded 2.0 mg/1 in only 6 of them. In the great majority of

the seepage lakes, therefore, these annual differences may be

regarded as small from a quantitative standpoint. Results for

a number of seepage lakes are given in Table XII.

Drainage Lakes

Surface waters . The drainage lakes fall into two general

classes, namely, those which are not fed by permanent streams

or springs and those which receive water from these sources-

The drainage lakes which do not have streams or springs flowing

0 T-C02 5 10 15 20 25

- - ..-.-.I -I. - - - .■ i I mi. I I ......

-J _ lY-1 _ I _ L

pH 5.0 5.5 6.0

Fig. 10. Vertical distribution of free (F) and bound (B) carbon dioxide, pH

and temperature in Big Carr Lake, July 12, 1929. A seepage lake.

20 Wisconsin Academy of Sciences , Arts, and Letters.

into them possess certain seepage characteristics in that they

have rather limited drainage basins and their waters contain

relatively small amounts of electrolytes in solution. They make

up the groups of drainage lakes which have the smallest amounts

of bound carbon dioxide. Since they are the sources of small,

more or less intermittent streams, however, they are classified as

drainage lakes.

In the drainage lakes that are fed by springs and streams,

the effect of the inflowing water upon the bound carbon dioxide

content of the lake water depends chiefly upon the quantity of

this gas in the former and upon the volume of the inflowing

water in comparison with that of the lake. A large volume of

stream or spring water flowing into a small lake will have a

marked influence on the chemical status of the lake water, while

the reverse would minimize the effect of the inflowing water.

In general then, the lakes that are traversed by the larger

streams, such as the Manitowish River, are affected to a greater

degree by the inflowing water than those that lie in the courses

of the smaller streams. In lakes of equal volume, the one re¬

ceiving a large amount of spring water would be affected more

than the one receiving a relatively small amount.

As a result of the modifications produced by the inflowing

water as well as by other factors, the bound carbon dioxide con¬

tent of the surface waters of the drainage lakes covers a much

wider range than that of the seepage lakes. The quantity in the

former ranged from a minimum of 1.0 mg/1 in Clear Lake

(Langlade County) to a maximum of 39.0 mg/1 in the Inkpot, a

small spring-fed lakelet lying a short distance north of Ander¬

son and Black Oak lakes. Twin Island Lake was second highest

with 31.5 mg/1 and Wild Cat was third with a mean of 30.5

mg/1. More than 60 per cent of the drainage lakes had a larger

amount of bound carbon dioxide than the maximum quantity

(11.3 mg/1) found in the seepage lakes.

The 280 drainage lakes have been separated into groups on

the basis of the bound carbon dioxide content of their surface

waters. These groupings are given in Table VII and the results

are shown graphically in Figure 16. As indicated above, the

drainage lakes with seepage characteristics are found in the

groups having the smallest amounts of bound carbon dioxide.

Juday, Birge & Meloche - — Lake Waters of N. E. Wisconsin 21

None of these lakes had less than 1.0 mg/1, but 7 of them fell

in the 1. 0-1.9 mg/1 group and 13 in the 2. 0-2. 9 mg/1 group. In

comparison with this, 186 seepage lakes, or 78 per cent of the

total number in this class, did not exceed 2.9 mg/1, while only

20 drainage lakes, or 7 per cent of the total number, yielded

such small amounts of bound carbon dioxide. The differences

in the groupings of the two classes of lakes are shown in Figure

16, where the distribution of the seepage lakes is shown in the

upper part of the diagram and the wider range of the drainage

lakes in the lower part. The figure shows a maximum of 48

lakes, or a little more than 17 per cent of the total number, in

the 14.0-15.9 mg/1 group of drainage lakes as compared with

a maximum of 135 lakes, or more than 56 per cent of the total

number, in the 0.0-1.9 mg/1 group of the seepage type. The 4

drainage lakes with 28.0 mg/1 of bound carbon dioxide or more

have been omitted from the diagram, but they are represented in

Table VII; they ranged from 28.0 mg/1 in Post Lake to 39-0

mg/1 in the Inkpot.

Fig. 11. Vertical distribution of free (F) and bound (B) carbon dioxide, pH

and temperature (T) in Crystal Lake, August 11, 1932. A seepage lake with high

transparency and unstained water. Note that the quantity of bound carbon dioxide

is smaller than that of the free. Very soft water.

Variations . A summary of the variations in the bound car¬

bon dioxide content of the surface waters of the drainage lakes

that were visited more than once is given in Table VIII. In 29

22 Wisconsin Academy of Sciences, Arts, and Letters.

of the 136 lakes included in the table, the maximum annual

differences did not exceed 1.0 mg/1 and in 47 others they fell

between 1.0 and 2.0 mg/1, so that in 76 of these lakes, or ap¬

proximately 56 per cent of the number in this group, the differ¬

ences did not exceed 2.0 mg/1. In comparison with this, 100

seepage lakes, or a little more than 94 per cent of the 106 that

were visited more than once, fall in this same category.

Fig. 12. Vertical distribution of the free (F) and bound (B) carbon dioxide,

pH and temperature (T) in Muskellunge Lake, August 25, 1932. A seepage lake.

In 28 of the drainage lakes, the difference exceeded 3.0 mg/1,

while only one of the seepage lakes exceeded this amount; the

seepage lakes, therefore, as might be expected from their smaller

bound carbon dioxide content, showed smaller quantitative an¬

nual variations than the drainage lakes, but their percentage

differences were larger in many instances. The maximum per¬

centile difference in the drainage lakes was found in Lake George

and in Lake Helen ; the waters of both of these lakes contained

relatively small amounts of bound carbon dioxide. In 3 obser¬

vations on Lake George, the surface water yielded a minimum of

2.0 mg/1 on July 10, 1927 and on August 4, 1930 and a maximum

of 4.0 mg/1 on August 24, 1929. In 5 surface samples from

Lake Helen, the bound carbon dioxide content varied from a

minimum of 2.5 mg/1 on July 17, 1928 and on August 4, 1930 to

a maximum of 5-0 mg/1 on August 18, 1929, so that there was a

Juday, Birge & Meloche — Lake Waters of N. E. Wisconsin 23

twofold variation in both of these lakes. In Harvey and Sand

lakes, the same readings were obtained in the two different

years that they were visited.

The maximum quantitative difference, 8.5 mg/1, was found

in Mann Lake ; the bound carbon dioxide content of the surface

water of this lake was 30.5 mg/1 on August 14, 1929 and 22.0

mg/1 on May 5, 1927. Considering the 5 July and August de¬

terminations, however, the difference was only 5.5 mg/1, since

the smallest summer reading was 25.0 mg/1. The second largest

difference was 6.9 mg/1, which was noted in the surface water

of Big Lake.

Fig. 13. Vertical distribution of the free (F) and bound (B) carbon dioxide,

pH and temperature (T) in Papoose Lake, August 2, 1928. A drainage lake. Com¬

pare with fig. 12.

Vertical Distribution

The vertical distribution of the bound carbon dioxide is given

for a number of lakes in Table XII, and the results for some of

them are shown in Figures 2-15 and 17-27. The character of

the vertical distribution is dependent to a large extent upon the

thermal stratification of the water, which in turn depends chiefly

upon the depth and area of the lake. In shallow lakes where the

entire body of water is kept in circulation during the summer,

the bound carbon dioxide is uniformly distributed from surface

24 Wisconsin Academy of Sciences , Arts, and Letters .

to bottom ; where the water is deep enough to become thermally

stratified, there may be more or less marked differences between

the bound carbon dioxide content of the epiliminion and of the

hypolimnion, but in lakes with a small bound carbon dioxide con¬

tent the distribution is usually about uniform from surface to

bottom even when the water is stratified. Owing to the small

quantity of bound carbon dioxide generally present in the waters

of the seepage lakes, the differences in vertical distribution are

not as marked as in the drainage lakes ; therefore, the two groups

are discussed separately.

Seepage lakes . As already indicated, the surface-bottom dif¬

ferences in bound carbon dioxide content are not so great in the

seepage lakes. A uniform distribution is shown by Bear Lake

(Fig. 2) which is shallow" enough to be kept in practically com¬

plete circulation in summer. A uniform distribution is shown

also by Crystal Lake (Fig. 11) which is deep enough to be therm¬

ally stratified. In Weber Lake (Fig. 8), there was a slightly

larger amount of bound carbon dioxide in the lower than in

the upper water. Similar results are shown for Big Carr, Blue,

Finley, Laura and Pallette lakes (Figs. 10, 9, 5, 4, 18).

The maximum difference between upper and lower strata was

noted in Bragonier and in Muskellunge lakes (Figs. 6, 12). In

the former lake, the surface Water yielded 2.5 mg/1 and the 8 m.

sample (0.5 m. above the bottom) 7.0 mg/1. In Muskellunge

Lake the surface water contained 10.0 mg/1 and that at 20.3

m- yielded 14.5 mg/1 of bound carbon dioxide. Anderson Lake

showed the next largest difference with 12.5 mg/1 at the surface

and 16.5 mg/1 at 19 m. ; Ike Walton was third with a difference

of 3.3 mg/1. Two other seepage lakes, namely Clear (Oneida

County) and Little John Jr., showed a difference of 2.0 mg/1 be¬

tween surface and bottom, but in 23 others the differences were

all less than 2.0 mg/1; in 12 of the latter, in fact, they were

less than 1.0 mg/1.

Drainage lakes . As might well be expected, the larger bound

carbon dioxide content of the great majority of the drainage

lakes gave rise to more marked differences between the quantity

of this gas in the epilimnion and that in the hypolimnion. In

33 drainage lakes on which series of samples were taken, the

maximum difference was found in Wild Cat Lake; on August

Juday, Birge & Meloche — Lake Waters of N. E. Wisconsin 25

0 T-CO; 5 10 15 _ 20 25

pH 6.5 7.0 7.5

Fig. 14. Vertical distribution of the free (F) and bound (B) carbon dioxide,

pH and temperature in Two Sisters Lake, August 13, 1929. A drainage lake. Com¬

pare with figs. 12 and 13

24, 1926 the surface water of this lake yielded 30.8 mg/1 and

that at 11 m. (1 m. above the bottom) contained 44.1 mg/1 of

bound carbon dioxide, a difference of 13.3 mg/1. In 5 other

series from this lake the differences between surface and bottom

ranged from 4.6 to 11.5 mg/1. The next largest difference was

noted in Nebish Lake; the bound carbon dioxide content of the

surface water on August 29, 1931 was 3.5 mg/1 and that of the

14 m. sample was 13.0 mg/1, a difference of 9.5 mg/1. A series

in which the bound carbon dioxide of this lake ranged from 4.0

mg/1 at the surface to 8.0 mg/1 at the bottom is shown in

Figure 15.

A surface-bottom difference of 9.2 mg/1 was noted in the

bound carbon dioxide of Lake Mary on July 12, 1926. A series

taken in this lake on July 11, 1928 is represented in Fig. 25;

in this instance the surface water yielded 3.7 mg/1 and that at

20 m- 11.5 mg/1. The surface water of Upper Gresham Lake

contained 18.3 mg/1 and that at 7 m. (1 m» above the bottom)

25.6 mg/1, a difference of 7.3 mg/1. In the other 29 drainage

lakes in which series were taken, the difference between the

bound carbon dioxide content of the surface and the bottom

strata ranged from zero in Stone Lake at Crandon to a maxi¬

mum of 6.5 mg/1 in Silver Lake; the difference was less than

26 Wisconsin Academy of Sciences, Arts, and Letters .

2.0 mg/1 in 11 of these lakes and did not exceed 1.0 mg/1 in 4

of them. Some of the moderate differences between the bound

carbon dioxide content of the upper and lower strata are shown

in the diagrams representing Adelaide, Black Oak, Crawling

Stone, Dead Pike, Fence, Presque Isle, and Trout (Figs. 20-26) ;

Lake Mary showed the largest difference represented in the dia¬

grams.

While the photosynthetic activities of the phytoplankton gave

the water of the thermocline an alkaline reaction to phenol-

phthalein in Anderson, Pallette and Silver lakes (Figs. 17, 19),

there was no decrease in the bound carbon dioxide of these

strata ; this indicates that the quantity of monocarbonate which

gave the water an alkaline reaction to phenolphthalein was not

large enough to cause a precipitation of the calcium. Such de¬

creases have been found in other Wisconsin lakes, however, where

the water contained a larger amount of bound carbon dioxide

than is found in the northeastern lakes.

Fig. IS. Vertical distribution of the free (F) and bound (B) carbon dioxide,

pH and temperature (T) in Nebish Lake, August 23, 1932. A drainage lake. Com¬

pare with fig. 14.

This phenomenon was observed in Beasley and Knights lakes,

for example, which are situated in central Wisconsin. (Birge

and Juday 1911). In Beasley Lake on August 3, 1908, the bound

carbon dioxide content of the surface water was 73.8 mg/1, that

at 5.5 m. was 63.8 mg/1 and that at 14 m. was 85.6 mg/1, so that

the amount at 5.5 m. was 10.0 mg/1 less than that at the sur¬

face. The smaller quantity at 5.5 m. was correlated with a

Juday, Birge & Meloche — Lake Waters of N. E . Wisconsin 27

marked alkaline reaction of the water to phenolphthalein and

with an excess of dissolved oxygen amounting to 20.3 mg/1, or

212 per cent of saturation. There was also a smaller amount of

calcium at 5.5 m. ; it decreased from 36.2 mg/1 at 4 m. to 30.0

mg/1 at 5.5 m. and then rose to 38.3 mg/1 at 7 m. Through the

removal of half-bound carbon dioxide by the phytoplankton at

5.5 m., the normal calcium carbonate exceeded the saturation

point and some of it was precipitated and sank into the lower

water.

In Knights Lake on August 25, 1909, the bound carbon diox¬

ide amounted to 82.4 mg/1 at the surface, 71.8 mg/1 at 4 m.

and 104.2 mg/1 at 11 m., so that the quantity at 4 m. was 10.6

mg/1 less than that at the surface ; the dissolved oxygen at 4 m-

amounted to 31.1 mg/1, or 328 per cent of saturation.

Bound Carbon Dioxide Content of Other Lakes

In 1911 Birge and Juday published results on the bound car¬

bon dioxide content of 151 Wisconsin lakes; 58 of these lakes

belong to the northeastern group, 63 are situated in the north¬

western quarter of the state and 30 in the southeastern quarter.

Those belonging to the northeastern group are included in the

present investigation. In the 63 lakes of the northwestern group,

the bound carbon dioxide content of the surface water ranged

from a minimum of 0.8 to a maximum of 43.1 mg/1 ; in 14 of

them the quantity did not exceed 5.0 mg/1, while 4 yielded 40.0

mg/1 or more. Calcareous material is more plentiful in the

southeastern quarter of the state, so that the lake waters of this

section contain a larger amount of bound carbon dioxide. With

one exception, the quantity ranged from 59.4 to 93.0 mg/1 in

these lakes. The exception is Devils Lake which occupies a

quartzite basin ; its water yielded only 6.3 mg/1.

In 10 of the Finger lakes of New York, the quantity varied

from 13-7 mg/1 in Canadice Lake to 47.6 mg/1 in Canandaigua ;

Canadice was the only one, however, that fell below 25.0 mg/1.

Seneca and Cayuga lakes, the largest and deepest members of

this group, yielded 43.6 mg/1. (Birge and Juday 1914).

Kemmerer, Bovard and Boorman (1923) found that the

bound carbon dioxide content of 53 lakes situated in the north¬

western part of the United States ranged from a minimum of

28 Wisconsin Academy of Sciences , Arts , and Letters.

2.5 mg/1 to a maximum of 947.6 mg/1. The waters of 21 yielded

less than 10.0 mg/1 and only 4 more than 60.0 mg/1. The deep¬

est one, Crater Lake, contained 12.5 mg/1 and the next deepest,

Lake Tahoe, 17.5 mg/1.

In some 70 Minnesota lakes, Johnson (1933) found that the

bound carbon dioxide varied from 10.7 to 90-0 mg/1 ; only 3 of

them yielded less than 25.0 mg/1.

Fig. 16. This diagram shows the grouping of the seepage and drainage lakes

on the basis of the bound carbon dioxide content of their surface waters. The

vertical spaces represent the percentage of lakes in each group and the horizontal

scale shows the amount of bound carbon dioxide. The diagram is based on re¬

sults given in Tables V and VII. Compare with fig. 1.

In comparison with the above results, Table V shows that

222 seepage lakes of northeastern Wisconsin had less than 5.0

mg/1 of bound carbon dioxide; this is a little nfore than 93 per

cent of the seepage lakes that were visited. The table also shows

that 29 of them yielded less than 1.0 mg/1 and gave a mean of

0.7 mg/1. In the drainage group (Table VII) , the surface waters

of 33 lakes, or almost 12 per cent of the total number, yielded

Juday, Birge & Meloche — Lake Waters of N. E. Wisconsin 29

less than 5.0 mg/1. One of the most important characteristics

of the waters of these northeastern lakes is the large percentage

of them with very small quantities of bound carbon dioxide.

Bound Carbon Dioxide and Specific Conductance

The relation of the bound carbon dioxide of these lake waters

to their specific conductance has been discussed in a previous

report (Juday and Birge 1933). A definite correlation between

these two factors was found; that is, small amounts of bound

carbon dioxide were correlated with low specific conductances

and large amounts with high conductances. The mean quantity

of bound carbon dioxide rose from 1.4 mg/1 in the 5-9 X 10~6 con¬

ductance group to 26.2 mg/1 in the 100-124 x 10 6 group. Rather

wide variations in the quantity of bound carbon dioxide were

found in the same conductance group, but the general means of

the various conductance groups showed a consistent correlation.

These results confirm what has been noted in other data regard¬

ing these lakes; while there may be a rather wide range in

the correlation between two factors in a single group, the means

of the various groups yield a consistent relation when they rep¬

resent an average of 15 cases or more in each group.

An average of the whole series of observations on the sur¬

face waters of the various lakes indicates that an increase of

approximately 2.3 mg/1 in the bound carbon dioxide content is

correlated with an increase of 10 x 10 6 in specific conductance.

It was noted that relatively small changes in the bound carbon

dioxide from year to year were not accompanied by similar

changes in the specific conductance of the water, but there was

a general shift in the latter in correlation with the more marked

variations in the former. The coefficient of correlation between

bound carbon dioxide and specific conductance was 0.89 in 241

seepage lakes and 0.94 in 292 drainage lakes.

Senior-White (1927) obtained similar results on 35 natural

waters of Ceylon which he investigated. There was a close re¬

lationship between the specific conductance and the total carbon

dioxide content of these waters. While he found considerable in¬

dividual divergencies in some cases, there was a general increase

in conductance with increasing amounts of carbon dioxide and

the coefficient of correlation of the entire series was 0.88.

30 Wisconsin Academy of Sciences , Arts , cmd Letters.

Fig. 17. Vertical distribution of the free (F) and bound (B) carbon dioxide,

pH and temperature (T) in Anderson Lake, August 18, 1929. A seepage lake. Com¬

pare with figs. 14, 18, and 19.

An increase in specific conductance with increasing depth

was noted in the Wisconsin lakes which yielded a larger quantity

of bound carbon dioxide in their lower strata.

Bound Carbon Dioxide and Calcium

Table IX shows the relation between the quantity of bound

carbon dioxide in the surface waters of 357 lakes and the amount

of calcium found in the corresponding samples of water. A large

percentage of these samples yielded only small amounts of cal¬

cium, and they also contained correspondingly small quantities

of bound carbon dioxide. The second group in the table includes

the largest number of lakes, namely 68, while the first group is

second with 60 lakes ; thus the first two groups include 128 lakes,

or a little more than 35 per cent of the number on which calcium

determinations were made. The surface waters of the lakes

belonging to these two groups yielded less than 4.0 mg/1 of

bound carbon dioxide and the mean quantity of calcium in them

was 0-8 and 1.0 mg/1, respectively.

There is a more even distribution of the lakes falling between

the 4.0-5.9 and the 18.0-19.9 mg/1 groups; 203 lakes, or about

57 per cent of those represented in the table, are included in

these groups, with a maximum of 39 lakes in the 14.0-15.9

group. There are only 25 lakes in the last three groups because

Juday, Birge & Meloche—Lake Waters of N. E. Wisconsin 31

Fig. 18. Vertical distribution of the free (F) and bound (B) carbon dioxide,

pH and temptrature (T) in Pallette Lake, August 22, 1928. A seepage lake. Com¬

pare with figs. 17 and 19.

lakes with such large amounts of bound carbon dioxide consti¬

tute a relatively small percentage of the total number found in

the Highland Lake District of Wisconsin.

While there is a wide range between the maximum and mini¬

mum amounts of calcium in the various carbon dioxide groups,

the means for the different groups show a general increase in

the quantity of calcium correlated with an increase in the amount

of bound carbon dioxide. There is a general rise in the mean

quantity of calcium from 0.8 mg/1 in the 0.0-1.9 mg/1 carbon

dioxide group to a maximum mean of 14.8 mg/1 in the 24.0-31.9

group. The table shows a fourteenfold difference between the

maximum and minimum amounts of calcium in the 0. 0-1-9 mg/1

group, a twentyfourfold difference in the second group and a

twentytwofold difference in the 4.0-5.9 mg/1 group. Beyond

the latter group, however, the percentile differences are not so

great; the difference between maximum and minimum in the

6.0-7.9 mg/1 group, for example, is only a little more than three¬

fold, while all of those in the higher groups are less. The small¬

est percentile difference, as well as the smallest quantitative

difference, is found in the 9 lakes belonging to the 20.0-21.9

mg/1 group. In spite of these marked differences between

maxima and minima in the various groups, the analyses yield

consistent means, especially where each group contains 18 lakes

32 Wisconsin Academy of Sciences , Arts , Letters .

0 T-C02 5 10 _ 15 20 25 30

5

10

15

pH 7.0 7.5 8.0 8.5

Fig. 19. Vertical distribution of free (F) and bound (B) carbon dioxide, pH

and temperature (T) in Silver Lake, August 13, 1932. A drainage lake. Com¬

pare with figs. 17 and 18.

or more. In these cases the high results counterbalance the low

ones in such a way as to yield a fair mean for the group.

Figure 28 shows graphically the relation between the bound

carbon dioxide and the calcium as represented in Table IX. The

quantity of calcium is platted at the middle of each bound carbon

dioxide group. The diagram contains curves representing the

maximum and minimum, as well as the mean, quantities of cal¬

cium in the various groups. The middle curve, represented by

a solid line, indicates the mean amounts ; there are three points

in this curve where the gradients are not as steep as they

are in the other groups, but the whole curve shows clearly the

general increase in bound carbon dioxide content with increasing

amounts of calcium.

The upper curve in Figure 28, consisting of a broken line,

connects the points representing the maximum amounts of cal¬

cium in the different groups, while the lower broken line curve

shows the minimum quantities. The irregularities in these two

curves are not very great, except the one for the minimum quan¬

tity of calcium in the 20-0-21.9 mg/1 group; this group has an

unusually large minimum, but a larger number of lakes would

undoubtedly modify this result. The small number of lakes in

the three groups above 18.0-19.9 mg/1 are hardly sufficient to

give fair means, but they match the means of the other groups

as well as could be expected for such a small number of lakes in

each group.

Juday, Birge & Meloche — Lake Waters of N. E. Wisconsin 33

Fig. 20. Vertical distribution of the free (F) and bound (B) carbon dioxide,

pH and temperature (T) in Adelaide Lake, August 5, 1927. A drainage lake with

a small amount of bound carbon dioxide.

CARBON DIOXIDE OF GROUND WATERS

Observations were made on the carbon dioxide content of

149 well waters and of 7 spring waters in this investigation for

the purpose of comparing them with the lake waters. All of the

wells except 9 and all of the springs except one are situated on

the immediate shores of 80 different lakes. Samples were ob¬

tained from only one well on each of 52 lakes, while from 2 to 19

samples were taken from wells and springs located on the shores

of each of the other 28 lakes. The maximum number of well

samples (19) was taken on Trout Lake; Plum Lake was second

with 10 and Muskellunge third with 6. In the other 25 cases,

the well and spring samples numbered only 2 to 4 on each lake.

Nine samples were obtained from wells that were situated half

34 Wisconsin Academy of Sciences , Arts, and Letters .

a kilometer or more from the nearest lake and one spring was

about half a kilometer from any lake.

The depth of 112 wells was ascertained, while that of 37

was unknown to the individuals in charge of them at the time

the samples were taken. The shallowest well was an open one

only 2.5 m. deep, while the deepest one was a drilled well with a

depth of 58.2 m. In 84 cases the depth did not exceed 15 m. and

only 6 were deeper than 30 m. The great majority of them were

driven wells, especially those that did not exceed 15 m- in depth.

Fig. 21. Vertical .distribution of the free (F) and bound (B) carbon dioxide,

pH and temperature (T) in Black Oak Lake, August 24, 1928. A drainage lake.

Compare with figs. 22 and 23.

Free Carbon Dioxide

The water of 3 wells gave an alkaline reaction to phenol ph-

thalein, thus showing the presence of normal carbonate carbon

dioxide ; the latter was equivalent to — 0.5 mg/1 of carbon diox-

Juday, Birge & Melocke—Lake Waters of N . E. Wisconsin 35

Me. Two of these wells are about a kilometer from the nearest

lake, but the third one is on the immediate shore of Beaver Lake.

The results obtained on some of the wells are given in Table XL

All of the other well samples gave an acid reaction to phe-

nolphthalein ; the free carbon dioxide in them ranged from a

minimum of 0.5 mg/1 in the Forestry Headquarters well at

Trout Lake to a maximum of 67.5 mg/1 in the Warner well on

Plum Lake. The second largest amount was noted in the Stover

well located on the shore of Clear Lake (Oneida County),

namely 58.0 mg/1. Only 8 wells, however, yielded more than

40.0 mg/1, but the amount reached 20 mg/1 or more in 49 cases ;

it did not exceed 10.0 mg/1 in 45 of the wells.

With one exception, the waters from wells situated on the

immediate shores of the various lakes yielded a considerably

larger amount of free carbon dioxide than the surface waters of

the corresponding lakes. Both the well water and the surface

water of Beaver Lake gave alkaline reactions to phenolphthalein ;

in both cases the amount of monocarbonate present was equiva¬

lent to —6.5 mg/1 of carbon dioxide. In 11 other instances, the

surface water of the lakes gave an alkaline reaction to phenol¬

phthalein, but all of the well waters of their respective shores

gave an acid reaction to this indicator.

The quantity of free carbon dioxide in the waters of the 7

springs ranged from a minimum of 4.0 mg/1 to a maximum of

41-6 mg/1; in the other 5 springs, it varied from 5.5 to 17.5

mg/1.

The general results indicate that there is no direct correla¬

tion between the free carbon dioxide content of the ground

waters as represented by these wells and springs and that of

the corresponding lake waters. Even the wells situated on the

shores of the same lake show a wide variation in free carbon

dioxide content. This fact is well shown by the results given

in Table XI for Muskellunge, Plum and Trout lakes.

Bound Carbon Dioxide

There was a wide variation in the bound carbon dioxide

content of the 149 well waters ; the minimum was 2.0 mg/1 and

the maximum 166.6 mg/1, which represents a fiftyfold range in

quantity.

86 Wisconsin Academy of Sciences , Arts, and Letters .

Fig. 22, Vertical distribution of the free (F) and bound (B) carbon dioxide,

pH and temperature (T) in Crawling Stone Lake, August 10, 1927. A drainage

lake. Compare with figs. 23 and 24.

The smallest amount (2.0 mg/1) of bound carbon dioxide

was noted in the water from a well at the State Fish Hatchery

near Woodruff ; this well is only 3.3 m. deep. This water yielded

25.0 mg/1 of free carbon dioxide and the hydrogen ion concen¬

tration was pH 5.5. The well is located along the outlet stream

of Madeline Lake and some 200 meters from the shore of this

lake, The surface water of the lake gave an alkaline reaction

to phenolphthalein and it yielded 20.5 mg/1 of bound carbon

dioxide, or more than ten times as much as the well water.

The second smallest amount of bound carbon dioxide was

2.7 mg/1; this quantity was found in the water of a well at

Sisson's Resort on Little St. Germain Lake, in that of a well

at the Pines Resort on Nokomis Lake and in that of Paquette's

well at Boulder Junction which is about half a kilometer from

Juday, Birge & Meloche — Lake Waters of N. E. Wisconsin 37

Little Rice Lake. The surface water of Little St. Germain Lake

yielded five times as much bound carbon dioxide as the well at

Sissons’s Resort, while that of Nokomis Lake was four times as

large as that of the Pines Resort well and that of Little Rice Lake

was four times as large as that of Paquette's well. Four other

well waters yielded only 3.0 mg/1 of bound carbon dioxide and

the amount did not exceed 5.0 mg/1 in 24 of the 149 wells.

In Table XII the wells have been divided into groups on the

basis of their bound carbon dioxide content. The maximum

number of wells (49) falls in the 5.0-9. 9 mg/1 group, whilejhe

10.0-14.9 group is second with 27 wells and the 0.0-4.9 group

is third with 19. Thus the waters of 95 wells, or more than 63

per cent of the 149, yielded less than 15.0 mg/1 of bound carbon

dioxide. This result compares favorably with the surface waters

of the 280 drainage lakes; approximately 59 per cent of them

contained less than 15.0 mg/1 of bound carbon dioxide. The

surface waters of all of the seepage lakes fell below this amount ;

the maximum amount in them was 11.3 mg/1.

Fig. 23. Vertical distribution of the free (F) and bound (B) carbon dioxide,

pH and temperature (T) in Dead Pike Lake, August 9, 1927. A drainage lake.

38 Wisconsin Academy of Sciences , Arts, and Letters.

The largest quantity of bound carbon dioxide (100.0 mg/1)

was observed in the water of the hotel well in the village of

Winegar ; this well is about 150 m. from the shore of Little Horse-

head Lake whose surface water yielded 11.1 mg/1. This well

water also showed the maximum specific conductance, namely

550 xlO'6. The second largest amount of bound carbon dioxide

was noted in a well water at Oxbow Park on Oxbow Lake; it

yielded 82-0 mg/1 and the specific conductance was 800 X 10-6.

These two wells were the only ones in which the quantity of

bound carbon dioxide exceeded 60.0 mg/1 and only 9 of the 149

wells yielded 50.0 mg/1 or more. The two wells with the largest

amounts of bound carbon dioxide and two springs whose waters

contained more than 60-0 mg/1 lie within the borders of the

Winegar Moraine; these results seem to indicate that the mo¬

rainic deposit contains a larger amount of calcerous material

than the outwash deposit in which the majority of the wells are

situated. Two wells located in the outwash plain, however,

yielded 55.5 and 56.0 mg/1 of bound carbon dioxide, respectively.

The results given in Table XI show that there is no correla¬

tion between the bound carbon dioxide content of the well waters

and that of the surface or bottom waters of the lakes on whose

shores they are situated. In some cases the well waters yielded

a larger amount of bound carbon dioxide than the corresponding

lake waters, in other instances it was smaller and in still others

both types of well waters were found on the shores of the same

lake. The first type is represented in Table XI by such lakes

as Adelaide, Anderson (spring) and Clear; the second type

includes Fishtrap, Little St. Germain and Wild Cat. The mixed

type is well illustrated by the wells situated on the shores of

Muskellunge, Plum and Trout lakes.

Two of the well waters from the shores of Muskellunge Lake

had a smaller amount of bound carbon dioxide than the surface

water of the lake and three had a larger quantity. Six of the

wells on Plum Lake had smaller amounts than the surface water

of the lake and three had larger amounts ; the well at Warner’s

Resort yielded approximately twice as much bound carbon diox¬

ide as the surface water of the lake and that at Wolff’s Cot¬

tages contained less than a quarter as much as the surface water

of the lake.

Juday, Birge & Meloche — Lake Waters of N. E. Wisconsin 89

Fig. 24. Vertical distribution of the free (F) and bound (B) carbon dioxide,

pH and temperature (T) in Fence Lake, August 23, 1929. A drainage lake. Com¬

pare with figs. 21-23.

In the case of Trout Lake, 11 wells had a smaller and 8 a

larger bound carbon dioxide content than the surface water of

the lake; the Kern well contained the smallest amount (4.0

mg/1) and that of the Forestry Headquarters house the largest

quantity (85.0 mg/1). The latter yielded almost nine times as

much as the former well.

The variations in the bound carbon dioxide content do not

seem to be correlated with the depth of the wells ; 12 wells rang¬

ing from 8 m. to 4-9 m. in depth varied from 2.0 to 21.0 mg/1.

In 55 wells between 5 m. and 9.9 m. in depth, the bound carbon

40 Wisconsin Academy of Sciences, Arts, and Letters .

dioxide ranged from 2.7 to 100.0 mg/1, while the second deepest

well (45.7 m.) contained only 7.2 mg/1. The variations in the

bound carbon dioxide content of the well waters, as well as of the

spring waters, appears to be due to the irregular distribution

of calcium and magnesium compounds in the glacial deposits of

this lake district.

The 7 spring waters showed a little more than a tenfold dif¬

ference in bound carbon dioxide content; the quantity ranged

from a minimum of 6.0 mg/1 in a spring situated on the shore

of Clear Lake (Manitowish waters) to a maximum of 64.0 mg/1

in one located on the shore of Little Horsehead Lake at Winegar.

Three of the spring waters yielded between 60.0 and 64.0 mg/1,

while the other four contained 44-0, 30.0, 9.5 and 6.0 mg/1, re¬

spectively. These marked variations in the bound carbon dioxide

content of the spring waters is also a good indication of the

irregular distribution of calcareous material in the strata through

which the water passes.

Fig. 25. Vertical distribution of the free (F) and bound (B) carbon dioxide,

pH and temperature (T) in Lake Mary, July 11, 1928. A drainage lake with

brown colored water. Compare with figs. 23 and 24. Note the unusual stratifi¬

cation of the pH.

Juday, Birge & Meloche — Lake Waters of N. E. Wisconsin 41

II. Hydrogen Ion Concentration

Introduction

Hydrogen ion observations have been made on natural waters

by many investigators during the past decade and a large amount

of literature dealing with this .subject has been published dur¬

ing this time. Some of the studies have consisted of single

or only a few determinations on a lake or a stream, while others

have included regular observations for considerable periods of

time. Investigations of the latter type have served to bring out

the diurnal, seasonal and annual changes of these waters. Under

natural conditions these changes were brought about largely

by biological processes; that is, the removal of carbon dioxide

from the dissolved bicarbonates by aquatic plants during the

process of photosynthesis gives the water a more alkaline re¬

action, while the addition of free carbon dioxide to- the water

in the processes of respiration and decomposition tends to give

it a more acid reaction. • The latter is true also of the carbon

dioxide obtained from the atmosphere.

These changes in reaction may be quite marked in the course

of a relatively short period of time. Skadowsky (1926) found

diurnal differences of 2.6 pH units in the surface waters of some

ponds and Philip (1927) noted daily fluctuations of 2.45 units

(pH 7.35 to 9.8) in the shallow water of Crystal Lake, Minne¬

sota.

In general, natural waters show a wide range in hydrogen

ion concentration; the range is from about pH 3.0 on the acid

side to pH 12.0 on the alkaline side. Jewell and Brown (1929)

obtained readings of pH 3.3 on the waters of pools in the sphag¬

num margin of a Michigan bog lake and Skadowsky (1926) re¬

ported readings of pH 3.4-3, 8 for the waters of some Russian

bogs. Strom (1925) has reported readings as low as pH 3.8

in the waters of peaty bogs of Norway. Yoshimura (1933) ob¬

tained a reading of pH 1.4 in a Japanese lake which received

water from a volcanic region ; the SO* content of the lake water

was 474 mg/1. In general however, readings below pH 4.0 are

not found in natural waters except under extreme bog condi¬

tions, or in cases of pollution with mineral acids.

42 Wisconsin Academy of Sciences , Arts , and Letters .

On the alkaline side, Jenkin (1982) obtained readings of pH

12.0 for the water of Lake Nakuru and of pH 10.7-11.1 for that

of Lake Elmenteita; these two lakes are situated in an African

Eift Valley and the high alkalinity is due to the presence of

sodium carbonate derived from volcanic rocks of their respective

drainage basins.

Methods

Regular observations were made on the hydrogen ion con¬

centration of the waters of the Highland Lake District during

the progress of these investigations. Up to the summer of 1982,

the readings were taken colorimetrically ; a standard La Motte

outfit with a range from pH 4.0 to 9-8 was employed most of

the time. The buffers covered 0.2 pH intervals so that the read¬

ings were made to the nearest 0.1 pH.

Certain differences were noted in the results when overlap¬

ping indicators were used on the same sample of water. These

differences were observed particularly in the soft waters which

had only small amounts of salts in solution and which, therefore,

Fig. 26. Vertical distribution of the free (F) and bound (B) carbon dioxide,

pH and temperature (T) in Presque Isle Lake, August 9, 1928. A drainage lake.

Compare with figs. 24 and 27.

Juday, Birge & Meloche—Lake Waters of N. E. Wisconsin 43

were weakly buffered. In order to obtain a better understanding

of the significance of these differences, both colorimetric and

electrometric methods were used during the summer of 1932. The

quinhydrone system was used for the latter method.

In general reasonably close agreements were found between

the results obtained by these two methods in waters where the

readings did not fall below pH 6; in such cases the differences

were not greater than 0. 1-0.2 pH. Below pH 6, however, larger

differences were noted at times; they ranged from 0.3 to 0.6 pH,

or even larger below pH 5. When the pH of the indicator was

adjusted to that of the weakly buffered waters, on the other

hand, the results obtained by the two methods agreed within

0.2 pH; most frequently the difference did not exceed 0.1 pH.

The larger differences obtained with the unadjusted indica¬

tors were due, apparently, to the buffering effects of the indica¬

tors. Investigations by Saunders (1926), Pierre and Fudge

(1928), Fawcett and Acree (1929) Acree and Fawcett (1930)

and Kolthoff (1931) have shown that colorimetric methods yield

unreliable results when employed on poorly buffered solutions

unless the indicators are specifically adjusted for the various

samples.

The general results obtained by the quinhydrone method

showed that most of the readings for the soft water lakes were

not greatly in error. In fact the differences were well within

the range of the seasonal and annual variations in pH noted in

these waters, so that it has not been necessary to discard any

of the colorimetric readings of the soft water lakes. Table XIII

shows a comparison between the quinhydrone readings and the

means of the readings obtained by colorimetric methods on 24

lakes. The colorimetric results represent the means of two to

27 determinations made on the surface water at different times

in the summer and in different years. The maximum difference

between the means of the colorimetric readings and of the quin¬

hydrone readings of 1932 in these 24 lakes was 0.4 pH ; there are

five cases showing a difference of this amount, while 14 cases

do not exceed 0.1 pH, leaving only two with a difference of 0.3

and three with 0.2 pH. Among those showing the maximum

difference, the colorimetric mean was above the quinhydrone

reading in four lakes and below in one, while the latter was

44 Wisconsin Academy of Sciences , Arts , and Letters .

higher in both instances showing a difference of 0.3 pH. In

the lakes showing differences of 0.1 to 0.2 pH, the colorimetric

means were above the quinhydrone readings in about half of

the cases and below in the other half.

Fig. 27. Vertical distribution of the free (F) and bound (B) carbon dioxide,

pH and temperature (T) in Trout Lake, August 22, 1932. A drainage lake. Com¬

pare with figs. 22-26.

The question arose also as to whether there was any change

in the hydrogen ion when a sample of water was brought to

the surface from the lower strata and exposed to the air while

the pH reading was being taken either color imetrically or elec-

trometrically. That is, do the changes in temperature and pres¬

sure, and the slight exposure to the air when the sample is

brought to the surface for the reading, have an appreciable

Juday, Birge & Meloche—Lake Waters of N. E. Wisconsin 45

effect upon the actual reaction of the water ? In order to answer

this question, a quinhydrone-calomel deep-water cell was de¬

signed which made it possible to take pH readings at various

depths in situ. A description of this instrument and some re¬

sults obtained with it have recently been published by Freeman,

Meloche and Juday (1933). The general results obtained with

this apparatus show that the readings taken with a standard

quinhydrone cell by bringing the samples to the surface are in

substantial agreement with those secured with the deep-water

cell at the various depths. The sample that is brought to the

surface, however, should be guarded against exposure to the air

as much as possible and the readings should be made promptly

in order to secure concordant results.

Fig. 28. This diagram shows the correlation between the bound carbon dioxide

and the calcium content of the surface waters of 357 lakes. It is based on the re¬

sults given in Table IX.

Surface Samples

Between 1925 and 1932 inclusive, pH readings were taken

on 1,225 surface samples; a summary of the results obtained

on them is given in Table XIV. The readings ranged from pH

4.4 in 4 very soft water lakes having specific conductances of

46 Wisconsin Academy of Sciences , Arts, and Letters.

8 to 10 x 10 ^ up to pH 9.4 m Clour Ldrkc ( Msmtowish w aters )

with a specific conductance of 71 x 106. The high alkalinity of

the surface water of Clear Lake on August 19, 1925 was correl¬

ated with a large crop of phytoplankton which yielded 7.0 mg/1

of dry organic matter ; the water contained — 2.2 mg/1 of mono¬

carbonate carbon dioxide at this time. In 3 surface samples

taken from Clear Lake in different years, the readings varied

from pH 7.2 to 9.4.

The 4 samples) with readings of pH 4.4 were obtained from

small lakes with much bog along their shores, but they were not

completely surrounded by bog. The color of their waters varied

from 16 to 41 on the platinum-cobalt scale, which indicates that

they do not belong to the extreme bog type-

No surface samples fell in the pH 4. 5-4. 6 group, but one gave

a reading of pH 4.7, 3 pH 4.8 and 5 pH 4.9. Some of these bodies

of water are lakelets entirely surrounded by bogs, while others

have varying amounts of bog tributary to them. The color

of their waters ranged from 8 to 230; the specific conductance

of these 9 samples varied from 7 to 19 x 10 6. The latter was

found in Knife Lake which belongs to the drainage type, since

it is the source of a small intermittent stream. All of the other

samples falling in the pH 4. 4-4. 9 group came from seepage lakes

and the maximum specific conductance found in them was

13 x 10-6.

At the alkaline end of the series, there were 31 surface

samples from 22 different lakes which gave readings of pH 8.5 to

9.4. All of these samples were obtained from drainage lakes.

The surface waters of all of these lakes except one had a com¬

paratively large amount of inorganic material in solution; the

specific conductance ranged from 37 to 111 x 10 6, the bound

carbon dioxide from 4.0 to 26.9 mg/1, the calcium from 2.4 to

18.8 mg/1 and the magnesium from 1.9 to 15.0 mg/1. Adelaide

Lake had the smallest amount of inorganic material and yielded

the above minimum amounts of these substances. The sec¬

ond lowest was Little Star Lake with a conductance of

39 x 10*6, bound carbon dioxide 9.5 mg/1, calcium 5.4 and mag¬

nesium 1.5 mg/1; thus the bound carbon dioxide and calcium

were more than twice as large in this lake as in Adelaide.

Three of the samples giving readings of pH 8.5 or more

were obtained in early May soon after the ice had disappeared

Juday, Birge & Meloche — Lake Waters of N. E. Wisconsin 47

from the lakes, but all of the others were found in July and

August, 10 in the former and 18 in the latter month. In some

cases these high alkalinities were correlated with large crops

of phytoplankton, others with large growths of the larger aquatic

plants and still others with large crops of both large aquatics

and phytoplankton.

Table XIV shows that 680 of the surface samples fall in the

groups between pH 4.4 and 6.9, and 595 of them in the groups

between pH 7.0 and 9.4, so that just a few more than half of

them were below the neutral point (pH 7-0) .

The seepage lakes, in general, contain a smaller amount of

dissolved inorganic substances which will serve as buffers than

the drainage lakes and this produces a characteristic difference

between the two types with respect to their hydrogen ion. The

differences found in the surface waters of the various lakes are

indicated in Table XV and they are shown graphically in Fig¬

ure 29. In the table and diagram, the results are grouped by

lakes and not by individual samples; where more than one ob¬

servation has been made on a lake, the mean of the various re¬

sults has been used to determine the group to which the lake

belongs.

Figure 29 shows clearly the characteristic difference between

the pH of the seepage and of the drainage lakes. In the seepage

type, there are 205 lakes, or 86 per cent of those belonging to

this class, lying btween pH 4.4 and 6.5, and only 33 lakes, or

14 per cent, above the latter group. The pH 5.8-5.9 group con¬

tains the largest number of lakes, namely 35, or 14.7 per cent of

the total number. In the drainage lakes on the other hand, 251

or approximately 90 per cent of those belonging to this type,

fall between pH 6.6 and 8.9, with a maximum of 42 lakes or 15

per cent of the total number in the pH 7.6-7.7 group. (Table

XV).

The 33 seepage lakes with readings above pH 6-5 overlap onto

some of the larger groups of drainage lakes. These seepage lakes

are located in areas where the glacial drift contains a larger

amount of calcareous material, so that their waters yield greater

pH values.

The 29 drainage lakes with readings below pH 6.6 possess

certain characteristics of the seepage type in that they have

48 Wisconsin Academy of Science, Arts, and Letters.

15

10

5

PH

15

10

5

pH

Fig. 29. This diagram shows the hydrogen ion content of the surface waters of

238 seepage and 280 drainage lakes. The vertical spaces show the percentage of

lakes in the various pH groups. Note the striking difference in the grouping of the

two types of lakes. Compare with fig. 1.

very soft waters and yield low pH values. While these bodies

of water have natural outlets and thus belong to the drainage

class, they are the sources of small intermittent streams which

contain out-flowing water only when the water level of the

lake reaches a rather high stage. They have no inlets, so that

their outlet streams may be dry for the greater part of the year,

Juday, Birge & Meloche — Lake Waters of N. E. Wisconsin 49

or for several years in some instances, depending upon the

amount of the annual precipitation.

Variations in pH Readings

Annual . During the progress of this investigation, surface

samples were obtained from 245 lakes in different years ; a few

of the observations were limited to two but most of them in¬

cluded three or more years. Table XVI shows the maximum pH

differences found in these lakes in the different years. The first

group includes those in which the readings did not differ more

than 0.4 of a pH in the various years ; 65 lakes, or 26 per cent

of the total number, fall in this group. Five of these 65 lakes

gave the same readings in two or more years and 16 others

showed differences of only 0.1 pH. In 106 lakes, or 43 per cent

of the total number, the pH differences in the different years

were between 0.5 and 0.9; this is the largest number of lakes

in any of the groups. These two groups, therefore, include 69

per cent of the lakes that were visited more than once- That is,

the greatest difference found in the surface waters of these

lakes over periods of two to six years was less than one pH unit

in 69 per cent of them. Adding the third group to the first and

second gives 222 lakes, or 90 per cent of the total number, in

which the difference did not exceed 1.4 pH units in the various

years.

In 6 lakes the annual differences exceeded two pH units;

the maximum difference was found in Adelaide Lake where the

readings varied from pH 6.3 on August 29, 1929 to pH 8.8 on

August 23, 1925. Nine readings were made on the surface

water of Adelaide Lake between 1925 and 1930, inclusive, and

the two obtained in the summer of 1925 where the only ones

which exceeded pH 6.8. On July 20, 1925, the surface water was

pH 7-1 and the free carbon dioxide amounted to 1.2 mg/1; on

August 23, 1925, the reading was pH 8.8 and this was correlated

with a phenolphthalein alkalinity equivalent to — 0.5 mg/1 of car¬

bon dioxide. The water of this lake is quite soft; the mean quan¬

tity of bound carbon dioxide was 3.2 mg/1 and the calcium con¬

tent 2.4 mg/1. Thus, a relatively small change in the carbon diox¬

ide content of the water during the process of photosynthesis

causes a marked change in the reaction. In 1925 the change in

50 Wisconsin Academy of Sciences , Arts, and Letters .

hydrogen ion was correlated with an increase in dissolved oxy¬

gen from 7.9 mg/1 on July 20 to 9.1 mg/1 on August 2&

Spring and Summer . Hydrogen ion observations were made

on the surface waters of 36 lakes in the spring and again in the

suirqner of the same year. Such sets of readings were taken on

some of the lakes for two or three different years, so that 57

pairs of readings were secured on the 36 lakes. In 13 of the

pairs, the summer pH value was lower than that of the spring;

the maximum decline was noted in Crystal Lake in 1930 where

the reading on April 26 was pH 5.8 and on the following July 9

it was pH 5.1, thus showing a decrease of 0.7 pH. Second in

rank were Weber Lake with a surface reading of pH 5.8 on

April 26 and pH 5.2 on August 14, 1930, and Little Arbor Vitae

Lake with pH 8.2 on May 4 and pH 7.6 on July 7, 1927. There

was a summer decline of 0-5 pH in Little John Jr. Lake, but

none of the other pairs showed a summer decrease of more

than 0.3 pH.

In 9 pairs of readings the spring and summer results were

the same, thus leaving 35 pairs in which the summer surface

samples were more alkaline than those taken in the spring. The

maximum increase in alkalinity was noted in Little Tomahawk

Lake where the readings were pH 7.2 on May 6 and pH 8.1 on

July 25, 1927. In four samples the increase in the summer

alkalinity amounted to 0.7 pH, in one case 0.6 and in three

cases 0.5; all of the other increases were below the latter amount.

Autumn . Readings were obtained on the surface water of

four lakes in the month of November 1930; all of them were

lower than those noted during the previous summer. The maxi¬

mum autumnal decrease was found in Weber Lake where the

reading was pH 6.7 on June 29 and pH 6.0 on November 16, a

difference of 0.7 pH. In Trout Lake the hydrogen ion was pH

7.7 on August 27 and pH 7-2 on November 15. The smallest

difference was noted in Crystal Lake with pH 6.0 on August 20

and pH 5.9 on November 16. The difference in High Lake was

0.3 of a unit. k.

Diurnal changes . In order to determine the extent of the

diurnal fluctuations in the hydrogen ion of the surface water of

Trout Lake, a series of observations was made on August 15,

Juday, Birge & Meloche — Lake Waters of N. E. Wisconsin 51

1933. On this date the sky was clear until about mid-forenoon,

then partly cloudy until mid-afternoon, after which it was clear

again. There was a slight southwest wind (on-shore) and the

disc reading was 4.5 m. Surface samples were taken at two

stations every hour from 8 a.m. to 4 p.m. and again at 8 and 10

p. m. One series of these samples was taken at the end of the

laboratory dock or about 15 m. from the shore, while the other

was taken about 200 m. further out in the lake where the water

had a depth of about 5 m. During the day time, the readings

obtained at the two stations fell between pH 7.5 and 7.7, but they

were somewhat lower in the evening, or pH 7.2 to 7.3. Thus the

maximum diurnal difference noted in these observations was

0.5 pH. Large aquatic plants are scarce in Trout Lake and the

crop of phytoplankton is usually of moderate size, so that only

a relatively small diurnal fluctuation in the hydrogen ion read¬

ings is to be expected in the open water.

Vertical Distribution

The pH values at different depths are given for a number of

lakes in Table X and some of the results are represented in

Figs. 2-15 and 17-27. The various types of lakes are included in

both the table and the diagrams.

In a shallow lake like Bear (Fig. 2) where the entire body

of water is kept in circulation during the summer, the pH read¬

ings were uniform from surface to bottom. In all of the lakes

which had a bottom stratum that did not take part in the gen¬

eral circulation during the summer, there was a decrease in the

pH values in the lower water. This phenomenon is well illus¬

trated in such shallow lakes as Dorothy Dunn, Finley and Brag-

onier (Figs. 3, 5, 6) and in the deeper ones which have a more

definite thermal stratification, such as Crystal, Fence, Muskel-

lunge, Presque Isle and Trout (Figs* 11, 12, 24, 26 and 27).

Crystal Lake showed a relatively small difference between

surface and bottom ; the former reading was pH 5.5 and that at

19 m. (1 m. above the mud) was pH 5.3 on August 11, 1932. In

Crawling Stone Lake (Fig. 22), the difference between surface

and bottom readings Was 0.4 pH, in Black Oak, Presque Isle, and

Trout lakes 1.0 pH and in Muskellunge 1.8. In Adelaide Lake,

the surface reading was pH 8.8 and that at 20 m. was 6.2 on

52 Wisconsin Academy of Sciences , Arts, and Letters .

August 23, 1925 ; this difference of 2.6 pH units represents the

maximum between surface and bottom readings. This maximum

is exceeded by that of Pallette Lake (Fig. 18) if the reading

at 8 m. is compared with that at 16 m- on August 22, 1928 ; the

former was pH 9.1 and the latter 6.3, a difference of 2.8 pH

units.

Some of the vertical series taken in Anderson, Pallette and

Silver lakes (Figs. 17-19) deserve special mention. The sam¬

ples obtained in the thermocline of these lakes were more alka¬

line in some cases than those of the epilimnion or the hypolim-

nion. A maximum reading of pH 9.1 was noted in Pallette Lake

at 8 m. on August 22, 1928, while that at the surface was pH

6.7 and at the bottom pH 6.3 ; the alkaline stratum was sharply

defined as the reading changed from pH 6.6 at 7 m. to 9.1 at

8 m. and then back to 6.9 at 9 m. The most alkaline water

found in Anderson Lake was pH 8.4 at 7 and 8 m. on August

8, 1929, and pH 8.3 in Silver Lake on August 13, 1932. These

more alkaline readings were correlated with excess oxygen and

also with a more or less marked decrease in the carbon dioxide

content of the water in these strata as shown in the diagrams ;

all of these changes were due to the photosynthetic activities of

the phytoplankton populations of these strata.

Calculated pH Values

The last column in Table X shows the pH values that have

been computed from the carbon dioxide determinations by means

of Kolthoff’s formula. This formula, however, is intended pri¬

marily for the computation of the free carbon dioxide from

the pH and bicarbonate results in as much as free carbon dioxide

titrations may be subject to certain inaccuracies under field con¬

ditions.

In the great majority of the samples, the calculated pH val¬

ues are in reasonable agreement with the actual readings; in

most cases represented in the table the difference did not exceed

0.2 pH. In some samples the difference amounts to as much as

0.4 pH; these larger differences are found most frequently in

the region of the thermocline where temperature and chemical

conditions show rapid changes with increasing depth and also

in lakes having very soft waters. A more complete study of

this problem is included in plans for future investigations.

Juday, Birge & Meloche — Lake Waters of N. E. Wisconsin 53

Even larger differences between calculated and observed pH

values are indicated in some of the well waters in Table XI. No

satisfactory explanation for these discrepancies has yet been

found and this problem also remains for future investigation.

Dichotomous Stratification of Hydrogen Ion

In a recent paper Yoshimura (1932) describes an unusual

type of vertical distribution of the hydrogen ion in some Japan¬

ese lakes. Minimum pH readings were obtained by him in the

upper part of the hypolimnion, with higher values both above

and below this more acid layer. He applied the term “dichoto¬

mous stratification” to this type of vertical distribution. In the

most common type, the lowest pH values are found in the lower

part of the hypolimnion, or in the bottom stratum.

Yoshimura computed the pH for some of the Wisconsin lakes

on the basis of their carbon dioxide content and found that they

also showed a dichotomous stratification. Two of the lakes men¬

tioned by him as showing this type belong to the group included

in this report, namely Allequash and Mary lakes. Allequash

Lake has a maximum depth of only 7.5 m., so that it does not

show a very marked thermal stratification in summer ; in fact it

can hardly be regarded as possessing a hypolimnion. A series

of samples taken at various depths on August 15, 1926 showed

the usual type of pH stratification; that is, the hydrogen ion

changed from pH 8.6 at the surface and 3 m. to a minimum of

pH 6.7 at 7 m.

The dichotomous type of stratification was observed in Lake

Mary, however; this lake has an area of 1.2 ha. and a maximum

depth of 22 m., which is an unusual depth for a glacial lake of

this size. During this investigation, 5 series of samples cover¬

ing the entire depth of the lake were taken, but only two of them

showed dichotomous stratification. On July 11, 1928, the read¬

ing was pH 6.2 at the surface and 54 at 3 m. ; below the latter

depth the readings gradually rose to pH 5.9 in the 15-21 m.

stratum. The results of these observations are given in Table

XVII and they are shown graphically in Figure 25: the latter

brings out clearly the minimum value obtained at 3 m. The

series taken on July 29, 1927 showed a similar type of pH strat¬

ification, with a minimum of pH 5.8 again at 3 m. and pH 6.2

54 Wisconsin Academy of Sciences, Arts, and Letters.

at both surface and bottom. In the series of August 21, 1926,

the reading was pH 6.2 at the surface, 5.8 in the 3-5 m. stratum,

and 5.9 from 8 to 20 m. ; this small, increase of 0.1 pH in the

lower water can hardly be regarded as a definite case of dicho¬

tomous stratification because it falls within the limit of error

of the colorimetric method of hydrogen ion determination. The

pH readings were uniform throughout the hypolimnion in the

other two series of observations, so that two series showed a

definite dichotomous stratification and the other three did not.

Serial observations were made on 71 other lakes which were

deep enough to have a well marked hypolimnion, but no clear cut

examples of dichotomous stratification were found in them. Some

evidence of it was noted in 4 of them, but the pH value of the

bottom water was only 0.2 pH above that of the minimum inter¬

mediate layer. In Mud Lake for example, with a maximum

depth of 14.5 m., the reading was pH 6.3 at the surface, 5.7 in

the 5-8 m. stratum, and 5.9 at 13 m. This difference, however,

is not regarded as large enough to constitute a definite dichoto¬

mous stratification.

In the Japanese lakes of this class, Yoshimura found that

the minimum pH reading always appeared in the layer of water

immediately above the anaerobic stratum. The same result was

obtained in Lake Mary; the minimum reading of pH 5.4 at 3 m.

was correlated with an oxygen content of 0.4 mg/1 at that depth

and no oxygen was found below this depth.

According to Yoshimura, this type of pH stratification is

found only in lakes with a bottom stratum which has no dis¬

solved oxygen but which possesses a larger or smaller amount

of free carbon dioxide. This water which is charged with free

carbon dioxide, attacks the calcium, iron and manganese com¬

pounds in the bottom deposits and converts them to bicarbonates

which readily pass into solution. These dissolved substances

increase the pH value in spite of the free carbon dioxide that

is present.

Table XVII and Figure 25 show that there was a marked

increase in the bound carbon dioxide with increasing depth in

Lake Mary on July 11, 1928. On this date also the specific con¬

ductance ranged from 21 x 106 at the surface to 58 x 10 6 at 21

m. ; likewise the total residue increased from 51.3 mg/1 at the

Juday , Birge & Meloche — Lake Waters of N. E. Wisconsin 55

surface to 82.8 mg/1 at 21 m. and the calcium rose from 1.3

mg/1 at the surface to 2.7 mg/1 at the bottom. No determina¬

tions of the iron and manganese have been made so that their

vertical distribution is not known. Similar increases in the

bound carbon dioxide, the specific conductance and the total

residue were noted in the lower water in the three series which

did not show a dichotomous stratification and this seems to

indicate that some other factor or factors play a part in pro¬

ducing the phenomenon. Iron and manganese are regarded

as the chief factors by Yoshimura, but the lake which showed

the most marked increase of these two substances in the lower

water in this investigation, gave no indication whatever of a

dichotomous stratification in three sets of observations.

A reverse type of hydrogen ion stratification is described

and illustrated for Anderson, Pallette and Silver lakes. (Figs.

18-20). In these bodies of water, the highest pH values were

found in the intermediate stratum, while in the type described

by Yoshimura the lowest pH values were located in the inter¬

mediate stratum. Both cases represent a dichotomous stratifi¬

cation of the hydrogen ion, but one is the opposite of the other.

In order to distinguish the two types, the term acid dichotomous

stratification may be used to designate that in which the minimum

pH value is found in the intermediate zone of the lake and alka¬

line dichotomous stratification for that in which the highest pH

value falls in the intermediate layer.

An alkaline stratification was observed by Kusnetzow and

Schtcherbakow (1927) in Lake Baikal. The readings in the

upper 2 m. of water varied from pH 7.6 to 7.8, rose to pH 8.15

in the 6-10 m. stratum and then declined to pH 7.6 at 25 m.

where it remained for the rest of the series, namely down to

104 m.

Hutchinson and Piekford (1932) also found an intermediate

stratum in Mountain Lake, Virginia with somewhat higher pH

values than those above and below it ; the hydrogen ion changed

from pH 6.4 at the surface to 6.6 at 5 m. and 7 m. and then

fell to 6.5 at 11 m. and 19 m.

Hydrogen Ion and Carbon Dioxide

Carbon dioxide plays a very important role in determining

the reaction of lake waters. The supply of this gas is derived

56 Wisconsin Academy of Sciences, Arts, and Letters .

from the atmosphere, from inflowing waters, especially that of

springs, and from the decomposition and respiration that take

place within the lakes. Aquatic plants withdraw a certain

amount of this carbon dioxide in their processes of assimilation,

so that the quantity actually found at any particular time is the

resultant of the processes which tend to augment the supply and

of those which reduce the supply. Thus the interplay between

these two types of activities bears a close relation to the hydro¬

gen ion content of the water.

Under certain conditions, the reaction of the water is af¬

fected by other factors; in typical bog lakes and lakelets for

example, the humic acids play a part in determining this re¬

action. In the great majority of the lakes that have been visited

in this investigation, however, the hydrogen ion concentration

was chiefly or wholly a function of the carbon dioxide content

of the water.

Free carbon dioxide. The relation between the free carbon

dioxide and the hydrogen ion of the surface waters is given for

the various pH groups in Table XVIII ; the results are also shown

graphically in Fig. 30. As already indicated, the bog waters

Fig. 30. Relation between the hydrogen ion and the free carbon dioxide con¬

tent of the surface waters of 499 lakes. The solid line repreesnts drainage lakes and

the broken line seepage lakes.

Juday, Birge & Meloche — Lake Waters of N. E. Wisconsin 57

gave a rather high acid reaction to phenolphthalein ; thus they

may be regarded as representing a special type of water. In

compiling the data for Table XVIII, therefore, all of the samples

which contained more than 4.0 mg/1 of free carbon dioxide have

been omitted.

In the seepage lakes, the mean quantity of free carbon diox¬

ide decreased from 2.3 mg/1 in the pH 44-4.9 group to 1.4 mg/1

in the 7.0-7.4 group ; this is a decrease of only 0.9 mg/1 of free

carbon dioxide for the entire range from pH 4.4 to 7.4, an un¬

usually small change for such a marked difference in hydrogen

ion activity. (Fig. 30).

In the drainage lakes, the free carbon dioxide fell from 3.7

mg/1 in the pH 5. 0-5.4 group to 2.4 mg/1 in the 5. 5-5. 9 group ;

the quantity remained substantially the same as the latter in

the two following groups, but the 7. 0-7.4 group showed an ap¬

preciable decrease in free carbon dioxide, with a further decline

in the 7-S-7.9 group. (Fig. 30). This marked decrease in free

carbon dioxide continued beyond the latter group. Both the

table and the diagram show that the most marked decrease in

the quantity of free carbon dioxide came in the region of pH 8.0

where the water begins to give an alkaline reaction to phenolph¬

thalein, thus indicating the presence of normal or monocarbo¬

nate carbon dioxide. It will be noted that the seepage lakes

yielded a smaller mean quantity of free carbon dioxide than the

drainage lakes belonging to the same pH groups; this fact is

shown clearly in Figure 30.

The vertical distribution of the hydrogen ion and the free

carbon dioxide is shown in Figures 2-15 and 17-27. In the shal¬

lower lakes a striking correlation between the increase of free

carbon dioxide in the lower stratum and the decrease of the pH

values was noted in Allequash Lake (Fig. 7). In Crystal Lake

(Fig. 11), the change in both pH and free carbon dioxide was

not very marked in the lower water. The changes were much

more marked in Muskellunge Lake (Fig. 12), as well as in

Fence and Trout lakes (Figs. 24, 27). An interesting correla¬

tion between hydrogen ion and free carbon dioxide in the ther-

mocline of Anderson, Pallette and Silver lakes (Figs. 17-19)

was found; the phytoplankton removed enough carbon dioxide

from this stratum in the process of photosynthesis to produce

a sharp rise in the pH values.

58 Wisconsin Academy of Sciences , Arts , and Letters.

Bound carbon dioxide. Table XVIII shows the relation be¬

tween the hydrogen ion concentration and the bound carbon

dioxide content of the surface waters of 499 lakes ; of this num¬

ber 227 are seepage and 272 drainage lakes. It will be noted

that the groups with low pH values contain only a small amount

of bound carbon dioxide. Between pH 4.4 and 5.9 for example,

the mean quantity of bound carbon dioxide in the surface waters

of the seepage lakes varies from 1.3 to 1.6 mg/1 ; between pH

6.0 and 7.4 however, there is an increase in the mean bound

carbon dioxide content of the water from 2.0 to 7.7 mg/1.

In the drainage lakes, the first two groups have rather small

carbon dioxide means, but the pH 6. 0-6. 4 group shows an ap¬

preciable increase over the first two; this increase continues in

the following groups so that it rises to 19.8 mg/1 in the pH

8. 5-8.9 group.

20

15

10

5

0

4.5 5.0 5.5 6.0 6.5 7.0 75 8.0 85 9.0

Fig. 31. Relation between the hydrogen ion and the bound carbon dioxide con¬

tent of the surface waters of 499 lakes.

Figure 31 shows the results for hydrogen ion and bound

carbon dioxide in a graphic form. The curves of this diagram

serve to bring out clearly the relatively slight variations in the

bound carbon dioxide of the low pH groups of both seepage and

drainage lakes and the increases in amount which are found in

the more alkaline waters of both types of lakes. They show

also that there is only a small difference in the mean bound

carbon dioxide content of the seepage and drainage lakes up

to about pH 6.0, but at the higher pH values the waters of the

drainage lakes contain a larger amount of bound carbon dioxide

for corresponding pH values than the seepage lakes- This dif¬

ference may be accounted for by the fact that the waters of the

Juday, Birge & Meloche — Lake Waters of N. E. Wisconsin 59

drainage lakes have a somewhat larger free carbon dioxide con¬

tent than those of the corresponding seepage lake groups.

Hydrogen Ion and Calcium

The relation between the hydrogen ion and the calcium con¬

tent of the waters of 358 lakes is given in Table XIX. In the

pH 4.8-6.1 range, the mean quantity of calcium varies from 0.5

to 1.49 mg/1 ; the latter amount is found in the pH 5.6-5.7 group

and it is followed by a decline to a mean of about 1.0 mg/1 in

the next two groups. The increased amount of calcium in the

pH 5.6-5.T group is due to the fact that three of the 8 lakes in

this group yielded from 2.1 to 2.8 mg/1 of calcium. A larger

number of lakes falling in this group would probably give a

lower mean.

Beginning with pH 6.2, there is an increase of the mean cal¬

cium content of the successive pH groups, rising to 10.2 mg/1 in

the 8.2-9.2 group. All of the results above pH 8.2 have been com¬

bined into one group in order to get enough lakes to give a

fair mean. Likewise, one lake with a reading of pH 4.4 has been

included in the 4.8-4.9 group because the calcium content of this

water fell within the range of that group. No calcium determin¬

ations were made on the other lakes that gave readings below

pH 4.8.

Fig. 32. Relation between the hydrogen ion and the calcium content of the

surface waters of 358 lakes.

60 Wisconsin Academy of Sciences, Arts , and Letters.

The results are shown graphically in Figure 82. The curve

in this diagram brings out clearly the variations in the mean

quantity of calcium between pH 4.8 and 6.1. These variations

are due, in part at least, to differences in the free carbon dioxide

content of the various lake waters falling within this hydro¬

gen ion range. Relatively small differences in free carbon diox¬

ide content have an important effect upon the hydrogen ion read¬

ings in waters with as small amounts of buffers as those found

in this particular group of lakes-

Beyond pH 6.1 the curve shows a gradual increase in the

mean calcium content of the water correlated with rising pH val¬

ues. It is interesting to note that the steepest part of the curve

falls in the region of neutrality (pH 7.0). The entire curve

falls roughly into three parts ; the first part includes that in which

there is very little gradient in the curve, extending from pH 4.8

to 6.1. The second part extends from pH 6.2 to 7.8 and includes

the region with the steepest gradient, while the third part is

represented by the more gentle gradient above pH 7.8. The

maximum quantity of calcium was 18.8 mg/1 ; it was correlated

with a hydrogen ion of pH 8.5. The highest pH value found in

the lakes on which calcium determinations were made (pH 9.2)

was correlated with 8.2 mg/1 of calcium.

Hydrogen Ion and Biota

During the past decade or two, many investigations have

dealt with the relation of the hydrogen ion activity of natural

waters to the various living organisms found in them- A large

amount of biological material has been collected from the Wis¬

consin lakes during these studies, but much of this material must

be studied more thoroughly before a profitable discussion of

the relation of the physical and chemical factors to the biological

phenomena can be presented.

A general physico-chemical correlation is planned, however,

rather than one between a single factor, such as hydrogen ion,

and the biota. The hydrogen ion activity of these lake waters

is regarded mainly as an index of certain underlying chemical

conditions which obtain in them and which represent the funda¬

mental basis for the particular reactions that have been found.

That is, a number of chemical factors must be taken into con-

Juday, Birge & Meloche — Lake Waters of N. E. Wisconsin 61

sideration in order to obtain a complete picture of~the relation¬

ships which exist between the reactions of the water and the

lake biota. This general idea has been well expressed by Strom

(1925) in the following statement: “The specific reaction of

the water is a potential factor in determining the character of

the biota, especially the algal flora, but it must always be re¬

garded in conjunction with the other physical and chemical

properties of the medium.”

Brief mention may be made of the work of Morrison (1932)

on the Mollusca of these Wisconsin lakes in relation to hydrogen

ion and bound carbon dioxide. He found Pisidium and Cam-

peloma in some of the very soft water lakes where the hydro¬

gen ion was pH 5.7 and the bound carbon dioxide content was

only 1.0 mg/1. In the bog type of soft waters, large specimens

of Pisidium were found where the reaction was pH 5.1. The

Valvatidae, on the other hand, were not found in waters with

a reaction below pH 7.1 or with less than 8.0 mg/1 of bound

carbon dioxide. In the other groups, some species preferred acid

and others preferred alkaline water.

Hydrogen Ion of Well and Spring Waters

Hydrogen ion readings were taken on 149 well and 7 spring

waters. In the well waters each tenth of a pH was represented

from pH 5.3 to 8.1; in addition, one sample gave a reading of

pH 5.0, so that only two groups were missing between pH 5.0

and 8.1, namely pH 5.1 and 5.2. Representative results ob¬

tained on various well waters are given in Table XI.

The maximum number of these well waters fell in the pH

6.3 group, namely 21. The next largest number was 13 in the

pH 6.2 group, which was followed by 10 at pH 5.9. Three of the

waters gave readings of pH 7.0, with a like number at pH 6.9

and 6 at pH 7.1, so that 12 of the 149 samples were grouped

around the neutral point. Two samples gave readings of pH

8-0 and 3 pH 8.1; the latter was the highest pH value obtained

in any of the well waters.

The various samples are combined into larger groups on the

hydrogen ion basis in Table XX. This table shows the maximum

number of well waters in the pH 6. 0-6.4 group, namely 54, or 36

per cent of the total number of wells represented. The pH 6.5-

62 Wisconsin Academy of Sciences, Arts, and Letters.

6.9 group was second with 27 wells, or 18 per cent of the total

number. Three of the waters gave a neutral reaction, while 104

were on the acid side of neutrality and 42 on the alkaline side.

Of the 140 wells situated on the immediate shores of lakes,

the water from 99 of them gave lower pH readings than the

surface waters of the respective lakes and 83 gave higher pH

values, thus leaving 8 samples in which the surface and well

waters were the same. The maximum difference between lake

and well water was noted at Madeline Lake where a reading

of pH 8.8 was obtained for the former and pH 5.5 for the latter,

a difference of 3.3 pH. The second largest difference was found

at Lost Lake where the surface water was pH 8.1 and the well

water pH 5.9, a difference of 2.2 pH. In the well waters which

were more alkaline than the lake waters, the maximum differ¬

ence was obtained at Lake George where the reaction of the lake

water was pH 6.0 and that of the well water was pH 7.8.

The waters from wells situated on the shores of very soft

water lakes usually had somewhat higher pH values than the sur¬

face waters of the respective lakes; this was true particularly

of lake waters which gave readings between pH 4.5 and 6.0.

Certain exceptions were noted, however. The well and surface

water of McGrath Lake were both pH 5.0. Some of the lake

waters with values above pH 6.0 were also exceeded by their

respective well waters. The surface of Oxbow Lake, for ex¬

ample, was pH 7.3 and two well waters on its shores gave read¬

ings of pH 8.1 and 7.8.

The 19 wells listed on Trout Lake in Table XIII show that

there is a marked variation in the hydrogen ion content of the

waters of wells situated on the shores of a single lake ; the range

of the hydrogen ion in these wells was from pH 5-6 in the Kern

well to pH 7.9 in the barn well at the State Forestry Headquar¬

ters ; one of these well waters was neutral, while 10 were on the

acid side and 8 on the alkaline side of neutrality. In the 9 wells

situated on the shores of Plum Lake, the range was from pH

5.7 to 7.5; only one of them was alkaline. These variations in

hydrogen ion content, together with those noted in the carbon

dioxide content, show clearly the marked local variations in the

chemical composition of the ground waters of this region as

represented by these wells.

Juday, Birge & Meloche — Lake Waters of N. E. Wisconsin 63

There was a general correlation between the depths of the

Trout Lake wells and the hydrogen ion content of their waters.

Depth records were obtained for 14 of the 19 wells on this lake.

Of this number, 6 wells ranged from 4 to 6.7 m. in depth and the

mean of the readings was pH 6.1; 5 of them varied from 8

to 13.7 m. in depth and their mean value was pH 7.0. The depth

of three of them fell between 21 and 25 m. and their mean was

pH 7.6- Thus there was an apparent increase in the alkalinity

of the water with increasing depth of the wells.

This did not hold true for the 9 Plum Lake wells, however;

the deepest one (13.7) in that group gave a reading of pH 6.3

and the shallowest one (4m.) pH 6.2 and this difference is too

small to have any significance.

The hydrogen ion content of the Trout Lake well waters was

not correlated with that of the lower water of the lake. In 25

series the mean reading at a depth of 32 m. was pH 6.9, while

the well waters varied from pH 5.6 to 7.9. This was true of

Plum Lake also where the mean for the bottom samples (17 m.)

in 4 series was pH 6.9 and the well waters ranged from pH 5.7

to 7.5.

Hydrogen Ion and Carbon Dioxide in Well Waters

Free carbon dioxide . Table XX shows the general relation

between the hydrogen ion activity and the free carbon dioxide

content of 149 well waters. The first group (pH 5.0-5.4) shows a

somewhat smaller mean quantity of free carbon dioxide than

the second and third, but there are only 3 wells in this group,

which is too small a number to give a fair mean. Beginning with

the pH 5.5-5.9 group, the mean quantity of free carbon dioxide

declines from a maximum of 22.0 mg/1 to a minimum of 1.9 mg/1

in the 8.0-8.4 group. It will be noted, however, that the mean

of the pH 7. 0-7.4 group is a little smaller than that of the 7.5-

7.9 group.

Bound carbon dioxide . The mean quantity of bound carbon

dioxide is given for the various pH groups in Table XX. The

amjount of the former rises from 2.9 mg/1 in the pH 5. 0-5.4

group to 44.4 mg/1 in the 7.5-7. 9 group. The mean then de¬

clines to 33.4 mg/1 in the 8.0-8.4 group, but this represents only

5 well waters, which is insufficient for a fair mean. This differ-

64 Wisconsin Academy of Sciences, Arts, and Letters .

ence is due in part also to the fact that the pH 7. 5-7.9 group con¬

tains two well waters with unusually large amounts of bound

carbon dioxide; one of them yielded 100.0 mg/1 and the other

82.0 mg/1. If these two were omitted, the mean for this group

falls to a little less than 40.0 mg/1 instead of the 44*4 mg/1

indicated in the table.

Hydrogen Ion of Spring Waters

In the 7 spring waters, the hydrogen ion ranged from pH

6.2 to 8.4 ; a second one gave a reading of pH 6.3, but the other

5 fell on the alkaline side of the neutral point. Four of them

varied from pH 7.3 to 7.8, while the fifth one was pH 8.4. The

spring water which gave a reading of pH 6.2 yielded 13.0 mg/1

of free and 6.0 mg/1 of bound carbon dioxide; the one giving a

reading of pH 8.4 yielded 9.5 mg/1 of free and 64-0 mg/1 of

bound carbon dioxide. These were the minimum and maximum

amounts of bound carbon dioxide in the spring waters ; the free

carbon dioxide ranged from a minimum of 4.0 to a maximum

of 41.0 mg/1.

SUMMARY

1. Quantitative determinations of free and bound carbon

dioxide and of hydrogen ion concentration were made on the

surface waters of 518 lakes.

2. The surface waters of 238 seepage lakes (those without

inlets or outlets) gave an acid reaction to phenolphthalein which

varied from 0.3 to 10.7 mg/1 when expressed in terms of free

carbon dioxide. The hydrogen ion readings fell on the acid side

of neutrality (pH 7.0) in the surface waters of 93 per cent of

the seepage lakes.

3. The surface waters of 36 drainage lakes (those with out¬

lets) gave a mean alkaline reaction to phenolphthalein which

was equivalent to — 0.1 to — 10.7 mg/1 when expressed in terms

of carbon dioxide. The surface waters of 244 additional drain¬

age lakes gave a mean acid reaction to phenolphthalein equiva¬

lent to 0.1 to 7.0 mg/1 of free carbon dioxide. The hydrogen

ion readings were on the acid side in 28 per cent of the drain¬

age lakes, while the others were either neutral or alkaline.

Juday, Birge & Meloche — Lake Waters of N. E. Wisconsin 65

4. Annual variations in free carbon dioxide range from zero

up to 14.7 mg/1 in the surface waters of the various lakes;

these were due to the photosynthetic activities of the phytoplank¬

ton. Similar variations were noted in the hydrogen ion content

of the water.

5. The free carbon dioxide was uniformly distributed from

surface to bottom in some lakes, while the lower water of others

yielded a much larger amount than the upper. A marked de¬

crease of free carbon dioxide was noted in the thermocline of 3

lakes.

6. The waters of seepage lakes yielded from 0.2 to 11.3 mg/1

of bound carbon dioxide; in the drainage lakes the range was

from 1.0 to 39.0 mg/1.

7. In most of the seepage lakes, the annual variations in

bound carbon dioxide were small ; in 56 per cent of the drainage

lakes, these annual differences did not exceed 2.0 mg/1, while one

gave a maximum difference of 8.5 mg/1.

8. The differences between the bound carbon dioxide con¬

tent of surface and bottom waters varied from zero to 13.3

mg/1.

9. An average increase of about 2.3 mg/1 of bound carbon

dioxide was correlated with an increase of 10 x 10 6 in the speci¬

fic conductance of the water.

10. The free carbon dioxide content of the 149 well waters

ranged from — 0.5 to 67.5 mg/1 and the bound carbon dioxide

from 2-0 to 100.0 mg/1.

11. In 7 spring waters, the free carbon dioxide varied from

4.0 to 41.0 mg/1 and the bound carbon dioxide from 6.0 to 64.0

mg/1.

12. The free and bound carbon dioxide content of the lake

waters was independent of that of the ground water as rep¬

resented in the springs and wells.

13. The hydrogen ion readings of the surface waters ranged

from pH 4.4 to 9.4.

14. The annual variations in the hydrogen ion activity of

66 Wisconsin Academy of Sciences, Arts, and Letters.

the surface waters varied from zero to 2.5 pH units. Likewise

the differences between surface and bottom waters varied from

zero to 2.6 units.

15. The acid type of dichotomous stratification of hydrogen

ion was found in one lake and the alkaline type in three.

16. The hydrogen ion concentration showed a fairly definite

correlation with both free and bound carbon dioxide and with

the calcium content of the water.

17. Three of the well waters were neutral (pH 7.0), 103

were acid (pH 5.0-6.9) and 43 were alkaline (pH 7.1-8.4).

Literature

Acree, S. F. and E. H. Fawcett. 1930. The problem of dilution in colorimetric

hydrogen ion measurements. II. Use of isohydric indicators and superpure

water for accurate measurement of hydrogen ion concentration and salt errors.

Indus, and Eng. Chem., Analyt. Ed. 2:78-85.

Beadle, L. C. 1932. Scientific results of the Cambridge expedition to the east

African lakes, 1930-31. IV. The waters of some African lakes in relation to

their fauna and flora. Jour. Lin. Soc. London, Zool. 38:158-211.

Birge, E. A. and C. Juday. 1911. The inland lakes of Wisconsin. I. The dissolved

gases of the water and their biological significance. Bui. No. XXII. Wis. Geol.

and Nat. Hist. Survey. 259 pp. Madison.

Birge, E. A. and C. Juday. 1914. A limnological study of the Finger lakes of

New York. Bui. U. S. Bur. Fish. 32:525-609. Doc. No. 791.

Fawcett, Edna H. and S. F. Acree. 1929. The problem of dilution in colorimetric

hydrogen ion measurements. I. Isohydric indicator for accurate determination

of pH in very dilute solutions. Jour. Bact. 17:163-204.

Freeman, S., V. W. Meloche and C. Juday. 1933. The determination of the hydro¬

gen ion concentration of inland lake waters. Internat. Rev. ges. Hydrobiol.

u. Hydrog. 29:346-359.

Hutchinson, G. E. and Grace E. Pickford. 1932. Limnological observations on

Mountain Lake, Virginia. Internat. Rev. ges. Hydrobiol. u. Hydrog. 27:252-264.

Jenkin, Penelope M. 1932. Reports on the Percy Sladen expedition to some Rift

Valley lakes in Kenya in 1929. I. Introductory account of the biological survey

of five fresh water and alkaline lakes. Ann. and Mag. Nat. Hist. 9:533-553.

Jewell, Minna E. and H. W. Brown. 1929. Studies on northern Michigan bog

lakes. Ecology. 10:427-475.

Juday , Birge & Meloche- — Lake Waters of N. B. Wisconsin 67

Johnson, M. S. 1933. Preliminary report on some Minnesota lakes and their pro¬

ductiveness of fish food. Univ. Minn. Agric. Exp. Sta. Tech. Bui. 90. 31 pp.

Juday, C. and E. A. Birge. 1933. The transparency, the color and the specific

conductance of the lake waters of northeastern Wisconsin. Trans. Wis. Acad.

Sci., Arts & Let. 28: 205-259.

Kemmerer, G., J. F. Bovard and W. R. Boorman. 1923. Northwestern lakes of the

United States: Biological and chemical studies with reference to possibilities in

production of fish. Bui. U. S. Bur. Fish. 39:51-140. Doc. No. 944.

Kolthoff, I. M. 1931. The colorimetric and potentiometric determination of pH.

New York. 167 pp.

Kusnetzow, S. I. and A.L. Shtcherbakow. 1927. Physico-chemical data of Lake

Baikal. Trav. de la Com. pour 1’etude du lac Baikal. 3:193-211.

Maucha, R. 1929. Zur Frage der aktuelle Reaktion als Milieufaktor der Gewasser.

Verh. Internal . Ver. Limnol. 4:435-450.

Maucha, R. 1932. Hydrochemische Methoden in der Limnologie. Die Binnenge-

wasser. Bd. XII. 173 pp. Stuttgart.

Morrison, J. P. E. 1932. A report on the Mollusca of the northeastern Wisconsin

lake district. Trans. Wis. Acad. Sci., Arts & Let. 27:359-396.

Neresheimer, E. and F. Ruttner. 1929. Der Einfluss der Abwasser des Magnesit-

werkes in Radenthein auf den Chemismus, die Biologie und die Fischerei des

Millstatter Sees in Karnten. Zeitsch. Fisch. 27:47-66.

posting, H. J. 1933. Physical-chemical variables in a Minnesota lake. Ecolog.

Monogr. 3:493-534.

Philip, C. B. 1927. Diurnal fluctuations in the hydrogen ion activity of a Minne¬

sota lake. Ecology. 8:73-89.

Pia, J. 1933. Kohlensaure und Kalk. Die Biennengewasser. Bd. XIII. 183 pp.

Stuttgart.

Pierre, W. H. and J. F. Fudge. 1928. The adjustment of the reaction of indicator

solutions and its importance in determining the hydrogen ion concentration of

slightly buffered solutions. Jour. Amer. Chem. Soc. 50:1254-62.

Saunders, J. T. 1926. The hydrogen ion concentration of natural waters I. The

relation of pH to the pressure of the carbon dioxide. British Jour. Exper.

Biol. 4:46-72.

Schutow, D. A. 1926. Die Assimilation der Wasserpflanzen und die aktuelle Reak¬

tion des Milieus. Planta — Ark. Wis. Bot. 2:132-151,

Senior-White, R. 1927. On the relationship existing between carbonates and pH and

conductivity in natural waters. Indian Jour. Med. Res. 15:989-996.

68 Wisconsin Academy of Sciences , Arts, and Letters.

Seyler, C. A. 1894. Notes on water analysis. Chem. News. 70:82-3, 104-S, 112-14,

140-41, 151-52.

Skadowsky, S. N. 1926. Ueber die aktuelle Reaktion der Siisswasserbecken und

ihre biologische Bedeutung. Verh. Internal. Ver. Limnol. 3:109-144.

Strom, K. M. 1925. pH values in Norwegian mountains and their bearings upon

the classification of freshwater localities. Nyt Mag. f. Naturv. 62:237-244.

Werestschagin, G. J., N. J. Anickova and T. B. Forsch. 1932. Methoden der hydro-

chemischen Analyse in der limnologischen Praxis. Arch. f. Hydrobiol. 23:1-64,

167-230.

Wiebe, A. H. 1931. Diurnal variations in the amount of dissolved oxygen, alkalinity

and free ammonia in certain fish ponds at Fairport, Iowa. Ohio Jour. Sci.

31:120-126.

Worthington, E. B. 1930. Observations on the temperature, hydrogen ion concen¬

tration and other physical conditions of the Victoria and Albert Nyanzas.

Internat. Rev. ges. Hydrobiol. u. Hydrog. 24:328-357.

Yoshimura, S. 1932. On the dichotomous stratification of hydrogen ion concentra¬

tion in some Japanese lake waters. Jap. Jour. Geol. and Geog. 9:155-185.

Yoshimura, S. 1933. Kata-numa, a very strong acid water lake on volcano Katan-

uma, Miyagi Prefecture, Japan. Arch. f. Hydrobiol. 26:197-202.

Juday, Birge & Meloche — Lake Waters of N . E. Wisconsin 69

Table I

Mean quantity of free carbon dioxide in the surface waters of 238 seepage lakes.

The lakes are grouped by 0.5 mg. intervals; the number of lakes in the various groups

is indicated, together with the percentage of the total number. The results for free

carbon dioxide are expressed in milligrams per liter of water.

Table II.

Mean quantity of free carbon dioxide in the surface waters of 280 drainage

lakes. The lakes are grouped by 0.5mg. intervals. The number of lakes in each

group is indicated, together with the percentage of the total number. The means

are expressed in terms of milligrams of carbon dioxide per liter of water. The

minus sign indicates that the water gave an alkaline reaction to phenolphthalein,

which was due to the presence of a certain amount of normal or monocarbonate

carbon dioxide.

70 Wisconsin Academy of Sciences, Arts, and Letters.

Table III

The 7 drainage lakes which gave the highest alkaline reaction to phenolphthalein.

The color of the water is indicated on the platinum-cobalt scale. Carbon dioxide

and the organic carbon are stated in milligrams per liter of water. The minus sign

in the free carbon dioxide column shows that the water gave an alkaline reaction

to phenolphthalein due to the presence of normal or monocarbonate carbon dioxide.

Table IV

The seepage and drainage lakes which gave the maximum acid reaction to

phenolphthalein. The color of the various waters is indicated on the platinum-

cobalt scale. The free and bound carbon dioxide and the organic carbon are stated

in milligrams per liter of water.

Seepage Lakes

Juday , Birge & Meloche — Lake Waters of N. E. Wisconsin 71

Table V

Mean quantity of bound carbon dioxide in the surface waters of 238 seepage

lakes. The lakes are grouped by 1 mg. intervals; the number of lakes in each group

is indicated, as well as the percentage of the total number. The carbon dioxide

means are expressed in milligrams per liter of water.

Table VI

Annual variations in the bound carbon dioxide content of the surface waters

of 106 seepage lakes.

72 Wisconsin Academy of Sciences , Arts, and Letters ,

Table VII

Mean quantity of bound carbon dioxide in the surface waters of 280 drainage

lakes. The lakes are grouped by 1 mg. intervals; the number of lakes in each group

is indicated, as well as the percentage of the total number. The means are expressed

in milligrams of bound carbon dioxide per liter of water.

Juday , Birge & Meloche — Lake Waters of N. E. Wisconsin 73

Table VIII

Annual variations in the bound carbon dioxide content of the surface waters

of 136 drainage lakes.

Table IX

Calcium content of the various bound carbon dioxide groups of lakes. The

results are expressed in milligrams per liter of water. The lake waters are grouped

by 2 mg. intervals, and the maximum, minimum and mean quantities of calcium are

given for the various groups, based on surface samples of 357 lakes.

74 Wisconsin Academy of Sciences, Arts, and Letters .

Table X

Carbon dioxide and hydrogen ion results obtained on a number of lakes rep¬

resenting the different types. The quantity of carbon dioxide is stated in milligrams

per liter . The waters that gave an alkaline reaction to phenolphthalein are in¬

dicated by a minus sign in the free carbon dioxide column.

Juday, Birge & Meloche — Lake Waters of N. E. Wisconsin 75

Table X — Continued

76 Wisconsin Academy of Sciences, Arts, and Letters,

Table X — Continued

Juday, Birge & Meloche—Lake Waters of N. E. Wisconsin 77

Table XI

Carbon dioxide content and hydrogen ion concentration of some of the well

waters, together with similar data on the surface and bottom waters of the lakes on

which the wells are situated. The carbon dioxide is indicated in milligrams per liter.

The minus sign in the free carbon dioxide column indicates that the water gave an

alkaline reaction to phenolphthalein.

78 Wisconsin Academy of Sciences , Arts, and Letters,

Table XI — Continued

Table XII

The 149 wells are divided into groups on the basis of the bound carbon dioxide

content of their waters ; the mean quantity found in the different groups is indicated

in milligrams per liter of water.

Juday, Birge & Meloche — Lake Waters of N. E, Wisconsin 79

Table XIII

Comparison between the mean of the colorimetric pH readings and those obtained

with the quinhydrone system in 1932 .

Table XIV

This table shows the range of the hydrogen ion concentration (pH) in 1,225

surface samples obtained from 538 lakes and lakelets of northeastern Wisconsin.

80 Wisconsin Academy of Sciences , Arts , and Letters.

Table XV

The 238 seepage and 280 drainage lakes are grouped on the basis of the hydrogen

ion concentration of their surface waters. The mean pH has been used in classifying

the lakes that were visited more than once .

Table XVI

Annual variations of the hydrogen ion content of the surface waters of 245

lakes that were visited in more than one year during this investigation.

J uday, Birge & Meloche—Lake Waters of N. E. Wisconsin 81

Table XVII

The dichotomous stratification of the pH in Lake Mary on July 11, 1928.

Results for other determinations made on this date are given for purposes of com¬

parison. The results for carbon dioxide, oxygen, chloride and residue or total solids

are indicated in milligrams per liter of water .

Table XVIII

Mean quantity of free and bound carbon dioxide in milligrams per liter in the

various hydrogen ion groups . The table includes the results for 227 seepage and

272 drainage lakes. See figs. JO and 31.

82 Wisconsin Academy of Sciences , Arts, and Letters,

Table XIX

Relation between the hydrogen ion and the calcium content of the surface waters

of 358 lakes; the results for the latter are indicated in milligrams of Ca per liter

of water.

Table XX

Relation of the hydrogen ion to , the free and bound carbon dioxide of the well

waters. The free and bound carbon dioxide are given in milligrams per liter of water.

FURTHER NOTES ON THE OCCURRENCE OF PARASITIC

COPEPODS ON FISH OF THE TROUT LAKE REGION,

WITH A DESCRIPTION OF THE MALE OF

Argulus biramosus*

Ruby Bere

From the Limnological Laboratory of the Wisconsin Geological and Natural

History Survey. Notes and reports No. 56.

In conjunction with the general fish parasite work being

conducted by the Wisconsin Geological and Natural History Sur¬

vey at the Trout Lake Laboratory during the summers of 1931

and 1932, certain observations were made regarding the parasitic

copepods which, together with the material collected, have been

turned over to the writer. It was noted that these parasites were

present in greater numbers in 1932 than in 1931. In the former

year as many as 43 per cent of the ciscoes and 30 per cent of the

perch in Trout Lake were parasitized while in 1931 only about

one-half this per cent were infected.

As in 1930, the Ergasilids were present in greatest

numbers in the two succeeding years and again all the specimens

collected belonged to the species Ergasilus confusus . Achtheres

micropteri, only one specimen of which was collected in 1930,

was quite abundant in 1932 but not in 1931. The majority of

the specimens were attached to the roof of the mouth cavity

and the gill arches of rock bass caught in Muskellunge Lake-

Achtheres coregoni , however, was no more plentiful than in

1930; no specimens were noted during the summers of 1931 and

1932 but in October, 1932, two specimens were taken from the

fin of a whitefish caught in Trout Lake. In both 1931 and

1932 many Arguli were observed, whereas in 1930 only two

specimens were found. However, most of the Arguli occurred

on the sucker, only 13 of which were examined in 1930. Of

the 128 Arguli comprising the 1931 and 1932 collections, 16 per

cent have proved to be Argulus biramosus , the remainder being

A. catastomi. A. biramosus was found on rock bass and possibly

*This work was supported by a grant from the Wisconsin Alumni Research Foundation,

University of Wisconsin.

83

84 Wisconsin Academy of Sciences , Arts, and Letters .

also on wall-eyed pike, and A. catastomi on the sucker. In the

present instance the hosts of the these two species can only be

conjectured as the various species of fish were brought to the

laboratory in one bucket and many of the parasites had left the

fish and were swimming about in the water of the bucket.

In 1930 no parasitic copepods were observed on any of the

fish from the five soft water lakes included in the study. Dur¬

ing the two succeeding years fish from three of these lakes,

Nebish, Weber and Crystal, were examined for parasites but

no copepods were found, although the same species of fish harbor

them in the lakes with harder water. A total of 1088 fish has

now been examined from the three lakes just mentioned. These

lakes, as well as Nelson, and Geneva in which no parasitic cope¬

pods were found in 1930, are of the seepage type, i.e., they

possess neither inlets nor outlets. On the other hand, parasitized

lakes in which these parasitic copepods do occur are of the drain¬

age type. Clear Lake is also a seepage lake and during 1931

and 1932, 146 fish, the majority of which were ciscoes, were ex¬

amined but no copepods were found.

The question arises as to why these soft water lakes are

without parasitic copepods- Internal helminth parasities are

also quite rare. Other forms of life occur abundantly in Nebish

and Weber lakes and to a lesser extent in Crystal. There are

at least two possibilities. The parasites may never have been

introduced into these lakes for the fish with which the lakes

were originally populated may have been free of these para¬

sites and invasion by new fish is impossible since the lakes

possess neither inlets nor outlets. On the other hand, parasitized

fish may have been among the original population but the cope¬

pods may not have been able to adapt themselves to the chem¬

ical and physical conditions existing, or which subsequently de¬

veloped, in these isolated bodies of water. This phase of the

problem is open to experiment and might be answered by placing

parasitized and non-parasitized fish in cages and setting them in

the lakes mentioned above. E. confusus and the two species

of Achtheres do not leave the host after they have become at¬

tached but the larval stages are free living. Therefore, in order

for non-parasitized fish to become infected the parasites would

first have to pass through their life cycle in the open water.

B ere— Parasitic Copepods on Fish

85

In 1980 when Argulus biramosus was first described, no males

were available and a short description of the male is given now.

It displays the three specific characters of the female, on the

basis of which the species was established, viz. 1) the biramous

character of the terminal portion of the first antennae; 2) the

shape of the respiratory areas (Fig. 1) ; and 3) the character of

the supporting rods in the margin of the sucking disks. The

third and fourth legs are highly modified in structure and as

a result it is possible to distinguish the sexes with the naked

eye.

The basipod of the fourth leg carries a very large leaf¬

like appendage (Fig. 2) . The peg on the anterior margin of the

basipod is much like that of many other species. Posterior to the

basal portion of the peg is a small narrow area which is covered

with minute spines. The endopod is two jointed but the distal

segment is much reduced in size; this modified segment and

the very large flap on the basipod give the leg a very character¬

istic appearance.

Attached to the lateral ventral wall of the basipod of the

third leg is a large lobe which is fringed with long setae on its

posterior border and short spines on its outer border (Fig. 3).

A large scythe-like appendage, partly covered with spines, ex¬

tends from the anterior margin of the basipod. Arising from the

proximal basipod segment are two lobes which are thickly cov¬

ered with small spines and which lie on either side of the scythe¬

like structure, partially covering its base. The semen receptacle

is located in the posterior portion of the distal basipod segment

and anterior to it, on the dorsal surface, is a circular concave

area bearing many spines.

The basal joint of the second leg bears a flap which carries

three plumose setae at the outer anterior corner, but the poster¬

ior border and part of the lateral surface are thickly covered

with short spines (Fig. 4).

The specimens had been preserved in alcohol and are almost

colorless, no trace of the prominent markings found on the

thorax of the female being apparent.

The ten males range in length from 4-6.5 mm. and in width

from 3-5 mm. The 12 females range in length from 2.5-7 mm.

and in width from 2-5 mm.

S6

Wisconsin Academy of Sciences , Arts, and Letters .

Plate I

Fig. 1. Dorsal view of Argulus biramosus, male.

Fig. 2. Ventral view of fourth swimming leg.

Fig. 3. Ventral view of third swimming leg.

Fig. 4. Ventral view of second swimming leg.

TRANS. W1S. ACAD., VOL. 29.

PLATE I

88 Wisconsin Academy of Sciences , Arts , and Letters.

Bibliography

Bere, R. 1931. Copepods parasitic on fish of the Trout Lake Region, with descrip¬

tions of two new species. Trans. Wis. Acad. Sci., Arts and Let. 26:427-436.

Wilson, C. B. 1903. North American Parasitic copepods of the family Argulidae,

with a bibliography of the group and a systematic review of all known species.

Proc. U. S. Nat. Mus. 25: 635-742.

- 1907. Additional notes on the development of the Argulidae, with de¬

scription of a new species. Proc. U. S. Nat. Mus. 32:411-424.

- 1911. North American parasitic copepods belonging to the family Erga-

silidae. Proc. U. S. Nat. Mus. 39:263-400.

- 1915. North American parasitic copepods belonging to the Lernaeopo-

didae, with a revision of the entire family. Proc. U. S. Nat. Mus. 47:565-729.

TWO NEW SUBSPECIES OF FISHES FROM WISCONSIN

Carl L. Hubbs and C- Willard Greene

During the course of the survey of the Wisconsin fish fauna,

which we have been conducting for the Wisconsin Geological

and Natural History Survey, three new subspecies of fishes have

been discovered. One of these, Notropis nux richardsoni , was

described by us in 1926 (Hubbs, 1926: 39, pi. 3, fig. 2) ; origin¬

ally this form was regarded as a subspecies of N. heterodon, but

later studies showed that it was connected rather with N. nux

(see Hubbs and Greene, 1928: 372). The other new subspecies

are described in this report, under the following names :

Campostoma anomalum oligolepis

Boleosoma nigrum eulepis

The very interesting distribution and the probable origin

and history of these forms, will be treated in a subsequent paper

by the junior author.

Most of the counts and measurements as tabulated in this

paper were made for us by Dr. Canuto G. Manuel.

1. Campostoma anomalum oligolepis , new subspecies

Plate II, Fig. 1.

Holotype, a breeding male 92 mm. long to caudal, collected

by Greene and Stuart in Little Rib River 2% miles east of

Hamburg, Marathon County, Wisconsin, on June 19, 1927; Cat.

No. 75582, Museum of Zoology, University of Michigan. Num¬

erous paratypes are preserved in that museum and also in the

University of Wisconsin.

In identifying the many hundred specimens of Campostoma

anomalum collected in Wisconsin, it early became apparent that

two rather sharply differentiated forms occur in this state. Sub¬

sequent study of the species throughout its wide range has in¬

dicated that one of these forms, occurring in southwestern Wis¬

consin, is of wide distribution toward the southwest, as far as

Mexico. Its proper name appears to be Campostoma anomalum

pullum (Agassiz). A photograph of a nuptial male of C. a.

89

90 Wisconsin Academy of Sciences, Arts, and Letters .

pullum is reproduced in Plate II, Figure 2. The other form which

we have not yet encountered outside of Wisconsin, appears to

have never been named. Although it occurs here and there

(with C. a. pullum) through the southwestern part of the state;

its main range is to the northward and eastward. It occurs in

both the Lake Michigan and the Mississippi River basins. The

distinctive features of the two subspecies of Campostoma ano -

malum occurring in Wisconsin are listed in tabular form in

Table I.

Table I

Distinctive features of Compo stoma anomalum in Wisconsin.

C. a . oligolepis C. a. pullum

Scales:

Hubbs & Greene— New Subspecies of Fish 91

In addition to these numerous average differences, the two

forms oligolepis and pullum differ also very interestingly in dis¬

tribution, as the junior author will indicate in a later paper. As

already mentioned, the main range of oligolepis in Wisconsin is

to the north and east of the territory occupied by pullum . They

were taken together at only seven localities, as follows :

1. Tributary of West Pecatonica River, 3 miles east of Dar¬

lington, Lafayette County.

2. Mineral Branch of West Pecatonica River, just west of

Mineral Point, Iowa County.

8. Tributary to Dodge Creek, 5*4 miles northeast of Blan-

chardviile, Iowa County.

4. Crawfish River, 2 miles northwest of Fall River, Columbia

County.

5. South Branch of Turtle Creek, at Allen Grove, Walworth

County.

6. Lake Geneva outlet, 3 miles east of Lake Geneva city, Wal¬

worth County.

7. Bark River, Merton, Waukesha County.

These localities all lie in the territory largely occupied by

C. a. pullum , and in each of these collections that subspecies

predominates, except in the lot from Lake Geneva outlet where

two specimens of each were caught. There is no real evidence

of intergradation even in these localities. The scale counts

for each form, where the two occur together (Tables II and III),

are almost typical, though apparently showing a slight approach

toward the mean of the other subspecies (which might reflect

some hybridization in the past). Furthermore, the other char¬

acters, such as contours of body, shape of head and size of mouth,

confirm the separation. These other characters are all of aver¬

age or usual significance ; even when taken together, they do not

invariably characterize individuals. Occasionally whole series

of either subspecies show the form characters of the other. With¬

in all of the same stream systems listed above, pure stocks of

both subspecies were also taken.

92 Wisconsin Academy of Sciences, Arts, and Letters.

Table II

Summary of scale counts in Wisconsin specimens of Campostoma anomalum.

In each column the median set of counts is for lateral line scales, the upper set for

scales around the body and the bottom set for the sum of these two counts ; the

circumference counts in the column for C. a. pullum, where occurring alone, are

given to the left of the lateral line counts.

Hubbs & Greene — New Subspecies of Fish

93

The two Wisconsin forms of Campostoma are regarded as

subspecies despite the fact that we have been unable to find

good evidence that they intergrade. This course is followed be¬

cause in other localities we find the distinctive features to break

down. Thus in the Ohio Valley form, typical C. a. anomalum ,

the scales are intermediate in number. (See Table III).

In fact, the distinction of C. a. oligolepis as individuals from

C. a. anomalum is often difficult, though the average characters

are sufficiently different to enable most series to be distinguished

rather readily. In addition to having the scales usually some¬

what fewer, oligolepis usually has the mouth somewhat smaller

(Table IV). The form of body, head and mouth is very similar.

C. a. anomalum differs only in average characters from C. a.

pullum ; the distinction of individuals often appears impossible

and the identification of series is difficult at times. These differ¬

ences involve scale size (Table III), mouth size (Table IV) and

shape and form of body and head. The characters of form, in

which typical anomalum resembles oligolepis closely, are most

subject to variation.

94 Wisconsin Academy of Sciences , Arts, and Letters.

Table III

Average of scale counts for different lots of C am po stoma anomalum. The

number of specimens on which each average is based is indicated in parenthesis .

Since C. a. anomalum appears to be incompletely differenti¬

ated from C. a. oligolepis on the one hand and from C. a. pullum

on the other hand, it seems logical to regard both oligolepis and

pullum as subspecies of C. anomalum, even though they them¬

selves appear to intergrade very slightly or not at all. When

other subspecies of anomalum (yet unnamed) are considered, the

treatment of oligolepis as a subspecies appears to be almost

demanded.

Description (based primarily on nuptial male holotype 92

mm. long to caudal, supplemented by measurements in paren¬

thesis of numerous paratypes). — This is usually an elongated

by rather thick fish: greatest depth 4.7 (4.3 to 4-7, usually about

4.5) in standard length; least depth of caudal peduncle 2.6 (2.05

to 2.8, usually 2.2 to 2.5) in head, 2.7 (2.0 to 3.0, usually 2.4 to

2.7) in depth of body, measured over curve of side, and 2.4 (2.1

to 2.6) in standard length of caudal peduncle. The anterior dor¬

sal profile is usually rather flattish, less arched than in pullum •

The head tends toward a squarish form (relatively speaking),

flattened above, broad, and rather globose in the muzzle region :

length of head 3.8 (to 4.3, usually about 4.0) in standard length;

depth of head, from occiput to isthmus, 1.4 (1.35 to 1.6, usually

about 1.5) ; width of head 1.8 (1.5 to 1.8, usually about 1.65) in

length of head; least bony interorbital width 3.5 (3.05 to 3.8,

usually 3.3 to 3.5) ; postorbital length 2-1 (2.0 to 2.25) ; sensory

lines in frontal region of skull nearly parallel, not so strongly

convergent and approximated anteriorly as in pullum ; suborbital

Hubbs & Greene — New Subspecies of Fish 95

width 5.0 (5.1 to 7.0) ; eye (cornea) 5.9 (4.2 to 6.0) ; snout 2.65

(2.6 to 2.95) ; length of upper jaw 3.1 (3.0 to 3.8) ; width of

mouth including lips 2.7 (2.7 to 3.35) ; width of gape, excluding

outer lip, 3-6 (3.9 to 5.2, usually 4.3 to 4.8 in non-breeding males

and females).

Distance from tip of snout to origin of dorsal 2.05 (2.0 to

2.1) in standard length; snout to pelvic 2.25 (2.05 to 2.2) ; in¬

sertion of pelvic to origin of anal 1.05 (1.05 to 1.35) in distance

between insertions of paired fins; longest dorsal ray 1.4 (1.2

to 1.5) in head; length of depressed dorsal 1.3 (1.3 to 1.8) in

distance forward to occiput; dorsal base 2.0 (2-0 to 2.65) ; length

of caudal fin 1.15 (0.95 to 1.1) in head; length of depressed anal

1.3 (1.3 to 1.5) ; anal base 2.15 (2.3 to 2.9 in non-breeding males

and females) ; length of pectoral fin 1.05 (1.1 to 1.5) in inter¬

space between insertions of paired fins; length of pelvic fin 1.2

(1.1 to 1.6) in pelvic-anal interspace.

Dorsal with 8 principal rays (constant) ; anal 8 in holotype,

but normally with 7 ; caudal 19 (18 to 20) principal rays; pec¬

toral 17 (13 to 17) ; pelvic 8 (rarely 9).

For scale counts refer to Tables II and III. The scale struc¬

ture) is about the same as in pullum. The focus is far basad.

Table IV

Width of gape in three subspecies of Campostoma anomalum. measured between

angles of gape, not across outer lips; measured into head .

C. a. anomalum C. a . oligolepis C. a. pullum

fisual variation in half-

grown and adults . 3.7 to 4.4 4.3 to 4.8 4.6 to 5.5

Extreme variation in half- z

grown to adults . . . 3.4 to 4.7 3.9 to 5.2 4.4 to 6.2

In large breeding males,

about . 3.3 3.6 3.8

The lateral field is much reduced by the encroachment of the

apical field, which bears numerous radii between which the

ridges are curved. An anterolateral angle may be fairly well

marked or absent. The lateral radii run obliquely to the scale

margin.

Gill-rakers 22 (19 to 24). Pharyngeal teeth 4-4 in all speci¬

mens examined, with an obliquely truncated grinding surface

and obsolete hook. Intestine coiled around air-bladder as usual

in the genus.

96 Wisconsin Academy of Sciences, Arts, and Letters.

The general color and color pattern is similar to that of the

other forms, with average differences noted in Table I. The

breeding male shows the usual large subbasal blackish markings

and orange shades on the fins, possibly more intensely than in

pullum.

The nuptial tubercles are similar to those of other subspecies.

Very large, suberect ones, with swollen bases, occur on top of

head and on sides, as shown in the figure. Much smaller soft

tubercles or papillae occur on all surfaces of the head, and are

enlarged and uniserially arranged along each branchiostegal ray

and on adjacent opercular edge. The tubercles on the upper

surface of the pectoral rays are moderately large, slightly hooked

forward and arranged in one series basally branching one dis-

tally. Tubercles are also scattered in one series, along the an¬

terior dorsal rays, strongest on last unbranched ray. At most

barest traces of tubercle^ can be detected on the caudal fin, or

on the thickened rays of anal and pelvic fins.

The name oligolepis emphasizes the character of low scale

number, which reaches its extreme in this subspecies*

2. Boleosoma nigrum eulepis, new subspecies

Plate III, Figs. 1, 2, 3.

Holotype. — An adult 45 mm. long to caudal, collected in a trib¬

utary of the West Branch of Rock River, 2 miles north of At¬

water, Dodge County, Wisconsin; Cat. No. 77225, Museum of

Zoology, University of Michigan.

A large number of paratypes from Wisconsin, Minnesota and

Iowa, grading in standard length up to 61 mm., are in the col¬

lections of the universities of Michigan and Wisconsin.

Forbes and Richardson (1909: 297-298) discovered an ex¬

tremely interesting trend of variation in Boleosoma nigrum in

Illinois. We quote their remarks:

In studying our collections, wide variation was noticed with respect to the

scaly covering of the breast and cheeks, ranging from complete nakedness to com¬

plete scaliness of both, and also a considerable variation in robustness of build. While,

generally speaking, specimens become more scaly northward and more slender south¬

ward, it was not possible to make out, even approximately, any line or area of divi¬

sion, either general or local, between the two forms, or to draw any definite dividing

Hubbs & Greene- — New Subspecies of Fish

9?

line among the variants themselves. This confusion of conditions may be illus¬

trated by the following analysis of a single collection of forty-six specimens (Ac¬

cessions No. 28180) obtained from the north fork of the Vermilion River in Ver¬

milion county June 6, 1901.

Variations of Boleosoma nigrum (46 specimens)

It was also impossible to distinguish any correlation, even approximately constant,

between robustness of form and scaliness of cheeks and breasts, both stout and

slender forms having these parts sometimes naked and sometimes more or less covered

with scales. The larger percentage of specimens with scaly breasts and cheeks came

from the Rock River basin, from the northwest district, and from the Lake Michigan

drainage; but in all these districts scaly and naked specimens were intermingled, the

latter preponderating. In collections from the Kaskaskia, the Saline, the Cache, and

the lower Wabash Valley, on the other hand, both cheeks and breasts were also in¬

variably naked, while in the upper Wabash streams and in the Illinois basin the two

forms were indiscriminately commingled. The larger number of the stouter specimens

came from the Rock River system and the northwest area, while those from the

Kaskaskia, the Cache, and the Saline were of more slender proportions, with the

depth usually nearer six times than five times the length. A similar study of speci¬

mens from a wider range would probably show that Illinois is in a region of tran¬

sition between two varieties of this species — the typical nigrum , with slender body

and naked breast and cheeks, and some scaly-cheeked variety, probably olmstedi ,

or perhaps identical with it.

Our work has disclosed the form toward which Forbes and

Richardson found Boleosoma nigrum grading in northern Illinois.

This form can not be B. n. olmstedi as Forbes and Richardson

rather naturally supposed, because it is separated from olmstedi

98 Wisconsin Academy of Sciences , Arts, and Letters .

by a wide band of ordinary nigrum through Ontario, Lake Erie,

Ohio and Kentucky.

B. n. eulepis in typical form we find to be limited to the

glacial lake districts of Wisconsin (Lake Michigan and Missis¬

sippi Eiver drainage basins), Minnesota, Iowa and probably of

Illinois. In areas of these states where glacial lakes are numer¬

ous, the Boleosoma population is almost consistently of the eulep¬

is type. The range of the subspecies is discontinuous. Around

each of its territories, eulepis intergrades extensively, completely

and rather regularly with the surrounding and wide-spread B.

n. nigrum.

B. n. eulepis differs from the Atlantic drainage form B. n.

olmstedi in having fewer dorsal rays (9 to 14, usually 11 or 12,

instead of usually 13 to 15), in having fewer scales on the aver¬

age (48 instead of about 50 in lateral line to caudal), and the

soft dorsal lower and less finely tessellated in breeding males.

From typical B. n. nigrum, as represented in the Ohio Valley,

and elsewhere, eulepis differs in its larger size, decidedly more

robust form, darker color, and perhaps in having more dorsal

soft rays on the average. Occasionally we encountered such

extreme types of B. n. nigrum in Wisconsin, notably in a few

very isolated colonies within eulepis territories. But as a rule,

there is virtually no difference between B. n. nigrum and B. n-

eulepis within Wisconsin, in characters other than the degree of

completeness of squamation. The number of scales in the lateral

line may average very slightly higher, and the soft dorsal fin

may average a trifle higher, but the differences are of doubtful

statistical significance and are wholly unusable for identification

purposes. The data are given in Table V.

Table V

Comparison of counts and measures of the two forms of Boleosoma nigrum in

Wisconsin. The figures refer to number of specimens showing each count or

measurement.

Hubbs & Greene— New Subspecies of Fish

99

B. Degree of sqmmation of nape

Degree scaled Slightly Moderately Completely

Grade . . . 12 3

B, n. nigrum .......................................... 2 202

Intergrades . . . .... 31 33

B, n. eulepis . . 1 19 110

C. Degree of sqmmation of opercle

Degree scaled Slightly Moderately Completely

Grade . . . 12 3

B. n. nigrum . . 59 146

Intergrades . . 14 49 1

B. n. eulepis .......................................... 15 102 13

D. Degree of sqmmation of cheek

Degree scaled Naked Slightly Moderately Completely

Grade .......................... 0 12 3

B. n. nigrum .............. 205

Intergrades ................ 49 11 4

B. n. eulepis .............. 22 31 70 7

E. Degree of sqmmation of breast

Degree scaled Naked Slightly Moderately Completely

Grade . . 0 12 3

B. n. nigrum .............. 201 4 .... 1

Intergrades . . 31 17 7 8

B. n. eulepis . 4 9 5 112

F. Degree of sqmmation of all four areas

(Figures obtained by adding “Grade” of squamation for each area in each specimen)

Degree scaled 2 3 4 5 6 7 8 9 10 11 12

B. n. nigrum 2 57 141 4 .... 1

Intergrades .... 5 20 12 14 3 4 2 4

B. n. eulepis .... 267347 24 65 47

G. Number of dorsal soft rays

9 10 11 12 13 14

B. n. nigrum .......... .... 4 69 122 10

Intergrades . . . .... 1 27 29 7

B, n. eulepis .......... 1 3 53 59 12 1

H. Highest dorsal soft ray measured into head

1.5 1.6 1.7 1.8 1.9 2.0 2.1 2.2

B. n. nigrum ...... 5 13 33 52 36 54 8 3

Intergrades ........ 1 4 7 19 10 22 1

B. n. eulepis. .... 3 14 21 27 15 38 8 3

I. Highest dorsal spine measured into head

1.7 1.8 1.9 2.0 2.1 2.2 2.3 2,4 2.5 2.6 2.7 2.8 2.9 3.0 3.1

B. h. nigrum ............ 1 3 8 45 56 52 25 7 8 1 1 .... .... .... 1

Intergrades . . 4 3 3 21 16 9 7 .... 1 .... .... .... .... .... ....

B. n. eulepis ............ 2 4 1 38 31 27 18 1 3 .... .... . . .

100 Wisconsin Academy of Sciences , Arts, and Letters.

J. Length of head measured into standard length

3.2 3.3 3.4 3.5 3.6! 3.7 3.8 3.9 4.0 4.1 4.2 4.3 4.4 4.5

B. n. nigrum . . . . 1 1 19 36 49 45 30 7 11 1 2 3 .... 1

Intergrades . . . . . . 2 14 12 16 11 2 7 . ....

B. n. eulepis . . . 1 .... 6 31 37 36 16 2 1 . . . .

K, Depth of body measured into standard length

4.5 4.6 4.7 4.8 4.9 5.0 5.1 5.2 5.3 5.4 5.5 5.6 5.7 5.8 5.9 6.0 6.1 6.2 6.3

B. it. nigrum . .... 2 1 4 4 26 39 23 30 28 17 16 4 6 2 .... 1

Intergrades ...... . . . 4233855 11 48316 .... 1 ....

B. n. eulepis .... 1 . . . 1 2 6 7 10 20 13 23 11 11 11 7 6 .... 1 ....

There is also a very noticeable difference in the roughness of

the scales, readily appreciated when one rubs typical examples

of the two forms toward the head. The difference is due to the

longer marginal ctenii of B. n. eulepis , as is easily seen under

the microscope. The ctenii in B. n. eulepis resemble those of B.

podostemone 1 as figured by Cockerell and Elder (1913: 157),

while the ctenii of B. n. nigrum resemble those of nigrum or

even those of olmstedi as figured by the same authors.

The usual variation in B. n- eulepis in some additional char¬

acters, of little apparent taxonomic significance, are ; least depth

of caudal peduncle 2.6 to 3.0 in head; head and trunk (to anus)

1.7 to 1.8 in total length; first dorsal base 1.1 to 1.5, second

dorsal base 1.1 to 1.4 and anal base 1.75 to 2.1 in head; highest

dorsal spine 1.1 to 1.3 in highest dorsal soft ray; pectoral fin

1.0 to 1.1, pelvic fin 1.4 to 1.5 and caudal fin 1.1 to 1.4 in head.

Width of head 1.55 to 1.75, snout 3.6 to 4.25, orbit 3.6 to 4,35,

eye 3.8 to 5.0, snout and eye 2.0 to 2.3, interorbital 8.2 to 9.5,

cheek 3.35 to 3-9, upper jaw 3.4 to 4.4 in head; least distance

from lateral line to anterior dorsal contour 2.0 to 2.1 in post¬

orbital. Scales 4 or 5 above and 5 to 7 below lateral line. Dor¬

sal spines usually 8 or 9; pectoral rays 11 or 12; anal I, 7 to I,

10 ; principal caudal rays 15.

TRANS. WIS. ACAD., VOL. 29.

PLATE II

Fig. 1. Holotype of Campostoma anomalum oligolepis.

Fig. 2. Nuptial male of Campostoma anomalum pullum .

TRANS. WIS. ACAD., VOL. 29

PLATE III

Fig. 1. Ventral view of the holotype of Boleosoma nigrum eluepis showing

completeness of squamation.

Fig. la. Ventral view of specimen of Boleosoma nigrum nigrum from Wisconsin.

Fig. 2. Dorsal view of the holotype of Boleosoma nigrum eulepis.

Fig. 2a. Dorsal view of a specimen of Boleosoma nigrum nigrum from Wisconsin..

Fig. 3. Lateral view of the holotype of Boleosoma ngrum eluepis.

Fig. 3a. Lateral view of a specimen of Boleosoma nigrum nigrum from Wisconsin.

Hubhs & Greene-New Subspecies of Fish

101

Literature Cited

Cockerell, T. D. A. and Mary Esther Elder. 1913. The validity of Boleosoma

olmstedi (Storer) as indicated by its scales. Bui. Bur. Fish. 32:157.

Forbes, Stephen Alfred and Robert Earl Richardson. 1909. The fishes of Illinois.

Nat. Hist. Survey Ill. 3:cxxi-f357 pp.

Hubbs, Carl L. 1926. A check-list of the fishes of the Great Lakes and tributary

waters, with nomenclatorial notes and analytical keys. Misc. Pub. Mus. Zool.

Univ. Mich. 15:1-77.

Hubbs, Carl L. and C. Willard Greene. 1928. Further notes on the fishes of the

Great Lakes and tributary waters. Pap. Mich. Acad. Sci. Arts and Let. 8:371-

392.

GROWTH OP THE YELLOW PERCH ( Perea flavescens

Mitchill) IN NEBISH, SILVER AND WEBER LAKES,

VILAS COUNTY, WISCONSIN

Edward Schneberger

From the Limnological Laboratory of the Wisconsin Geological and Natural

History Survey.* Notes and reports No. 57.

Introduction

For nearly a decade the Wisconsin Geological and Natural

History Survey has been carrying on limnological investiga¬

tions on the lakes of the Highland Lake District of North¬

eastern Wisconsin. These investigations have been extended

to over 500 lakes in which there are a variety of environmen¬

tal conditions. Because of these facts this region is particu¬

larly favorable to the study of the growth of fresh water fish

in relation to their environmental conditions. With the co-op¬

eration of the U. S. Bureau of Fisheries, the Survey began

such a study in the summer of 1927, and these studies have

been continued to the present time. In 1930, six type lakes

were selected, and collecting limited to these lakes. Dr. Ralph

Hile of the Bureau of Fisheries is studying the ciscoes, rock

bass and bluegills, while the author is studying the perch and

gamefish. Recently, Mr. W. A. Spoor has undertaken to in¬

vestigate the ecology and growth of the common sucker. The

present paper deals with the growth of the perch from three

of the selected type lakes.

The author takes this opportunity to express his thanks to

Dr. E. A. Birge and Dr. Chancey Juday for allowing him to

work on the problem while in the employ of the Survey; to his

colleague, Dr. Hile, who has been very generous with helpful

suggestions, and to Mr. H. J. Deason of the Bureau of Fisher¬

ies for the advice and plans for the construction of a micro¬

projection apparatus.

*This investigation was made in co-operation with the U. S. Bureau of Fisheries and the

results are published with the permission of the Commissioner of Fisheries.

103

104 Wisconsin Academy of Sciences , Arts , and Letters.

METHODS

The fish were caught in gillnets and by hook and line fish¬

ing. Some of the smaller sizes were taken with a minnow

seine. Records were kept for all gear except the 1928 col¬

lections. The fish were taken to the field laboratory to be

measured and weighed as soon as possible after having been

removed from the nets. The standard length of each fish was

taken to the nearest millimeter by means of a steel tape. Weights

were taken to the nearest gram by means of a Chatillon spring

balance. These data were recorded on standard scale envelopes

furnished by the Bureau of Fisheries. Along with these data

were also placed the following : Field Number, Species, Locality,

Date, Sex, State of Organs, and Gear. A few scales were

taken from the left side of the fish near the middle of the body

and just above the lateral line, and placed on the inside of the

envelope in order to study the scales.

Typical scales were selected from the sample, soaked and

cleaned in water and then mounted in glycerine jelly prepared

after Van Oosten (1929). Later it was learned that they could

be mounted in ordinary white Karo syrup with greater ease and

with quite satisfactory results.

The scales were projected upon a piece of ground glass of

an apparatus similar to that described by Van Oosten (1928).

The 1932 material was studied, however, by means of an appar¬

atus fashioned after that used at present by the Bureau of

Fisheries at Ann Arbor, Michigan. The projected scale was

measured from the focus outward along the inter-radial space

to each annulus and to the outside circulus. Length of previous

years of life were calculated by the formula given by Van Oosten

(1923).

Due to the fact that additional and valuable material has

been collected during the past summer (1933), the validity of

the scale method and the ratio between the body and scale are

to be considered in a later paper. Space and time do not permit

the inclusion of this material in the present report. The calcu¬

lated lengths of the fish at the end of each year of life previous

to capture are given in the tables for the fish from each lake, but

wherever comparisons are made, they are based on the average

actual standard length of the specimens as much as possible.

Schneberger — Growth of Perch

105

Description of the Lakes

The perch for this study were obtained from Nebish, Weber,

and Silver Lakes. These three lakes do not have inlets, but

Nebish and Silver have outlets which function only during per¬

iods of high water.

Table I, taken from the records of the Wisconsin Geological

and Natural History Survey, gives a brief description of these

lakes. The physical and chemical data cited are the means of

a large number of determinations of surface waters. Silver

Lake is the largest ; its surface area is 22 times as large as that

of Nebish Lake, and 5.6 times as large as that of Weber Lake.

Nebish is almost twice as large as Weber. In volume, Silver is

4.4 times as large as Nebish, and 8.7 times as large as Weber.

All three lakes are deep enough for thermal stratification in the

summer. Weber Lake has the highest degree of transparency,

as the Secchi disc can be seen to an average depth of 7.2 meters.

The visibility of the same disc is lost at an average of 6.0 meters

in Nebish, and 5.5 meters in Silver. In regard to conductivity

and bound CO, Silver Lake is the highest and Weber the low¬

est. Nebish and Weber have an equal amount of organic matter

in the centrifuge plankton (0.77 mg/1), while Silver contains

somewhat more (0.85 mg/1). On the whole, the conditions for

the production of fish food in Silver Lake are more favorable

than in the other lakes.

Besides perch these lakes contain other species of fish. Weber

contains chiefly perch and small-mouthed bass. Other species

have been taken, but as the catches were infrequent and in small

numbers these are considered as remnants of bait discarded by

fishermen and not as an established species. Nebish Lake con¬

tains perch, rock bass, large- and small-mouthed bass, and a

few minnows. The following species are found in Silver Lake :

perch, rock bass, bluegills, suckers, ciscoes, small-mouthed bass

and muskellunge. Minnows and darters are also present. The

perch occurs more abundantly in this lake than the other species,

and is also more abundant in this lake than in the other two

lakes. The rock bass population of Nebish Lake is very large.

Weight Length Relationship

The weight and length of fishes are closely correlated. This

relationship is described within reasonable limits of variation

106 Wisconsin Academy of Sciences , Arts, and Letters.

Table I

Some physical and chemical characteristics of Nebish, Weber and Silver Lakes .

by the formula W = cL3 ; W = weight, L = length, and c is a

factor which varies for any species only as the weight fluctuates

in proportion to the cube of the length. It is assumed that the

specific gravity and form of the species do not change materially.

The factor c then is an index to the condition of the specimen and

is called the “coefficient of condition”.

In order to simplify the formula, another factor, K, is intro¬

duced and is equal to 100c. Substituting this in the original

formula gives

where W is expressed in grams and L in centimeters.

The exactness of the proportion of the weight to the length

has been considered by several authors. Crozier and Hecht

(1914) and Hecht (1913, 1916) conclude that, as a rule, the

weight is proportional to the cube of the length. More recent

authors, however, have found that the weight is proportional to

a power somewhat higher than the cube of the length.

Keyes (1928) shows that the weight of the herring, Fundulus

and sardine increases more rapidly than the cube of the length.

The value of the exponent is always greater than 3 and less than

4. This author concludes that the more rapid increase of the

weight is due to changes in form rather than in specific gravity.

Since fishes are in close hydrostatic equilibrium with their en¬

vironment, there is little or no chance of a change in specific

gravity. Similar differences were found by Clark (1928), Hart

(1931), Tester (1931), and others for different species.

S chneb erg er— Growth of Perch

107

The relationship between weight and length for the perch

from Nebish and Weber lakes (1932 material) was obtained by

plotting the average weights and lengths on double logarithmic

paper. A straight line was drawn through the points and the

slope determined. It was found that the weight increased at a

slightly higher rate than the cube of the length. For Weber

Lake the value of the exponent is 3.10 and for Nebish it is 3.19.

For the purposes of this paper the approximate formula is suffi¬

ciently accurate.

The chief factors that are reported as influencing the weight

of fish are sexual maturity and the condition of the individual

(fatness). In regard to sex, Heincke (1907), and Crozier and

Hecht (1914) found that sex does not influence the constant

ratio between weight and length. Heincke’s constant, “Ernah-

rungskoeffizient”, however, showed seasonal fluctuations.

Hecht (1916) found it unnecessary to use a correction fac¬

tor for the weight of the contents of the stomach. In the sardine,

Clark (1928) found that the undigested food in the alimentary

tract had no effect on the weight of the fish. The weight-length

factor fluctuated in the same way for the eviscerated fish as it

did for the total weight.

As these collections were made during the summer months,

the fish were well through their spawning ; so it was decided to

determine the effect, if any, of the contents of the alimentary

tract on the value of K. This factor was computed for fish with

different percentages of undigested food in the alimentary tract.

The results are shown in the following table :

Weber Lake, 1932

The probable error of the difference of the means (Edm) for

the mean K of the empty fish and the 1-24% is 0.0180 ± 0.14.

108 Wisconsin Academy of Sciences, Arts, and Letters.

When the mean K of the 25-50% full fish is compared with the

mean K of the empty fish, the Edm is 0.0713±0.153. In both

comparisons the difference of the means is smaller than the cor¬

responding probable error, indicating that the difference is in¬

significant.

Since the perch in Weber Lake feed largely on insect larvae,

it was thought that perhaps the weight-length factor might be

influenced by a different type of food. In Silver Lake the perch

had been feeding on small perch; therefore, comparisons are

made on perch with different percentages of fish in their aliment¬

ary tract. The results of these comparisons are as follows : the

Edm of K between the empty and 25-50% contents for Silver

Lake is 0.094 ± 0-137. A comparison of the mean K for all fish

with the 25-50% group shows that the Edm is 0.055 ± 0.136.

A similar. comparison of the mean K of the empty fish and the

mean K of all fish gives an Edm of 0.039 ± 0.110, which shows

that there is no significant difference. From these data it can be

concluded that the undigested food in the alimentary tract has

no appreciable effect on the value of the weight-length factor. It

is also true that the type of food present has no direct effect upon

this value.

Table II shows the average K for each age-group for the years

1930-1933, inclusive. The table reveals that for the most part

there is a general decline in the value of K for Nebish and Weber

Table II

Average K of each age-group from each lake.

* VIII /Weber (1932), 1.54; Silver (1932), 1.58.

Schneberger — Growth of Perch

109

lakes. In Silver Lake the average of K declines slightly in 1931,

followed by an increase in 1932. The increase is not sufficient,

however, to equal the value of K for 1930. It will also be seen

that the perch from Nebish are in the best condition while those

from Silver Lake are in the poorest condition. Weber Lake forms

an intermediate group. The same situation is found in the rate

of growth, as will be seen later.

Sex Ratio

In many instances various authors have reported that one of

the sexes is more numerous than the other. The variation de¬

pends somewhat upon the species; that is, the males are more

numerous in some species, while in others the females are more

numerous. For the perch of Lake Erie, Jobes (1932) makes no

statement regarding sex ratio, but gives a table (II, p. 647) of

the males and females of age-groups II and III taken in 1927 and

1928. His data show substantially a 1:1 sex ratio in age-groups

II and III in Lake Erie. This table includes 476 perch, of which

240 were females and 236 were males.

In Nebish Lake the males are slightly more numerous than

the females. The ratio of females to males is 1 :1.26. In Weber

and Silver lakes the reverse is true, as there are more females

than males. In these two lakes the male to female ratio is 1 :1.31

and 1:1.40, respectively. Table III gives the number in each

length frequency class for each of the three lakes. In Nebish

Lake the sexes are almost equally distributed through all sizes.

In Weber Lake the larger groups, 215-245 mm., contain only fe¬

males; no males larger than the frequency value of 205 were

found. There is a sharp decline in the number of males after

the 145 mm. group.

In Silver Lake the 115-135 mm. length frequency classes

contain 94% of all males. Of the larger males, the 165 and 185

mm. groups contain only one male each. From the data in this

table it is seen that the males are more numerous than the fe¬

males in the smaller sizes, but they are less numerous in the

larger sizes. The number of males decreases with increasing

size.

110 Wisconsin Academy of Sciences , Arts , and Letters.

Table III

Sex compostion of the 1932 catches.

* Mid-points of classes.

Schneberger — Growth of Perch

111

Gill Net Selectivity

Since gill nets are known to be highly selective in the size

of fish they take, a variety of sizes of mesh was used in the col¬

lecting. During the seasons 1931 and 1932 the following sizes

of mesh were used: %, %, %, 1, 1%, 1 %, and 1*4 inches.

These measurements are bar measure instead of stretched mesh.

The % inch bar measure is equivalent to 1% inch stretched

mesh, etc. The meshes were made from linen thread and each

net was six feet deep and 150 feet long. In each setting the

nets were tied together to form a gang in which each mesh size

was represented. When the fish were removed from the net they

were placed in a container upon which was painted the corres¬

ponding net size. The size of the gear was recorded on each of

the scale envelopes.

Not only are gill nets known to be selective in the size of

fish they take, but they may take only the larger individuals of

one age-group and the smaller individuals of the next larger

age-group. Sometimes a net will take specimens that are very

much above or below the average size of that net. In these cases

the fish are not gilled but tangled in the meshes, or, in the case

of larger specimens, they are caught by some structure such as

the parts of the mouth. These are accidental catches and are

not frequent enough to play a part in the selectivity of the net.

Table IV

Gill net selectivity. Number of perch per age-group per mesh size.

Nebish Lake , 1932.

112 Wisconsin Academy of Sciences, Arts, and Letters.

Figure 1 and Table IV show the agesgroups and lengths of

fish taken by the different sizes of gill nets from Nebish Lake.

In the graphs in Figure 1 are shown the length frequency curves

for each mesh size for each of the age-groups. There is an over¬

lapping of sizes of fish in each of the age-groups. The net-

length frequency curves for each of the age-groups show that

the maximum length of fish taken by the smaller sizes of mesh

is overlapped by the minimum lengths of fish taken by the next

larger size. Only two nets, the % and % inch mesh, took fish

belonging to age-group I. The % mesh took the smaller speci¬

mens, and the % inch mesh took the larger specimens of this

group. In age-group II, four mesh sizes (%, %, %> and 1 inch)

functioned in taking specimens. Although the % and 1 inch

nets took only a few fish, they took specimens of the two extreme

limits of size of the age-group. While the % and % took the

bulk of the age-group, neither would be sufficient alone to take a

representative sample of this age-group, because the majority

of the fish taken by the % inch mesh fall below the modal length

of the age-group, and those taken by the % inch net fall well

above the modal length.

Fig. 1. Catch per net by age-groups. Nebish Lake, 1932.

In age-group III the % and 1% inch meshes took fish of the

two extreme lengths, while the % and 1 inch took most of the

fish of this group. Here again the % inch specimens fall below

Schneherger — Growth of Perch

113

the mode and the 1 inch fall above. Age-group IV is represented

by only five specimens taken by nets. Of these, three were

taken by the 1 inch mesh and two by the lVs inch mesh. The

1% inch meshes caught but one fish, which belonged to age-

group III. No perch were taken by the IV2 inch net.

Table V shows the relation between mesh size and length fre¬

quencies of the various age-groups of perch taken from Silver

Lake. This table shows that the smaller fish of one age-group are

taken by one net size, and that the larger specimens of the same

group are taken by the next larger size of mesh.

Table V

Gill net selectivity. Number of perch per age-group per mesh size.

Silver Lake , 1932

From the foregoing data and discussion it can be concluded

that on the whole the sampling of the different age-groups has

been adequate- Although the individual nets show a decided

selectivity, the difficulty was surmounted by having nets of

various sizes. The difference in sizes of mesh was small enough

so that no large gaps in ages and sizes occurred.

Not all of the perch collected for this study were taken

by gill nets, as hook and line fishing was carried on to some

extent. The most extensive collecting by hook and line fishing

was carried on in Silver Lake in 1931. Of the 463 perch taken

from this lake, 220 were taken by hook and line fishing. The

remaining 243 were taken by the gill nets. Table VI shows

the length frequencies of each age-group caught by hook and line

compared with those taken by gill nets.

114 Wisconsin Academy of Sciences , Arts , and Letters .

All age-groups with the exception of VI and VII are fairly

well represented by the hook and line catches. Since there

are but 220 specimens represented in seven age-groups, it

seems that this hook and line sample is quite representative

because it does not show any selectivity. These data indi¬

cate that hook and line catches of large numbers of perch will

present a representative sample, provided small hooks are used.

Table VI

Silver Lake, 1931 collections. Net catches compared with hook and line catches.

Rate of Growth

Nebish Lake

Collecting was carried on in this lake for three years, 1930,

1931, and 1932. A total of 632 perch was taken during this

period. Only 55 were collected in 1930, 232 in 1931, and 245

during the 1932 season.

The sexes were not determined in 1930 and 1931, but were

determined in 1932. The females numbered 157, and the re¬

maining 188 were males. Data regarding the growth of males

and females are given in Table VII, which shows the average

actual length, weight, K, and calculated lengths. In most cases,

the females show a slightly higher (2-4 mm.) rate of growth

than the males. The males of age-group V, however, are 11 mm.

Schneberger— Growth of Perch

115

longer than the females. As there are but two females and four

males in this group, the difference loses its significance. The

difference between the sexes of the other ages is also considerd

not significant, especially when compared with the average of

the combined data.

Table VII

Nebish Lake , 1932 . Averages of weight, K, actual and calculated lengths for

each age-group

Assuming that the same relation exists between the sexes of

other years, the growth of the other years may be compared.

Table VIII gives the data regarding each of the other years. The

youngest fish belong to age-group I, and the oldest to age-group

VI (7th year of life). Since age-group VI contains but one

specimen, which is smaller than the average of age-group V of

the same year, it is given no further consideration. The curves

in Figure 2 are based on the average actual length of each age-

group, with the exception of age-group 0. The value of this

group is based on the average calculated lengths. It will be

noted that the curve for the 1930 collection varies considerably

from the other years. Since there are but 55 specimens repre¬

sented in this curve, a great deal of significance cannot be given

to this difference.

116 Wisconsin Academy of Sciences , Arts, and Letters.

J _ I III

2 3 4 5 6

YEARS OF LIFE

Fig. 2. Growth of perch from Nebish Lake.

When the grand average (Table IX, Fig. 2) is taken, it is

seen to fit the 1932 data rather closely. This is to be expected,

inasmuch as the 1932 collection comprises over 50% of the

total collection. There is a sharp rise in increment the first two

years : 56 mm. the first year, and 68 mm. the second year. During

the third and fourth years there is a sharp decline in the incre¬

ment curve. This decline may be correlated with the attain¬

ment of sexual maturity. After the fourth year there follows an

increase in increment.

Jobes (1932) gives a preliminary report on the growth of

perch from Lake Erie. He finds that the growth is very rapid

in the first two years, but that there is a decline in growth rate

during the third summer. In regard to this decline Jobes says,

“The decided decrease in the growth-rate during the third sum*

mer may possibly be correlated with the attainment of sexual

maturity.” Figure 3 shows a comparison of the growth of perch

from, Nebish Lake with those from, Lake Erie. The curves of

this diagram show a very close agreement in the type of growth.

The Lake Erie perch, however, are consistently larger than the

Nebish specimens. The average increments show the same de-

Schneherger— Growth of Perch

117

cline during the third summer in Lake Erie and also in the Nebish

Lake perch. If this decline is due to the attainment of sexual

Table VIII

Nebish Lake , 1930 and 1931. Averages of weight, K, actual and calculated lengths

for each age-group.

Number Average Average calculated lengths

maturity, it has a greater effect on the Nebish perch during the

fourth summer than it does on the Lake Erie specimens. The

recovery of the Nebish perch occurs during the fifth summer,

while the Lake Erie specimens do not show an increase until a

year later.

Table IX

Nebish Lake. Summary of average lengths for each year of life.

Years of Life

* Calculated lengths, the rest are actual lengths.

1 Number of specimens in parentheses.

118 Wisconsin Academy of Sciences , Arts, and Letters.

Fig. 3. Average growth of perch from Lake Erie (Jobes) and Nebish Lake.

Weber Lake

Collecting was carried on during the summers of 1928, 1980,

1931, and 1932. Table Xa shows the number of fish in each

age-group, average length, K, and the average calculated lengths

for each of the four collections. Table Xb gives a summary of

the average length for each age group.

Two collections were made in 1932, one in the early season

during the first part of July, and one in the late season during

the latter part of August- The purpose of the two collections

was to determine whether or not any significant growth occurred

within the limits of the collecting season- Data regarding the

early and late seasons are given in Table XI. This table shows

that only age-group II contains sufficient numbers for statistical

comparisons. When these data are treated statistically, the

following means are found :

Early Season Late Season Edm

Females . 130.0 ± 0.299 134.16 ± 0.147 4.16 ± 0.347

Male . 126.1 ± 0.270 128.17 ± 0.148 2.07 ± 0.308

Edm . 3.9 ± 0.403 5.99 ± 0.209

S chneb erg er— Growth of Perch

119

The increment of the females from July 6 to August 17 was

4 mm., while the males increased 2 mm. during the same period

(42 days). The probable error of the difference of the means

Table Xa

Weber Lake , 1928, 1930 , 1931 and 1932. Averages of weight , K, actual and

calculated lengths for each age-group.

Number Average

Total 407

between the early and late season females is 4.16 ± 0.847, and

the same value for the early and late season males is 2.07 ±

0.308. The difference, though slight, is significant. When the

sizes of the sexes are compared, it is seen that the difference be-

tween the early season females and males is 8.9 ±0.403, and be¬

tween the late season females and males is 5.99 ± 0.209. In each

of these cases the difference of the means is small but significant.

These data indicate that there is a significant difference be¬

tween the growth of males and females, as well as between the

early and late season collections.

120 Wisconsin Academy of Sciences , Arts, and Letters.

Table Xb gives the grand average lengths of each age-group

based on the actual lengths of these specimens with the excep¬

tion of age-groups 0 and I, which are the average of all calcu¬

lated lengths for these two age-groups. The growth of the fish

Table Xb

*VII; 1928, 183(4), 1932, 223(9). VIII; 1932, 224(3). "Calculated lengths.

**Calculated, but not included in grand average.

Note: Gr. Av. VII, 210(13) ; VIII, 224(3). Av. Inc. VII, 12; VIII, 14.

Table XI

Weber Lake , 1932. Data on early and late season catches.

Number Average

Age- of Average Average actual Average calculated lengths per age-group

Note: For combined data

lection in table 13a.

on early and late season males

IV V VI VII VIII

157

162 187

167 195 212

143 165 184 209

143 171 186

127

150 166

135 156 171

130 144 160 181

and females, see 1932 col-

Schneb erger — Growth of Perch

121

taken in 1928 is the slowest of all. The growth of the other

years agrees quite closely to the grand average. The average

increment shows that the greatest growth was made during the

first year of life.

Silver Lake

A total of 1405 perch from Silver Lake were studied. Table

XII shows the data for each year. Of the total number of fish,

187 were collected in 1928, 412 in 1980, 463 in 1931, and 343

in 1932. As with the other lakes, the sexes were not determined

until the 1932 season. Table XIII gives the data regarding the

Table XII

Silver Lake, 1928, 1930, 1931, 1932. Averages of weight, K, actual and calculated

lengths for each age- group.

Number Average

122 Wisconsin Academy of Sciences, Arts, and Letters.

Table XIII

Silver Lake, 1932. Averages of weight, K, actual and calculated lengths for each

age-group

Number

Age- of Av’ge Av’ge Avg. act’l Average calculated lengths per age-group

group fish weight K length I II III IV V VI VII

Females

Total 136

See table XII, 1932 collections, for combined data.

growth of the males and females. Of the 343 perch taken in

1932, 207 were females and 136 were males. There seems to

be a decided difference between the growth of the two sexes, the

females growing at a much higher rate than the males. The av¬

erage of the combined data of the males and females is also

shown in the table. The females seem, to have a longer span of

life than the males-

The 1932 data are so conflicting that it is not possible to

make comparisons with other years, and they are, therefore,

omitted from the grand average shown in Table XIV. Such

differences may be due to the mixing of two local races. The

1932 collection was made about 200 meters from the station

where the previous collections were taken. The possibilities of

two local races in this lake are small because of its small size

(87.2 hectares) and also because a thorough sounding and map¬

ping of the lake revealed no barriers. If there are local races

present here, there must be a sharp localization of the two

groups. However, further data are necessary before this point

can be settled.

The growth of the three years 1928, 1930, and 1931 (Table

XIV) is in close agreement with the other years. In age-group

VI the average of those taken in 1928 is low. This is undoubtedly

Schneberger — Growth of Perch

123

due to the inadequacy of the data, inasmuch as there are only two

specimens. These are not included in the grand average. The

average increment, obtained from the grand average, shows a

sharp decline between the third and fourth years of life, which

may perhaps be interpreted as due to sexual maturity occurring

at this time.

Comparison of Growth Rates

Table XV and Figure 4 show the rate of growth of perch

from each of the three lakes, Nebish, Weber and Silver. The

lengths given are the grand averages of the actual lengths of

each age-group, with the exception of age-group 0 from Nebish

Lake, and age-groups 0 and I from Weber and Silver lakes.

These values are based on the average of all calculated lengths.

The data from Nebish are compiled from the material collected

in 1930, 1931, and 1932; Weber, 1928, 1930, 1931, and 1932;

and Silver, 1928, 1930, and 1931- Figure 4 shows that the

Nebish Lake perch have the most rapid growth; they attain an

average length of 245 mm. during six years of life. The aver¬

age of age-group VIII (9th year of life) of the Weber Lake

perch was 224 mm. The growth of the Weber Lake perch was

not sufficiently rapid for those specimens to attain a size equal

to the largest from Nebish Lake in three years additional time.

The early growth of the Silver Lake perch proceeds at a much

250

200

z

1 150

{5 100

2

LJ

"J 50

0

Fig. 4. Average rate of growth of perch from Nebish, Weber, and Silver lakes.

_ _J _ T I - - -

23456789

YEARS OF LIFE

124 Wisconsin Academy of Sciences, Arts, and Letters.

slower rate than that of the other two lakes, until the fourth

summer. Following the fourth summer there is an increase in

growth rate, so that at the end of the 7th year of life the Silver

Lake perch have surpassed those from Weber of the same age

by three millimeters. The figure shows that Nebish ranks first

in growth rate, Weber second, and Silver third.

Church (1927), studying the effect of crowding on the rate of

growth of the tropical fish, Platypoecilus maculatus rubra ,

found that the rate of growth was always greatest when the

fewest fish were present. These experiments were later repeated

by Shaw (1929) who obtained similar results.

Table XIV

Silver Lake. Summary of average lengths for each year of life.

Table XV

Comparison of the growth of perch from Wisconsin lakes with the growth of those

from other localities.

Lake O I II III IV V VI VII VIII

Nebish . 56 124 157 173 209 245

Weber . 53 95 130 158 174 191 198 210 224

Silver . 45 77 109 120 145 173 201

Erie (Harkness).. 44 144 168 187 217 234 244

Wawasee (Hile).. 39 86 129 167 198 220 230

Erie (Jobes . 69 150 176 197 316 250

The existence of two sizes of perch in the same body of water

or in closely connected or neighboring bodies of water is a phen¬

omenon that has been observed. In two of the Madison (Wis¬

consin) lakes, Lake Mendota and Lake Monona, differences in

size of perch are very noticeable. Lake Mendota, which is much

larger than Monona, and drains into the latter, contains a much

larger population of perch, but they are small. In the smaller

and shallower body of water, Lake Monona, the perch are almost

Schneberger — Growth of Perch

125

twice as large, but the population is much smaller. Wagner

(1910) assigns this difference to the fact that the conditions for

hatching and nourishing the young are better in Mendota than

in Monona. Naturally the abundance of young as well as of

older perch in Mendota brings on a struggle for food. This

competition for food does not have its effect in the elimination

of the weaker fish, but manifests itself in a decreased growth

rate. In Monona, such a high proportion of the perch does not

survive, and the competition for food is not so keen. Wagner also

mentions that the pike-perch ( Stizostedion ) might be a possible

factor in keeping down the population of perch in this lake.

Birge (1922) states that in 1883 or 1884 there was an epi¬

demic among the perch of Lake Mendota which resulted in an

enormous mortality. Those that died were only the larger speci¬

mens, and they were decidedly larger than those now found. The

removal of these larger specimens which had preyed upon the

smaller sizes, made conditions more favorable for the survival

of small perch. Insufficient food for this increased number had

its effect in that the small fish could not reach the larger size.

Dymond (1926) shows the same correlation between size of

fish and population in Lake Nipigon. In the sheltered southern

bays of this lake the perch occur in great numbers. With one

haul of about forty yards with a fifty foot seine, 1,760 perch were

taken. These were all small, the largest measuring less than

six and one-half inches. In the northern bays, which presum¬

ably have a smaller population, specimens measuring ten and

one-half inches are taken regularly.

Although a definite knowledge of the total population in

Nebish, Weber and Silver lakes is not available, it is possible

to obtain an idea of the relative abundance of perch present in

these lakes through the rate of capture in gill nets. This neces¬

sitates uniformity in gear and methods. During the 1932 season

all perch were taken by gill nets. The same nets were used in

each lake and set specifically for perch. The following table is

constructed from the 1932 records, and these are very similar

to the previous years.

From the above table it is seen that the perch are most

abundant in Silver Lake, and least abundant in Nebish Lake.

The relative abundance of perch in Weber Lake lies approxi¬

mately midway between Nebish and Silver lakes. The ratio

of fish caught per hour between Nebish and Silver is 1:11,

while the ratio between Nebish and Weber is 1:6; that is, dur¬

ing the period of time necessary to catch one perch in Nebish

the same nets would catch 11 and 6 fish from Silver and Weber

lakes respectively. When relative abundance of perch is cor¬

related with their rate of growth (Fig. 4), it is seen that an

indirect correlation exists. Nebish Lake perch are character¬

ized by the most rapid growth and least abundance. Silver

Lake perch have the slowest growth and are the most abundant.

Weber forms an intermediate group, both in growth and in

abundance. These data give evidence that the density of the

population is a contributing factor affecting the rate of growth

of fish.

It is interesting at this point to compare the growth of the

perch from the lakes of Northeastern Wisconsin with those from

different localities. Harkness (1922) and Jobes (1932) report

the growth of perch from Lake Erie, while Hile (1930) describes

the growth of perch from Lake Wawasee, an inland lake in

Indiana. The data given by these workers are included in

Table XV, and they are shown graphically in Figure 5. The

data for the growth of perch obtained by Harkness and Jobes in

Lake Erie are not in agreement, but their collections were

made in different localities and at a different time.

Attention has already been called to the striking similarity

between the growth of Lake Erie perch (Jobes) and those of

Nebish. The curves for these lakes are very similar except that

the Erie perch are consistently larger than the Nebish perch.

The greatest variation occurs between age-groups I, II and III ;

after this the Nebish perch have a higher rate of growth, and

at age-group V (6th year of life) the Nebish specimens are

only 5 mm. shorter than those from Lake Erie. The growth

Schneberger — Growth of Perch

127

of the perch from Lake Wawasee agrees closely with the data

of Harkness for Lake Erie. The first four years of Weber

(age-groups 0, I, II, and III) agree to a certain extent with

Wawasee and Erie (Harkness) ; after age-group III, the Weber

perch have a slower growth than that of the other two lakes.

With the exception of age-group 0 and VI, the Silver Lake speci¬

mens are consistently smaller than those from the other lakes.

It will be noticed in Figure 5 that the perch with the high¬

est rate of growth, those from Erie (Jobes) and Nebish, are not

as old as those with a lower rate of growth. Wawasee and Erie

(Harkness) perch have a lower rate of growth and are older.

In Weber Lake specimens as old as age-group VIII (9th year

of life) are represented and show slow growth. These data in¬

dicate that where the growth rate proceeds at a low rate the

fish live longer, and where the growth rate proceeds at a higher

rate, the span of life is shortened. In both Nebish and Silver

lakes a year older age-group than is shown in the diagram was

obtained, but since these groups contained only a few specimens

whose average lengths are shorter than the preceding groups,

they are not included in the figure. Further evidence is found

in the position of the first annulus of the scale. In the younger

fish (age-groups II and III) cases are found where the annulus

is far from the focus, indicating an extremely rapid growth.

Fig. 5. Growth of perch from Wisconsin compared with those from other

localities.

128 Wisconsin Academy of Sciences , Arts , and Letters.

However, this type is rarely found in the older fish, suggesting

that the rapid growers are short-lived.

This condition is not limited to perch, as Titcomb (1928) and

McCay (1981) have shown by their experiments on trout that

those that failed to grow lived longer than those that grew on

a similar diet. There is also evidence of this occurring in other

animals, particularly mammals. McCay (1933), briefly dis¬

cussing longevity and optimum growth, points out that those

diets that stimulate maximum growth are not conducive to

longevity. This author states, “It is possible that longevity and

rapid growth are incompatible and that the best chance for

an abnormally long life span belongs to the animal that has

grown slowly and attained a late maturity.”

Summary

1. The growth of perch from Nebish, Weber and Silver

lakes in Vilas County, Wisconsin, has been studied.

2. The weight of perch increases at a slightly higher rate

than the cube of the length. For the perch from Weber and

Silver lakes the value of the exponent is 3.10 and 3-19 respect¬

ively.

3. The weight-length relationship in the yellow perch is

not affected by the undigested food in the alimentary tract, nor

by the type of food present.

4. The sex ratio varies with the different lakes. In Nebish

the ratio of females to males is 1:1.26, while the ratio of males

to females is 1:1.31 and 1:1.40 in Weber and Silver lakes re¬

spectively.

5. Gill nets were selective in that they took only the larger

specimens of one age-group and the smaller of the next higher

age-group. Adequate sampling was obtained through the use

of a series of sizes of meshes, in which the difference between

sizes was small.

6. Hook and line samples of perch are adequate if the sam¬

ples are large and small hooks are used.

7. The males and females from Nebish Lake grow at ap¬

proximately the same rate.

Schneberger — Growth of Perch

129

8. The females in Weber Lake grow faster than the males.

Age-group II, 1932, shows significant statistical differences be¬

tween the growth of the males and females. The growth of age-

group II during a period of 42 days, from July 6 to August 17,

1932, is significant.

9. According to the data collected in 1932, the females from

Silver Lake grow more rapidly than the males. There is also

evidence that there are local races present in Silver Lake.

10. The highest rate of growth occurs in Nebish Lake perch.

Silver Lake perch exhibit the slowest growth, while those from

Weber Lake form a group intermediate between these two lakes.

11. The rate of growth is in inverse proportion to the rela¬

tive abundance. The fish caught per hour by gill nets averaged

1.57 in Nebish Lake, 9.78 in Weber Lake, and 16.5 in Silver

Lake.

12. Nebish Lake perch have a growth curve that is very sim¬

ilar to that reported by Jobes for the Lake Erie perch, except

that the Erie perch are consistently a trifle larger.

13. Weber Lake perch grow at about the same rate as the

perch from Wawasee (Hile) and Erie (Harkness) for the first

four years, after which the Weber perch grow at a slower rate-

14. The slower growing specimens have a longer span of

life. The rapid growing specimens seem to have a shorter span

of life.

130 Wisconsin Academy of Sciences, Arts, and Letters.

Literature

Birge, E. A. 1922. The plankton of the lakes. Trans. Amer. Fish. Soc. 52:128-129

Church, Frances. 1927. The effect of crowding on the growth of fishes. Master’s

Thesis, University of Chicago Library, 32 pp.

Clark, F. N. 1928. The weight-length relationship of the California sardine ( Sardina

caerulea ) at San Pedro. Fish Bull. No. 12, Calif. Fish and Game Comm., 58 pp.,

11 figs.

Crozier, W. J. and Selig Hecht. 1914. Correlations of weight, length and other

body measurements in the weakfish, Cynoscion regalis. Bull. U. S. Bur. Fish.

33:139-147. 4 figs.

Dymond, J. R. 1926. The fishes of Lake Nipigon. Univ. Toronto Stud. (biol. ser.),

Pub. Ontario Fish. Res. Lab. 7:97-107. 4 figs.

Harkness, W. J. K. 1922. The rate of growth of the yellow perch in Lake Erie.

Univ. Toronto Stud. (biol. ser.), Pub. Ontario Fish. Res. Lab. 6:87-97.

Hart, J. L. 1931. The growth of the whitefish, Coregonus clupeaformis (Mitchill).

Contrib. Canad. Biol, and Fish., n.s. 6(20) :429-444.

Hecht, Selig. 1913. The relation of weight to length in the smooth dog-fish,

Mustelus cams. Anat. Rec. VII: 39-41.

Hecht, Selig, 1916. Form and growth in fishes. Jour. Morph. 27(2) : 3 77-400.

Heinecke, F., et al. 1907. Untersuchungen der biologischen Anstalt auf Helgoland

zur Naturgeschichte der Nutzfische. Beteiligung Deutschlands a.d. Internat.

Meeresforschung. Bd. 4/5, 66-155.

Hile, R. 1931. Investigations of Indiana lakes. The rate of growth of the fishes of

Indiana. Pub. No. 107, Dept. Conserv., State of Indiana, pp. 9-55.

Jobes, F. W. 1933. Preliminary report on the age and growth of the yellow perch

( Perea flavescens Mitchill) from Lake Erie, as determined from a study of its

scales. Papers Mich. Acad. Sci. Arts and Letters. XVII: 643-652.

Keys, A. B. 1928. The weight-length relation in fishes. Proc. Nat. Acad. Sci. 14:922-

925.

McCay, C. M., et al. 1931. The nutritional requirements of trout and chemical com¬

position of the entire trout body. Trans. Am. Fish. Soc. 61:58-79.

McCay, C. M. 1933. Is longevity compatible with optimum growth? Science.

77:410-411.

Shaw, Gretchen. 1929. Further studies on the effect of crowding on the rate of

growth of fishes. Master’s Thesis, Univ. of Chicago Library.

Tester, Albert L. 1932. Rate of growth of the small-mouth black bass (Micro p-

terus dolomieu) in some Ontario waters. Univ. Toronto Stud. (biol. ser.), Pub.

Ontario Res. Lab. 47:207-221.

Titcomb, J. W.,* et al. 1928. The nutritional requirements and growth rates of brook

trout. Trans. Am. Fish. Soc., 58:205-231.

Van Oosten, J. 1923. The whitefishes ( Coregonus clupeaformis) . A study of the

scales of whitefishes of known ages. Zoologica. 2(17) :380-412.

Van Oosten, J. 1929. Life history of the lake herring ( Leucichthys artedi Le Sueur)

of Lake Huron as revealed by its scales, with a critique of the scale method.

Bull. U. S. Bur. Fish. 44:265-428.

Wagner, George. 1910. On the stickleback of Lake Superior. Science, n.s. 32:28-30.

FISH FOOD STUDIES OF A NUMBER OF NORTHEASTERN

WISCONSIN LAKES

Faye M. Couey

Notes from the Limnological Laboratory of the Wisconsin Geological and

Natural History Survey*. Notes and reports No. 58.

Introduction

This paper is based on the results of the examination of the

alimentary tracts of 5,767 fish obtained from twenty-four north¬

ern Wisconsin lakes during the summers of 1931 and 1932. It

was done as part of the program of the Wisconsin Geological

and Natural History Survey in the limnological observations

which have been carried on since 1925 at the Trout Lake Lab¬

oratory, Vilas County. The work has consisted of biological,

chemical, and physical observations on the waters of the lakes

of this region. The biological aspect includes a qualitative as

well as a quantitative study of the plankton and of bottom fauna.

For a good discription of the Highland Lake District of North¬

eastern Wisconsin and the Trout Lake Limnological Laboratory,

the reader is referred to Juday and Birge (1930).

In 1927, 1928, and 1930, with the cooperation of the U. S.

Bureau of Fisheries, considerable work was done on the age and

rate of growth of fish in these lakes. As this afforded a large

amount of material, the present work was begun at the sug¬

gestion of Professor Chancey Juday.

Methods

The fish were secured by means of gill nets, fyke nets, seines,

and hook and line. The work on age and growth rate was con¬

fined principally to six lakes, Trout, Muskellunge, Weber, Nebish,

Silver, and Clear, and most of the fish from these lakes were

caught with gill nets. In order to obtain a representative group

of fish, a string of these nets was used as a unit; a string con¬

sisted of seven one hundred-fifty foot nets ranging in size from

five-eights inch to one and one half inch bar mesh. This yielded

*This investigation was made in co-operation with the U. S. Bureau of Fisheries and the

results are published with the permission of the Commissioner of Fisheries.

131

132 Wisconsin Academy of Sciences, Arts, and Letters.

a larger number of fish and a larger number of species than

could be caught any other way. The fishing in the remaining

lakes was carried on mostly with hook and line and the number

of fish from some of them was small- The few small forms

that were studied were caught along the shore with a seine.

The fish were brought to the laboratory where they were

weighed, measured (standard length), scale samples taken, and

these data placed on the outside of the envelope containing the

scales. This work on the age and growth rate was done by the

fish-catching crew. Each fish was given a serial number which

was placed in its mouth so that reference could be made to its

data envelope.

Only the stomach contents were examined in the 1931 mate¬

rial, but in 1932 both stomach and intestinal contents were

taken into consideration. In the procedure with the 1931 mate¬

rial, the contents of the stomach were removed, placed in a

dish, and the amount of each article present estimated in terms

of percentage volume. This was done as follows: the amount

of the specific food ingestion was estimated as a certain per¬

centage of the amount of food present, and, to get a better idea

of the relative importance of the various articles, the degree

of fullness of the stomach was estimated, i.e., 100%, 50%,

10%, etc. This was taken into consideration in tabulating the

data by arriving at a weighted average between the first per¬

centage volume and the second.

With the 1932 material, the per cent fullness of the intestine

was estimated as well as that of the stomach and the two aver¬

aged, making allowance for their difference in size. The con¬

tents of the whole alimentary tract were placed together and

the amount of each article, in terms of the per cent of the total

amount of food, was calculated, again making allowance for the

percentage fullness of the alimentary tract. These figures were

analogous to those arrived at with the 1931 material.

Of the total number of fish studied, 3,015 were caught in

1931 and of these, 2,082 contained food. In 1932, 2,752 fish were

studied, and 1,944 of them contained food. Over one-third of

those studied during the two years were empty. This seems an

unusually large number to be empty and it was thought that

possibly complete digestion had taken place in those fish which

Couey — Fish Food Studies

133

had remained alive in the nets for some time. Another reason

for so many empty fishes might be regurgitation ; some of those

brought up from deep water had stomaches everted, due pos¬

sibly to pressure change. Some fish apparently regurgitated

their food while trying to escape from the nets.

For those lakes which had sufficient numbers of fish to have

some significance, graphs were made. The different foods were

condensed to seven chief articles of diet (fish, insects, amphi-

pods, crayfish, entomostracans, molluscs and plants) which are

represented by different lines. The original foods from which

these were condensed were: fish, Chironomus larvae and pupae,

Chaoborus larvae and pupae, Tanypus larvae and pupae, Pal-

pomyia larvae, Hexagenia nymphs, Coenis nymphs, caddis fly

larvae, Odonata nymphs, Sialis larvae, ants, corixids, miscellan¬

eous insects, hydrachnids, Mysis, amphipods, crayfish, ostracods,

copepods, cladocerans, snails, clams, leeches, algae, plants, Gor¬

dius, sponges, rotifers, and miscellaneous materials (debris etc.).

This list was also condensed somewhat for the tables.

In order to correlate type of food with rate of growth or

change of growth, the fish were graphically arranged in groups

with a length frequency of 10 millimeters . These, plotted against

the per cent volume of each article of diet, yield a graph from

which may be read the different foods of the different sizes of

fish. Due to a lack of the smaller fish, a knowledge of the food

of the young is wanting.

List of fishes studied

Perea flavescens (Mitchill) — yellow perch.

1931 — 1,078 examined and 710 contained food.

1932 — 1,210 examined and 788 contained food.

Ambloplites rupestris (Rafinesque) — rock bass.

1931 — 727 examined and 446 contained fo'od.

1932 — 718 examined and 546 contained food.

Helioperca macrochira (Cuvier and Valenciennes) — bluegill.

1931 — - 259 examined and 233 contained food.

1932 — 94 examined and 82 contaned food.

Micropterus dolomieu (Lacepede) — small-mouthed black bass.

1931 — 286 examined and 212 contaned food.

1932— 141 examined and 81 contained food.

Huro floridana (LeSueur) — large-mouthed black bass.

1931 — 27 examined and 20 contained food.

1932 — 8 examined and 4 contained food.

Stizostedion vitreum (Mitchill) — wall-eyed pike.

1931 — 46 examined and 23 contained food.

1932 — 64 examined and 44 contained food.

184 Wisconsin Academy of Sciences, Arts, and Letters .

Cristivomer namaycush (Walbaum) — lake trout.

1931 — ■ 15 examined and 7 contained food.

1932 — 7 examined and 5 contained food.

Esox masquinongy (Mitchill) — muskellunge.

1931 — 4 examined and 2 contained food.

1932 — 4 examined and 2 contained food.

Leucichthys artedi (LeSueur) — cisco.

1931 — ■ 359 examined and 266 contained food.

1932 — • 342 examined and 281 contained food.

Coregonis clupeaformis (Mitchill) — whitefish.

1931— 21 examined and 17 contained food.

1932 — - 31 examined and 14 contained food.

Catostomus commersonii (Lacepede) — common sucker.

1931 — 124 examined and 76 contained food.

1932 — 133 examined and 97 contained food.

EXPOSITION OF DATA

The fish of each lake and their foods are taken up separately

below, but only those percentages that are not obvious from the

tables will be mentioned. The various species of Diptera, Eph-

emerida, Entomostraca, Mollusca, and plants which were found,

are also mentioned, as tables could not be made comprehensive

enough to include them.

Trout Lake

This is one of the largest and is the deepest of the lakes in

the region studied. It is a drainage lake (has an outlet) .

Perch. The most common fish in the lakes studied is this

species, and more yellow perch were obtained from Trout Lake

than any other species.

Tables I and II show that the principal foods of the Trout

Lake perch are fish and insects, both of equal importance. In

the 1931 material, the total food was made up of 36% fish and

33.5% insects. In the 1932 perch, insects were 43% and fish

39% of the total. Those fish found in the perch diet which

were identifiable consisted of small perch, minnows and Johnny

darters. The Diptera larvae were Chironomus, Chaoborus (Cor-

ethra), Tanypus, and Palpomyia, but the 1932 perch contained

just Chironomus. The Ephemerida nymphs were all Hexagenia.

The miscellaneous insects, i.e., forms not regularly found, con¬

sisted of beetles, beetle larvae, mayflies (adult), leaf hoppers,

wasps, katydids, grasshoppers, and unidentifiable remains. In

1931 crayfish were rather important (16%) but were rare in

Couey — Fish Food Studies

135

the 1932 specimens. The Entomostraca consisted of Copepoda

(Cyclops, Diaptomus, and Canthocamptus) and Cladocera

(Daphnia, Holopedium, and Camptocercus). The Mollusca were

all snails (Planorbis and Physa) . Plants were mostly Potamoge-

tons. The “miscellaneous” column includes leeches, sponges,

rotifers, Gordius, unidentifiable food, and bottom debris.

The graphs of the food of the perch were not included because

of the varying nature of the data concerned. The fish ranged in

size from 60 to 220 mm. and there seemed to be little change of

diet, from the smallest to the largest. However, the smallest

fish which were caught showed a slightly higher insect content

in their food ; the larger perch showed a sudden drop in insect

food and an increase of fish as food. This seems to indicate that

the perch from this lake do not have a gradual change of diet

with growth, but that this change is sudden. Perhaps this phen¬

omenon is due to the fact that no fish sufficiently small was

caught to indicate the exact time of this sudden change in diet.

Rock Bass . The principal foods of the 1931 rock bass were

insects, crayfish and fish, in order of importance. In the 1932

rock bass the principal foods were insects and crayfish. The

fish diet consisted of minnows and darters. The Ephemerida

were all Hexagenia nymphs. A great difference in this food for

the two years was noted (30% of the total food was Hexagenia

in 1931; 2% in 1932). This difference may be due to the fact

that the 1931 fish were caught in nets in fairly deep water. Those

in 1932 were caught with hook and line in 4 to 8 feet of water in

the upper part of the lake near an old submerged pier and a

weed bed. The miscellaneous insects of the 1931 rock bass were

mostly grasshoppers with a few beetle remains. Those of

1932 were Gyrinid beetles, midges, dragon flies, and grass¬

hoppers. The larger percentage of insects here was probably

due to the shallow water and the overhanging branches along

the shore. The plants found were Potamogetons, with a few

algae. The Entomostraca were Daphnia and Leptodora.

The rock bass ranged in length from 60 to 200 mm. and

these, as well as the perch, presented an inconsistent and varying

set of curves when the lengths were plotted against the foods.

The smaller rock bass ate principally insects until they reached

about 130 mm., at which time there was a decrease in insects

and an increase in crayfish and fish in their diets.

136 Wisconsin Academy of Sciences , Arts , and Letters.

Cisco. The food of the ciscoes consisted almost entirely of

plankton. In Trout Lake, they ate a few Chironomus and Chao-

borus larvae, but by far the most important element of the food

was Entomostraca. In 1931 Copepoda constituted 49% and

Cladocera 33% of the food. In 1932 the Copepoda made up

42% and the Cladocera 47.5%. Some fish eggs were found in

a few of the 1931 specimens. There was not enough variation

in the size of these ciscoes to make any correlation between

length and diet.

Whitefish. Whitefish were taken only in Trout Lake. A

large percentage of the food came from the bottom and con¬

sisted of dipterous larvae and molluscs; a few entomostracans

were noted also. The larval Diptera were Chironomus, Chao-

borus and Tanypus. Mysis was a favorite food, but it was not

found in any other species of fish except in the ciscoes from

Black Oak Lake. The molluscs were represented by small speci¬

mens of Pisidium. The Entomostraca included Ostracoda, Can-

thocamptus, Leptodora, Acroperus and Ophryoxis. The large

amount of miscellaneous material was mostly bottom debris,

pine seeds, fish eggs, and rotifers (Conochilus).

Small-Mouthed Black Bass. Representatives of this species

were caught with a minnow seine and hook and line. In 1931

ten small bass were caught along the shore with a seine. They

were 30 to 40 mm. in length. The main food was Diptera larvae

(Chironomus — 11%, Chironomus pupae — 5%, Chaoborus pupae

— 8.5%, Tanypus — 3%, and Palpomyia — 10% ) . The Ephemer-

ida were all Hexagenia nymphs. The miscellaneous insects were

midges, leaf hoppers, and damsel flies. The Entomostraca were

all Sida. The miscellaneous column consisted of land spiders.

Insects seem to be the main food of small bass. The one small¬

mouthed bass caught in 1932 contained a large crayfish.

Wall-Eyed Pike. The wall-eyed pike fed almost entirely on

other fish. Of the 52 examined from this lake during both sum¬

mers, only one had eaten anything besides fish and that one had a

stomach full of Hexagenia nymphs. The fish eaten were perch,

cisco, and minnows.

Lake Trout. There are only five lakes in Wisconsin where

this fish is caught, and of these lakes, three are situated in this

Couey—Fish Food Studies

137

region. The only specimens caught during this investigation

were from Trout Lake. Some of the trout were caught in gill

nets and some with hook and line.

Lake trout inhabit deep water and live on fish. The food

of those included in this study consisted mostly of ciscoes, but

suckers were found in two of them. One trout measuring 810

mm. in length had eaten a sucker 830 mm. long, which weighed

520 grams.

Sucker . Because the sucker is a bottom feeder, its food varies

somewhat, consisting of molluscs, dipterous larvae, entomostra-

cans, bottom ooze and debris. The principal food was molluscs

(Amnicola, Planorbis, Bythinia, and Pisidium). Chironomus

larvae made up 18% of the food in 1931 and 17.5% in 1932.

There was a considerable variety of Entomostraca but they were

rather few In number. There were some Ostracoda, but most

of the Entomostraca were Copepoda (Canthocamptus and Cy¬

clops) and Cladocera (Bosmina, Camptocercus, Daphnia, and

Ophryoxis) .

The size of the suckers ranged from 120 to 380 mm., no small

ones being caught. The curves for 1931 show a gradual de¬

crease in insect food as the length of the fish increases, and a

correspondingly increased mollusc consumption. In 1932 no

sucker was caught measuring less than 280 mm. in length.

Mollusc food was predominant in those caught. In suckers be¬

tween 320 and 360 mm., insects replaced the mollusc diet, which

again prevailed in the larger fish.

Nebish Lake

This is a small lake of medium depth (15.8 m.) with rather

clear water. The marshy weed beds found in, so many of the

other lakes are lacking here. The food of the fish from this lake

differs from that of other lakes in the great predominance of in¬

sects.

Perch. Insects made up 81% of the food of the 1931 perch

and 53% of those caught in 1932. There was a further de¬

parture from the usual ephemerid findings in that 28.5% of

the 1931 food consisted of Coenis; against 1% Hexagenia in

1931 and entirely Coenis (14.5%) in 1982. This was unusual

because Coenis nymphs were found in hardly any of the other

138 Wisconsin Academy of Sciences, Arts, and Letters.

lakes as food. The Entomostraca were almost entirely Daphnia

with a few Leptodora. In 1931 the Moll u sea were Pisidium

with a few Planorbis and in 1932, Pisidium with a few Planorbis

and Amnicola.

31 —

90 110 130 150 170 190 210

Fig. 1. This diagram shows the percentage composition of the food of different

sizes of perch taken in Nebish Lake in 1931. Compare with fig. 2.

Figure 1 represents the type and amount of food eaten

by the different sized perch. Here insects predominate; the

other foods have little significance. In Figure 2, insects are

also the most important food, but there is an increase in En¬

tomostraca in specimens between the lengths of 110 and 150

mm. This increase of Entomostraca during a certain period

of growth seems to be characteristic of the perch in all the

lakes studied. It occurs as a rule in those perch from 120 to

150 mm. in length.

Rock Bass. The food of this fish is quite similar to that of

the food of perch, the main food being insects. Of the Diptera

larvae, Chironomus were most important (1931 — 19.5% of the

total food; 1932—42.5%). Of the Ephemerida, Coenis predom¬

inated (1931—16.5%; 1932 — 24.5%). The Entomostraca con¬

sisted entirely of Cladocera; in order of their importance, (1931)

Holopedium, Leptodora, Daphnia, Sida; (1932) Daphnia, Lep¬

todora, and Holopedium. The molluscs consisted mostly of Pis-

Couey—Fish Food Studies

139

90 110 130 130 170 190 210

Fig. 2. This diagram shows the percentage composition of the food of differ¬

ent sizes of perch taken in Nebish Lake in 1932. Compare with fig. 1.

idium with a few Planobis in 1931 ; in 1932 a similar condition

existed except that a few Amnicola and Physa were found.

The 1931 rock bass were all so much alike in size that they

presented no significant curves. Figure 3 shows the curves for

the 1932 rock bass. In these the insect food increases irregu¬

larly; fish appears in the stomachs of the 140 mm. group and

predominates in the 190 mm. rock bass.

In October of 1932, twenty-seven rock bass were taken from

Nebish Lake; Table II shows the marked change of diet that

took place with change of season. The main food was Ento-

mostraca, to be followed by insects and molluscs. The ento-

mostracans were all Daphnia pulex • They were winter forms,

their ephippia containing winter eggs. The molluscs were Plan-

orbis (11% of the total food) and Pisidium (1%). Insecta

included no Diptera larvae and, as shown in Table 1, consisted

of caddis fly larvae, dragon fly nymphs and Sialis larvae. The

miscellaneous insects were mostly the remains of small aquatic

beetles. Of note is the fact that no Coenis nymphs were found.

Figure 4 shows the Entomostraca gradually decreasing with

increase in the size of the rock bass, and the insect food coming

into prominence. In the largest fish, insects are, in their turn,

replaced by molluscs.

140 Wisconsin Academy of Sciences, Arts , and Letters.

J %J V /‘‘•s, t l£. ^

70 90 110 130 150 170 190 210

Fig. 3. This diagram shows the percentage composition of the food of differ¬

ent sizes of rock bass taken in Nebish Lake in July and August 1932. Compare with

fig. 4.

Small-Mouthed Black Bass . The principal food of the large¬

mouthed bass was insects (1931 — 61% ; 1932 — 51%), which was

followed in order by fish and Entomostraca. The Diptera larvae

were mostly Chironomus. The Ephemerida were largely Coenis

(1931 — 12.5%; 1932 — 5%). The miscellaneous insects were

all grasshoppers for both years. Cladocera were the only rep¬

resentatives of the Entomostraca, consisting chiefly of Lepto-

dora, with a few Daphnia and Sida. The Mollusca were rep¬

resented by Pisidium.

The graphs of these fish were not included, but it is worthy

of note that Cladocera were found in the 110 to 160 mm. fish.

Insects predominated in fish up to 220 mm., when fish became

the main food. However, as the 1932 material contained no

fish less than 150 mm. long, the curves were of little significance.

Large-Mouthed Black Bass. One large-mouth bass 118 mm.

long was taken from this lake in 1932. It was 50% full and had

eaten Hexagenia (60%) and Gyrinid beetles (40%).

Weber Lake

This is the clearest and the smallest of the lakes studied. It

is a seepage lake with soft water, and has an area of about 38

acres and a maximum depth of 13 m.

Co^ey — Fish Food Studies

141

0 — . — _jl_ _ I _ Z-~f V

80 120 120 140 160 ! 80

Tig. 4. This diagram shows the percentage composition of the food of different

sizes of rock bass taken in Nebish Lake in October 1932. Compare with fig. 3.

Perch . The principal food was insects (1931 — 84% and 1932

— 79.5%). Caddis fly larvae were the principal insect food for

both years (1931 — 65.5% ; 1932 — 45.5%). The Diptera larvae

for 1931 were Chironomus (35%), Chaoborus (11.5%), and

Tanypus (0.5%); for 1932, Chironomus (19%), Chaoborus

(8%), and Tanypus (3-5%). Noteworthy is the absence of

Ephemerida, especially Hexagenia which was found in most of

the lakes, '"he miscellaneous insects for 1931 were grasshoppers,

caterpillars, damsel flies, midges, parnid larvae, and adult

dragon flies. For 1932 they were grasshoppers, Hydroporus lar¬

vae, beetle remains, and unidentified beetle larvae. The En-

tomostraca were, for both summers, entirely Cladocera (1931:

Ophryoxis, Holopedium, and a few Camptocercus ; 1932: Holo-

pedium, Daphnia, Bosmina, Chydorus, Sida, Acantholeberis, and

Diaphanosoma) . A few molluscs were found ; one Planorbis and

some specimens of Pisidium were observed.

In 1931 insects predominated in all sizes of perch, the other

foods having little significance. In Figure 5 (1932), the young

fish subsisted chiefly on Cladocera, changing to insects at 120

to 130 mm. Insects continued to be the principal food through¬

out the rest of the size range.

142 Wisconsin Academy of Sciences, Arts, and Letters.

Small-Mouthed Black Bass. The 1931 material consisted of

five bass with an average length of 230 mm., and twenty-four

small ones between 50 and 80 mm. in length. Of the first five,

one 110 mm. specimen was 10% full, contained Chaoborus

100

80

60

40

20

Fig. 5. This diagram shows the percentage composition of the food of different

sizes of perch taken in Weber Lake in 1932. Compare with figs. 2 and 6.

pupae (60%), Holopedium (20%) and Camptocercus (20% ).

One, 234 mm. in length and 50% full, had eaten two minnows

(80%) and plant material (20%). The third fish, 232 mm.

long and 10% full, contained fish (90%) and Chironomus

(10%). One, 217 mm. in length and 100% full, had eaten one

small fish (10%), 10 grasshoppers (60%), and one frog

(30%). The fifth one was empty.

The smaller bass ate mostly insects but there was a great

variation in their diet. For instance, of four small ones caught

on the same day, one was empty, one was 100% full and con¬

tained Palpomyia larvae (100%); one, 30% full, had eaten

Chironomus larvae (20%), Palpomyia (20%), caddis fly larvae

(40%), Daphnia (10%), Sida (5%), and Bosmina (5%); one,

65% full, had eaten Chironomus larvae (5%), Palpomyia

(80%), Acantholeberis (5%) and Bosmina (5%). Twenty

small ones taken on another day had eaten mostly insects which

were, in order of importance: ants, an unidentified fly larva,

caterpillar, damsel fly, midge, and Chironomus larvae. One,

Coney — Fish Food Studies

143

25% full, was quite unusual in having eaten 100% Polyphemus.

Acantholeberis and Acroperus were found in small amounts in

the others.

Only two small-mouthed bass were obtained in 1932. One

of these, 15% full, contained a grasshopper; the other, 100%

full, had eaten a perch.

Muskellunge Lake

This is a rather shallow seepage lake with fairly soft water.

Data on fish caught in South Bay were kept separate from those

of North Bay during 1931. In 1932 four stations were estab¬

lished and these were designated North Bay, South Bay, West

Bay, and Station Four. This was done to see if there was any

difference in rate of growth of fish in different parts of the lake.

As there was not enough difference in food to warrant a sep¬

arate treatment of the various stations, the results for the en¬

tire lake are grouped together.

Perch. In 1931 fish were the main food of the perch with

insects (20%) ranking second and Entomostraca third. The

Diptera larvae consisted of Chironomus (5.5% of the the food

total) and Chaoborus (0.5%). The Ephemerida were all Hex-

agenia. The Entomostraca were Copepoda (0.5%) and Clado-

cera (15.5%). Molluscs were all snails.

In 1932 insects were first in importance in the perch diet

(36%), fish second and Entomostraca third. Among the in¬

sects, Chironomus made up 12.5% of the food, Chaoborus 6%,

Hexagenia 2% and Coenis 1%. The Entomostraca were Os-

tracoda 1%, a few Copepoda, and Cladocera 14.5%. The mol¬

luscs consisted of 4.5% snails and 0.5% bivalves. In 1931 the

perch ranged from 90 mm. to 240 mm. in length. The smallest

perch ate chiefly insects, but the percentage of insect material

showed a marked decline in the 100-130 mm. groups- Correlated

with a decrease in insects was a corresponding increase in En¬

tomostraca and fish. Perch that were 150 mm. or more in

length subsisted chiefly on small fish. One perch with a length

of 190 mm. had a stomach full of plant material.

The 1932 perch from Muskellunge Lake ranged from 80 mm.

to 230 mm. in length. Figure 6 shows that the young perch ate

Entomostraca chiefly and these organisms continued as an im-

144 Wisconsin Academy of Sciences, Arts, and Letters.

portant element of the food up to a length of 160 mm. Insects

appeared in the smaller perch and continued to be important

even in the larger sizes, where small fish became the chief item

of food.

The perch from North Bay in 1931 fed chiefly on fish and En-

tomostraca. In 1932 the perch from this bay ate insects (Chir-

Fig. 6. This diagram shows the percentage composition of the food of different

sizes of perch taken in Muskellunge Lake in 1932. Compare with figs. 2 and S.

onomus larvae and Odonata nymphs) and Entomostraca. The

perch from South Bay ate mostly fish in 1931, but in 1932 their

diet was more varied, consisting of insects and fish, with lesser

amounts of snails and amphipods. The perch of Station Four

in 1932 ate mostly Cladocera, with some insects and fish, while

the West Bay perch fed on Diptera larvae and fish.

Rock Bass. In 1931 insects were the principal food (40-5%)

of rock bass. Diptera larvae consisted of Chironomus (1%

of the food total) and Chaoborus (3%). Ephemerida were all

Hexagenia. The molluscs were all snails.

In 1932 insects were also the main food (58%). The Dip¬

tera larvae were Chironomus (2%) and Chaoborus (8.5%).

Ephemerida were Hexagenia (17%) and Coenis (2%). Mol¬

luscs were represented by snails only.

In 1931 the food of the smaller rock bass consisted chiefly

of insects ; there was a gradual decrease in insects with increase

Couey — Fish Food Studies

145

in the size and a corresponding increase in the amount of fish

and crayfish consumed in specimens above 120 mm, in length.

In 1932 (Fig. 7), the food relationship was similar to that of

1931 except that crayfish were not so important and insects

were more so.

Fig, 7 . This diagram shows the percentage composition of the food of different

sizes of rock bass taken in Muskellunge Lake in 1932. Compare with figs. 3 and 4.

The rock bass from North Bay in 1931 ate fish and insects

with some crayfish. In 1932 insects were the most important

food. The rock bass from South Bay ate mostly insect larvae

(Odonata nymphs) with some fish and plants. Their food was

about the same for the two years. At Station Four insect larvae

(Hexagenia and Odonata nymphs) and crayfish were the chief

foods. In West Bay the rock bass ate mostly fish and Hexagenia

nympmhs.

iBluegilL In 1931 insects were the principal food (49.5%).

The Diptera larvae were Chironomus (19.5%), Chaoborus

(1%), and Tanypus (0-5%). Ephemerida were all Hexagenia

and molluscs were all snails.

In 1932 plants were the main food (34% ) . (Fig. 8) . Seven

per cent of the food total were green algae and 27%, large

aquatic plants. Insects constituted 29.5% of the food. The Dip¬

tera larvae were Chironomus (8.5%), Chaoborus (2%) and

146 Wisconsin Academy of Sciences , Arts, and Letters.

Palpomyia (1.5%). The Ephemerida were Hexagenia (1%)

and a few Coenis. Entomostraca were Ostracoda (2.5% of all

food present) and a few Cladocera. Molluscs were snails with

a few clams.

In 1931 insect food predominated in the small bluegills, de¬

creased and was replaced by plant material in the medium sizes,

and again increased in the larger specimens. The pronounced

decrease in the amount of insect material in the food of bluegills

ranging from 90 to 140 mm. in length seems to be characteris¬

tic of this species. Figure 8 shows a similar food distribution

in specimens obtained in 1932-

100 - ENTOMOSTRACA

. AMPHIPODA

- MOLLUSCA

- INSECTS

80 - FISH -

w --PLANTS /

/

60- I -

✓v. y.i

40 — Vr —

20- \ \ -

\ '*/

01 i _ r \ 4

60 80 100 120 140 160

--ENTOMOSTRACA

-—AMPHIPODA

—MOLLUSCA

-INSECTS

— FISH

—PLANTS

A

A., y

tT

/

/ -]

/

\

V

..Ak-"

/

y

/

\

Fig. 8. This diagram shows the percentage composition of the food of different

sizes of blue-gills taken in Muskellunge Lake in 1932.

Bluegills were taken from North Bay in 1932 only and they

ate mostly insect larvae (Diptera, caddis fly, and Odonata).

Those from South Bay in 1931 ate chieflly plants, Diptera larvae,

and ants, with some molluscs. In 1932 the bluegills from this

bay fed on plants with lesser amounts but greater variation

of the other foods. The three bluegills from West Bay had

eaten plants and some insect larvae.

Small-Mouthed Black Bass. Fish were the chief food of

this species of bass in both years; in 1931 they constituted 75%

Couey — Fish Food Studies

147

of the diet. In the 1932 specimens, insects constituted 48.5%

of the food. Fragments of large aquatic plants were found in

some cases. Fish constituted an important element of the food

of small-mouths ranging from 110 to 140 mm. in length in 1931,

but no fish material was found in those ranging from 140 to

160 mm. in length; the percentage of fish material increased

in those ranging from 160 to 230 mm. in length. Plant material

formed an important item in those between 160 and 190 mm.

in length.

In 1932 two small-mouths with lengths of 100 and 110 mm.,

respectively, had eaten only Entomostraca. The other specimens

varied from 140 to 250 mm. in length; fish material was the

main article of diet in them, but insects were quite important in

the 140-190 mm. group.

The small-mouths from North and South bays in 1931 ate

chiefly fish. Plants occurred in the diet to some extent. In

1932 the specimens from North Bay ate chiefly terrestrial in¬

sects and fish. No specimens were obtain from South Bay in

1932. The West Bay specimens had eaten only fish and those

from Station Four ate mostly fish.

Large-Mouthed Black Bass. The unusual appearance of the

food of the large-mouthed bass in Table I is due to the combin¬

ing of both the small and the large fish of this species from this

lake. The large per cent of fish in the diet is due to the large

sized fish, and the large amount of Entomostraca can be ascribed

to the fact that the small ones fed mostly on Cladocera. The

“miscellaneous” insects labeled in the table were all found in

the small fish.

In 1932 all specimens caught were rather large and they ate

fish entirely. The plant food was probably accidentally ingested.

Large-mouthed bass were caught in South Bay only in 1931

and their food consisted of fish and Entomostraca. The latter

were found in the smallest specimens. In 1932 this species was

taken from North and West bays and they were found to have

fed mostly on fish.

Muskellunge . Four of these fish were caught in 1931 from

South Bay and their food consisted entirely of fish (Johnny

darters and chubs).

148 Wisconsin Academy of Sciences, Arts, and Letters.

Cisco. Entomostraca were the main food of ciscoes. The

Entomostraca in the 1931 speicmens consisted of Cyclops (50%),

Cladocera (24%), and a few Ostracoda. The Diptera larvae

were Chironomus (1%) and Chaoborus (3%).

The Entomostraca of the 1932 ciscoes were Cladocera

(59%), copepods (42%), and a few Ostracoda. The Diptera

were Chironomus larvae (1.5%) and Chaoborus (1.5%).

Not much can be said concerning the food of the different

sized ciscoes as there was little difference in the size of those

caught. Ciscoes were taken from North and West bays only and

there was little difference in type of food.

Sucker. In 1931 insects were the main food (26%). The

Diptera larvae were Chironomus (13.5% of all food present),

Chaoborus (1.5%), Tanypus (6.5%) and Palpomyia (2%).

The Ephemerida were all Hexagenia. The Entomostraca were

Ostracoda (1%), Copepoda (5%), and Cladocera (9%). Mol¬

luscs were snails (8%) and clams (4.5%). Plants were mostly

fragments of higher aquatic forms.

In 1932 insects were also the main food (28% ) . The Diptera

larvae consisted of Chironomus (17.5%), Chaoborus (1.5%),

Tanypus (0.5%), and Palpomyia (1%). The Ephemerida

were Hexagenia (1%) and Coenis (2%). Entomostraca were

Ostracoda (1.5%), Copepoda (3.5%), and Cladocera (15.5%).

Molluscs were snails (6%) and clams (2.5%).

The 1931 suckers were 110 to 310 mm. in length. Because

of lack of numbers, frequencies of 20 mm. were used in the

curves in place of 10 mm, but even so the curves did not show

much. Cladocera were most important in the small suckers,

molluscs increased in importance in those between 170 and 190

mm., and insects predominated from 170 mm. up.

The 1932 suckers were also divided into groups on a 20 mm.

basis, extending from 150 to 330 mm. Entomostraca were pre¬

dominant in the fish up to 210 mm. and then decreased in num¬

ber as insects increased- Insects continued to be the main food

of the larger suckers.

The suckers from North and South bays in 1931 ate foods

that were quite alike, consisting of dipterous larvae Entomos¬

traca and small molluscs. The South Bay suckers ate more En-

Couey—Fish Food Studies

149

tomostraca than those from North Bay. In 1932 suckers were

taken from North and West bays and Station Four and their

foods were quite similar to those of the previous summer.

Silver Lake

This is a small lake that has about the same type of water

as Trout Lake and it drains into Trout Lake in times of high

water.

Perch . In 1931 insects were the principal food (54.5%) of

Silver Lake perch. The Diptera larvae were Chironomus (4%),

Chaoborus (0.5%), Tanypus (1%), and Palpomyia (1%). Eph-

emerida were all Hexagenia. The miscellaneous insects were

grasshoppers, gyrinid beetles and tabanid larvae. The Ento-

mostraca consisted of Copepoda (4.5%), and Cladocera (5%).

Molluscs were all snails (Physa). Plants consisted of algae

(1%) (Aphanocapsa) and large aquatics (Potamogetons) .

Figure 9 shows the 1931 food curves for the various sized

Silver Lake perch, which ranged from 100 to 230 mm. The

young ate mostly Entomostraca, but insects appeared in the

110 mm. specimens and predominated in the larger sizes. The

characteristic change in food, previously described, occurs in the

140 to 160 mm. perch, at which time there is an increase in the

amount of fish material consumed.

100

80

60

40

20

0

90 110 130 150 170 190 210 230

Fig. 9. This diagram shows the percentage composition of the food of different

sizes of perch taken in Silver Lake in 1931. Compare with figs, 1, 2, 6 and 8.

150 Wisconsin Academy of Sciences, Arts, and Letters .

In 1932 a considerable change in diet was noted. The main

food was fish; this item made up 90% of all food and consisted

almost entirely of young perch. Entomostraca were found only

in the smallest specimens.

Rock Bass. In 1931 the principal food of rock bass was in¬

sects (51.5%). The specimens were 60 to 140 mm. in length.

Crayfish were eaten by the smallest rock bass and they increased

in importance until they became the main food in the 120 mm.

specimens. Only two rock bass containing food were obtained in

1932. One of them (15% full) contained only fish and the other

(25% full) had eaten only dragon fly nymphs.

Bluegill Bluegills were caught only in 1931; they were 70

to 130 mm. in length. The main food was insects (86%). Chir-

onomus larvae (10.5%) were most important and Chaoborus

(2%) next. Hexagenia and damsel fly nymphs were found in

some numbers, as well as a few Copepoda and Cladocera. Physa

represented the mollusc group.

Small-Mouthed Black Bass. In 1931 the small-mouthed black

bass from this lake ranged from, 70 to 270 mm. in length; the

chief constituent of their food consisted of insects, with fish

material second in importance. Both of these foods were eaten

by all sizes. Five specimens were secured in 1932 ranging from

133 to 200 mm. in length ; all of them had eaten small perch.

Cisco. Ciscoes ranging in length from 130 to 210 mm. were

taken only in 1931. Their chief food consisted of Copepoda

(59%) and Cladocera (30.5%) ; this was true of all sizes.

Sucker. Insects and molluscs constituted the chief food of

the suckers in 1931; the former made up 36.5% and the latter

33.5%. The Entomostraca were represented by 5% Cladocera

and 4.5% Copepoda. These suckers varied from 155 to 305 mm.

in length, but all of them ate about the same type of food.

Clear Lake

This is a seepage lake with clear, soft water. The fish studies

on this lake were concentrated on the ciscoes, so that the other

species are not well represented.

Perch. The perch obtained in 1931 were between 115 and

177 mm. in length. Insects constituted the main food item

Couey — Fish Food Studies

151

(86.5%), with Chironomus and Hexagenia as the chief forms.

Nostoc was found in some specimens. Only three specimens

containing food were obtained in 1932; their food consisted of

65 % insects and 35 % Leptodora.

Rock Bass. Insects were the chief element in the food of

three rock bass taken in 1931, namely 37.5%; the three speci¬

mens taken in 1932 contained 97% insects, chiefly Hexagenia

(92%).

Small-Mouthed Black Bass. One small-mouthed black bass

was taken in 1931. It was 125 mm. long and contained 100%

fish.

Wall-Eyed Pike. The 1931 pike measured 220 to 380 mm.

in length and fed chiefly on fish (darters and minnows). Insects

(19.5%) consisted of Diptera larvae (Chironomus) and Ephem-

erida (Hexagenia). Entomostraca were Cladocera (Leptodora).

Plants were green algae (Coleochaete) .

The 1932 pike measured 221 to 390 mm. and fed chiefly on

insects (59.5%). One specimen had eaten 84 large Hexagenia

nymphs. The fish eaten consisted of darters and minnows.

Cisco. Entomostraca were the chief food of the ciscoes. In

1931 Entomostraca made up 96.5% of the food and consisted of

Cyclops (0.5%) and Cladocera (96%) ; the latter were chiefly

Daphnia, with some Leptodora and a few Holopedium. Insects

were 3.5% of the total food and consisted of Chironomus 2%,

with a few Hexagenia and Tanypus.

In 1932 Entomostraca made up 99.5% of the food, consisting

of Cladocera (Daphnia 93% of total food, Leptodora 6.5%, and

a few Holopedium). One cisco (no. 963) had eaten 3,052 large

Daphnia and nothing else. The only insects found in these cis¬

coes were Chironomus larvae (0.5%).

Vieux Desert

This is the largest lake in this area although it is quite shal¬

low. It has deeply colored water due to vegetable extracts.

Perch. Insects constituted 72% of the food of the perch

obtained from this lake in 1931; 41% consisted of Chironomus.

While insects predominated in the food of the younger perch,

152 Wisconsin Academy of Sciences, Arts, and Letters.

they showed the characteristic decline in percentage in the 110

to 140 mm. groups; insects were replaced by fish in the larger

perch. Hyalella was eaten to a considerable extent by perch

ranging from 170 to 190 mm. in length.

In the 1932 specimens from this lake, Hyalella was the most

important item of food (51%) ; this crustacean did not form

such a large percentage of the food in any other lake although

it was present in them. Insects were next in importance

(32.5%) ; molluscs furnished 4.5% of the food and a few Daph-

nia were found in one specimen. The 1932 perch were caught

at two different times (55 in July and 15 in August) and the food

was very different for the two periods- The large percentage of

Hyalella was found in the July specimens, while most of the

snails were noted in the August catch. The youngest perch,

beginning at 70 mm., had eaten chieffy insects, but the per¬

centage of insects decreased sharply in the 100-140 mm. perch

and was replaced by Hyalella which remained predominant up

to 150 mm. Insects, however, continued to be important until

supplanted by fish in the larger perch (170-190 mm.)

Pallette

This is a small seepage lake with clear soft water.

Perch. Perch were taken from this lake only in 1931. The

principal food was fish, consisting of small perch and minnows.

Insects amounted to 33 % and snails ranked third in importance.

Five small specimens, averaging 30 mm. in length, were ob¬

tained; their food consisted entirely of Entomostraca (Diapto-

mus 82%, Cyclops 14% and Chydorus 4%).

Rock Bass. Only five rock bass containing food were taken ;

insects were the chief article of diet and consisted mainly of un¬

identifiable remains.

Small-Mouthed Black Bass. The principal food of the 1931

small-mouths consisted of small perch and minnows. The insect

material contained an appreciable amount of grasshoppers; one

small specimen had eaten Daphnia- Fish was the principal con¬

stituent in the food of the 1932 specimens. Chaoborus made up

2% of the food and Chironomus larvae 1%. Leptodora was

found in a 169 mm. specimen.

Couey — -Fish Food Studies

153

Cisco. Ciscoes were obtained only in 1931. The principal

food was Entomostraea (89%), consisting of 36% Cyclops and

53% Cladocera (Daphnia, Chydorus and Bosmina).

Sucker. A 351 mm. sucker was secured ; it had eaten chiefly

Entomostraea, consisting of 50% Copepoda and 20% Cladocera.

Chaoborus larvae made up 10%, other insect remains 10% and

plant material 10%.

Long Lake

This is a small, shallow lake with fairly clear water. Speci¬

mens for food studies were taken from this lake only in 1931.

Perch. Entomostraea were the principal food of these perch ;

they consisted entirely of Cladocera (Holopedium, Daphnia,

Bosmina and Sida). Insects (37.5%) were next in importance,

with 5% Chironomus larvae and 2% Chaoborus larvae; the rest

were unidentifiable fragments- These perch ranged in size

from 100 to 160 mm. The smaller ones ate chiefly entomostra-

cans and those between 130 and 150 mm. chiefly insects. Fish

appeared in the food of the 150 mm. group.

Large-Mouthed Black Bass. Three small specimens of large¬

mouthed black bass were examined. The principal food was

fish, but one of them contained remains of a moth.

Black Oak Lake

This lake has an area of 230 ha. and a maximum depth of

26 m. The water is fairly soft.

Ciscoes were obtained from this lake in 1932; they ranged

from 170 to 210 mm. in length. Entomostraea formed the

principal food, but Mysis and insects played an important role

also. Ostracoda made up 9% of the total food, while the cope-

pods Cyclops, Canthocamptus and Diaptomus furnished 15.5%,

and the cladocerans Daphnia, Leptodora, Chydorus and Bos¬

mina 16.5%- Mysis with 40% was a close second; the results

for it are included in the “miscellaneous” column in Table II.

This large percentage of Mysis agrees with results reported by

other investigators for ciscoes. Some plant material consisting

of Nostoc was found in some of the specimens. The smaller

ciscoes ate chiefly Entomostraea, but this type of food decreased

154 Wisconsin Academy of Sciences, Arts, and Letters .

sharply in the 190 mm. group, with a corresponding increase

in Mysis ; at 200 mm. then, Mysis decreased and Entomostraca

increased. The insect part of the food was consumed chiefly by

the larger ciscoes.

Other Lakes

Only a few specimens of fish were obtained from several

additional lakes, but the numbers were not large enough to give

any definite idea of the food relationships of the different sizes.

The general type of food material consumed by these specimens

is indicated in Table I and no further discussion of them is

necessary.

Summary and Discussion

From the data presented, it is evident that the food of young

fish differs considerably from that of the older ones. In some

species, notably the perch, rock bass, and black bass, three dis¬

tinct food periods may be observed. These periods are, first, one

when Entomostraca and small insect larvae are eaten, second,

when insects are the major food, and third, when fish and cray¬

fish are the chief foods.

Perea flavescens

As the perch were the most numerous and presented the

greatest variability of food in the different lakes, two graphs

for the two summers were prepared (Figs. 10 and 11). These

include all the perch studied from Trout, Muskellunge, Nebish,

Silver, Weber, Vieux Desert, and Long lakes ; these perch ranged

from 60 to 290 mm. in length. These curves are as significant

as those for the perch from each lake, for they are modified by

the unusual or different foods from certain lakes. One exam¬

ple of this is the 1932 Silver Lake perch which ate 90% fish as

compared with 6.5% fish in 1931. A second example is the

Nebish Lake perch, which are partly responsible for the high

insect curves, due to their eating so predominantly of insects,

especially Cqenis and Chironomus (1931—81% insects and 1932

— 53%). In general, however, insects were the principal food

of perch (1931 — 48%). Those for 1931 consisted of Diptera

larvae (13.5%), Ephemerida (14%), caddis fly larvae (7.5%),

Couey — Fish Food Studies

155

dragon fly nymphs (9.5%) Sialis larvae (1%), and miscellane¬

ous insects (8%). For 1932, insects were 42% and consisted

of Diptera larvae (10%), Ephemerida (4%), caddis fly larvae

(11%), dragon fly nymphs (3.5%), ants (1%), and miscellan¬

eous insects ( 13 % ) . Fish were the next important food, repre¬

senting about 22% in 1931 and 21% in 1932.

Fig. 10. This diagram shows the percentage composition of the food of differ¬

ent sizes of perch taken in all lakes in 1931. Compare with fig. 11.

Of note is the occurrence of the previously mentioned decline

in the percentage of insects (Fig. 11) which is found in the major

food curve of perch between 120 and 150 mm. in length. This

is a temporary decrease in insects and an increase of fish as

food. This raises a question as to why this particular size of

perch should suddenly turn cannibalistic.

From a study of the few small perch obtained it can be con¬

cluded that the very young fish eat Entomostraca, but quickly

change to an insect diet, consisting usually of small dipterous

larvae. These, along with other aquatic insect larvae, consti¬

tute the principal food until the perch is rather large. The

eating of fish as food begins early, but this usually does not be¬

come a major food until the perch has reached a length of 180

mm. or more.

As an example of the versatility of perch in feeding habits,

the foods of five perch from the same haul in Weber Lake in

1931 are cited. One perch, 185 mm. in length, contained 200

small caddis fly larvae ; one, 200 mm., contained a perch 65 mm.

156 Wisconsin Academy of Sciences, Arts, and Letters.

long; one, 196 mm., had eaten 28 large caddis fly larvae; one,

211 mm., had its stomach much distended with 250 Chaoborus

larvae; one, 155 mm., had its stomach half full of Ophryoxis.

Forbes and Richardson (1908) state that perch are wholly

carnivorous, but differ in their food according to habitat. River

perch fed on crayfish, insect larvae, small molluscs and fish, but

those from lakes ate entirely fish and crayfish. Reighard (1915)

found perch in Douglas Lake eating insects, fish and crayfish but

resorting to cannibalism in midsummer. This seems to be true

of the 1932 Silver Lake perch also, indicating that the condi¬

tions in the lake were not the most favorable for perch during

that year. Pearse (1918), in studying the food of shore fish in

lakes near Madison, found perch eating mostly insects (38.2%),

amphipods (24.5%), and entomostracans (25.8%). In 1920 the

same writer found insect larvae and entomostracans as the main

food. Clemens et al (1924) found in Lake Nipigon that the

perch below 4 cm. in length were plankton feeders and that above

this size, fish and insects rapidly increased.

Fig. 11. This diagram shows the percentage composition of the food of different

sizes of perch taken in all lakes in 1932. Compare with fig. 10.

Ambloplites rwpestris

In 1931 insects were the most important food of the rock

bass constituting about 43% and consisting of Diptera larvae

(4.5%), ephemerid larvae (6.5%), caddis fly larvae (5%),

dragon fly nymphs (11.5%), ants (4.5%), and miscellaneous

insects (11%). The Diptera larvae were mostly Chironomus

Couey — Fish Food Studies

157

and Chaoborus and the Ephemerida were Hexagenia with some

Coenis from Nebish Lake. Crayfish were the next important

food, amounting to about 36% of the total. This percentage is

rather high due to the fact that a few rock bass from some lakes

had eaten largely crayfish. Fish constituted 8% of the total

food.

In 1932 insects were also the principal food, making up

54% of the whole. They were similar to the 1932 foods, con¬

sisting of Diptera larvae (9%), Ephemerida nymphs (19.5%),

caddis fly larvae (4%), dragon fly nymphs (13%), Sialis larvae

( 1 % ) , ants ( 0.5 % ) , and miscellaneous insects ( 7 % ) . Fish were

next (9%) with Entomostraca (8%) third in importance. Cray¬

fish (6%) were not so important because fish were not taken

from those lakes where crayfish seemed to be so abundant the

previous summer.

It is evident from figures 3 and 7 that young rock bass eat

largely insects which give way to fish and crayfish in the larger

sizes. This change occurs around 140 mm. However, the rock

bass caught in October from Nebish Lake (Fig. 4) give a differ¬

ent picture in regard to the type of food eaten, showing that

there is a seasonal variation of food. Noteworthy is the large

amont of Entomostraca eaten by those from 90 to 150 mm. in

length and the increase of molluscs in the larger sizes. Seasonal

investigations in other lakes of this district should reveal some

interesting results.

Forbes and Richardson (1908) state that rock bass feed on

insects and small crustaceans, with a few fish. Hankinson

(1908) found that crayfish were the most important food of

these fish in Walnut Lake, Michigan. Baker (1916) found

crustaceans amounted to 75% of the food, the rest being insects

and plants (including algae). Pearse (1915) showed that the

food of young rock bass was largely insects and the adults

mostly crayfish, although insects were important. Evermann

and Clark (1920) examined 260 rock bass and found that those

under four inches in length had eaten Entomostraca, a few in¬

sect larvae and small fish, while the larger ones ate crayfish

and fish* Forbes (1880) mentions three small rock bass under

one inch which contained Cladocera, Cyclops, Chironomus and

Neuroptera larvae.

158 Wisconsin Academy of Sciences , Arts, and Letters.

The food of the rock bass seems to depend upon the habitat

of the fish. If it is in deep water, the foods will be much alike,

consisting of bottom fauna. If in shallow water, there will be

a greater variety, consisting of more insects (both surface and

aquatic), crayfish, fish and other forms which are abundant

in weed beds.

Helioperca macrochira

The food of bluegills is what one would expect from fish

adapted for life among tall, aquatic plants. In 1931 the food

of the bluegills of three lakes consisted of insects principally

(65% ) . They were Diptera larvae (24% ) , Ephemerida nymphs

(22.5%), caddis fly larvae (4%), dragon fly numphs (2%),

ants (7% ) , and miscellaneous insects (5.5% ) . The diptera were

mostly Chironomus and the ephemerids were Hexagenia. Plants

constituted 10% of the food of the bluegills, but made up 30%

of the food of those taken from Muskellunge lake. They consisted

of a small amount of filamentous algae (Spirogyra and Vau-

cheria) but mostly fragments of leaves and terminal buds of

large aquatic plants (Potamogeton and Elodea). The molluscs

were all snails (Physa, Planorbis and Amnicola).

The 1932 bluegills were all taken from Muskellunge Lake

and, as can be seen from Table II, their diet was about the

same as that of the previous summer.

An interesting relationship between the plant and insect

foods may be seen in Figure 8. Insects predominate except be¬

tween 90 and 140 mm., where plants constitute the most im¬

portant food- This is true only of the Muskellunge Lake blue¬

gills, for those from Silver and Allequash lakes had eaten no

plant food. Most of the bluegills from Muskellunge lake were

from the South and West bays which were quite shallow, and

aquatic plants were in great abundance.

Forbes (1880) found this species eating caddis fly larvae,

dragon fly nymphs, amphipods, entomostracans and crayfish.

Some fed largely on aquatic plants. Forbes and Richardson

(1908) found the bluegill food to be 45% insects, some crus¬

taceans, snails and a few fishes. Rjsighard (1915) found these

fish eating largely plant material with many insects, bryozoans

and a few hydrachnids and ostracods. He concluded as did

Couey — Fish Food, Studies

159

Forbes, that plants are a regular food and are not taken acci¬

dentally. Pearse (1915) found bluegills feeding mostly on in¬

sects (46.2%) and entomostracans (24.9%).

The food of the smallest bluegills consisted of small dipterous

larvae (mostly Chaoborus and Chironomus) with a few entomos¬

tracans. Moore (1920), however, reports the food of fingerlings

to consist of Cladocera, Copepoda, Ostracoda, chironomids and

nymphs of damsel flies and may flies.

Micropterus dolomieu

In 1931 insects were the chief food of the small-mouthed

bass, making up 47% of the diet, and fish were next (35%).

The insects consisted of Diptera larvae (5%), Ephemerida

(7%), caddis fly larvae (2.5%), dragon fly nymphs (3.5%),

ants (10.5%), corixids (3%), and miscellaneous insects

(15-5%). Hyalella was present as 3% and crayfish constituted

5% of the total food.

In 1932 fish were the main food (54%). Insects (25.5%)

were next, consisting of Diptera larvae (1% ) , ephemerid nymphs

(11.5%), caddis fly larvae (7%), dragon fly nymphs (2%),

ants (6%), and miscellaneous insects (8%). Crayfish made up

about 16.5% of the food.

In general the food of the small-muothed bass of this region

is similar to that described by investigators of other regions.

The young are not very well represented in this report but, from

the few obtained, it was found that small insect larvae, small

surface insects and Entomostraca form the main food. Insects

continued to be an important food until the small-mouths were

fairly large, at which time they were replaced by fish or cray¬

fish. Crayfish seem to be a favorite food when available. In

several lakes where crayfish were plentiful, they were an im¬

portant item in the diet of the small-mouthed bass. More surface

insects were found in this species of bass than in any other

species of fish. They consisted of grasshoppers and ants, with

lesser amounts of various other terrestrial insects. Although

fish were important as food, no cases of cannibalism were noted ;

the fish eaten, consisted of darters, minnows and small perch.

It is of interest to note that Corixidae, which have been reported

as quite important as food of small-mouthed bass (Forbes — 1880,

160 Wisconsin Academy of Sciences , Arts, and Letters .

Lydell 1904, and Pearse — 1918 and 1921a) were not found in

the specimens of this region although they were found in some

of the lakes. One small specimen (60 mm.) from Star Lake

had eaten three corixids and a small crayfish. The literature

on foods of small-mouthed black bass is well summarized by

Tester (1932).

Hnro floridana

Adult large-mouthed black bass seem to be quite piscivorous ;

fish made up about 60% of its food in 1931. The largest ones ate

almost entirely fish, but those around 100 mm. in length ate

largely insects. The small one (about 50 mm.) ate Cladocera,

with a few Cyclops and a few midge larvae.

The 1932 large-mouths were from Muskellunge Lake only

and they had eaten almost entirely fish with a few plants; the

latter were probably taken accidentally.

Fish and crayfish are reported by Forbes and Richardson

(1908), Tracy (1910), Reighard (1915) and Evermann and

Clark (1920) as the principal food of this fish. Hankinson

(1908) found crayfish to be the most important food in Walnut

Lake.

Although there were not sufficient quantities of the various

sizes of bass studied to show definitely where the changes in

diet occur, still it can be said in general that the food of the

smallest sizes consists of entomostracans, changing to insects

soon and finally to fish.

Stizostedion vitreum

The wall-eyed pike were all quite large and fed mostly on

fish. In 1931 fish made up 85% of the diet, consisting of darters,

minnows and perch. The insects were present as Diptera larvae

(1%), Ephemerida (6%), Entomostraca (4%) and plant mate¬

rial (1%). In 1932 the food was fish (85%), Ephemerida

(4.5%) and some caddis fly larvae, dragon fly nymphs, and cray¬

fish. As can be seen from Tables I and II, fish made up the

main food of the wall-eyed pike from all the lakes except Clear

Lake. Here insects were an important item of food, even in

the largest ones caught.

Forbes (1880) reports small pike eating entomostracans

and small fish, and adults eating fish entirely. Forbes and

Couey — Fish Food Studies

161

Richardson (1908) found fish the chief food along with a few

crayfish. Pearse (1918 and 1921a) reports fish as the principal

food with aquatic insect larvae occasionally found. Clemens et

a l (1924) found small pike (3 to 10 mm.) had eaten mostly En-

tomostraca, a few Chironomus larvae and a few fish. Those 10

to 40 mm. long had eaten largely Ephemerida nymphs (Hexa-

genia) and some Trichoptera larvae, with some fish scattered

throughout. Those above 40 mm. ate fish entirely. Adamstone

(1924) found fish usually as food but occasionally Ephemerida,

Trichoptera and Odonata nymphs.

Cristivomer namaycush

The food of the lake trout was entirely fish, mostly ciscoes

with a few suckers. No small ones were examined. Clemens

et al (1924) found about the same food. Although the specie*

of small fish eaten were more varied, ciscoes were the main

food.

Esox masquinongy

The food of the muskellunge was found to be entirely fish,

consisting of perch, darters, and chubs. No very small ones

containing food were taken.

Leucichthys artedi

Ciscoes are preeminently plankton feeders. Tables I and II

show that entomostracans were the principal food taken by them.

Exception may be taken here to the ciscoes of Black Oak Lake

in which Mysis was an important article of diet (40% Mysis

and 41% entomostracans). Mysis has been reported by other

investigators as a favorite food of ciscoes, but Black Oak Lake

is the only one in this region in which this crustacean played an

important role.

The 1931 ciscoes had eaten about 47.5% Cladocera, 34%

Copepoda and small amounts of Diptera larvae. In the 1932

ciscoes, Cladocera made up 54.5%, Copepoda 25%, Ostracoda

2%, Mysis 10%, with small amounts of dipterous larvae.

Clemens et al (1924) found ciscoes feeding chiefly on Mysis

relicta and, in smaller amounts, on Limnocalanus macrurus.

Those taken in shallow bays fed on Daphnia and other plankton

animals. Adamstone (1924) reported that bottom forms were

162 Wisconsin Academy of Sciences, Arts, and Letters.

eaten occasionally, especially by the younger specimens; these

included mysids, amphipods, cladocerans, copepods, ostracods,

chironomids, ephemerids, hydrachnids and molluscs. Eawson

(1928) found that the ciscoes were essentially plankton feeders

in Lake Simco, but they took some bottom forms also. The

latter included emphemerids, chironomids, amphipods and oli-

gochaets.

Cor eg onus clupeaformis

Whitefish were taken only in Trout Lake; they proved to be

chiefly bottom feeders. The principal foods, in order of im¬

portance, were Diptera larvae, molluscs and Entomostraca. In¬

cluded in the “miscellaneous” column of Table I is Mysis

(11%). In the same column of Table II is Rotifera (14%) and

Mysis (5%). Also there was a considerable amount of bottom

debris consisting of ooze, plant fragments, pebbles, and some

unidentifiable fish eggs.

Clemens et al (1924) state that the food of whitefish, in order

of importance, was Pontoporeia, Chironomus larvae, molluscs and

Ephemerida nymphs. Hart (1931) found the chief items of

food of adults to be amphipods, molluscs and insect larvae. The

food of young whitefish is reported to be Entomostraca by

Forbes (1883) and Hankinson (1916). Hankinson (1911) re¬

cords whitefish of Walnut Lake as eating mostly Chironomus

larvae and Daphnia and he notes a seasonal variation.

Catostomus commersonii

Of the 1931 suckers, molluscs (39%) were the main food,

with Diptera larvae (23%) second and Entomostraca (7%)

third. Debris was 19%. In 1932 Diptera were 27%, molluscs

24.5% and Entomostraca 12%. Debris was 30%.

The sucker eats a larger variety of food than any other

species examined. It is entirely a bottom feeder, which accounts

for so much mud and bottom debris in its food. Pearse says,

“the sucker is remarkable for the fineness of the food it is able

to select. No other fish shows such a high percentage of pro¬

tozoans, unicellular algae and rotifers in its food” (1915). Hank¬

inson (1908) found the suckers in Walnut Lake eating caddis

fly larvae, midge larvae and other insects, small bivalve moL

Couey — Fish Pood Studies 163

luscs, amphipods and Entomostraca. In Oneida Lake, Baker

(1916) found that the suckers had eaten mud, plant remains,

molluscs and insects. Bigelow (1924) found that the young

suckers from Lake Nipigon were plankton feeders. Clemens

et al (1924) state that, as the sucker grows, it takes more and

more of the larger bottom organisms such as Chironomus larvae

and later ephemerid nymphs, caddis fly larvae, molluscs, and

amphipods. Algae and diatoms are also reported as being eaten.

Bibliography

No attempt has been made to give a complete summary of the literature as

that has been done quite adequately by several recent workers. C. C. Adams and

T. L. Hankinson (1928) in their “Ecology and Economics of Oneida Lake Fish” give

a complete survey of the literature, including most of the species of fish mentioned in

this paper. Therefore, in the preceding discussion and summary, reference was made

to only a few authors for purposes of comparison.

Adams, C. C. and T. L. Hankinson. 1928. The ecology and economics of Oneida

Lake fish. Roosevelt Wild Life Annals. Vol. I.

Adamstone, F. B. 1924. The distribution and economic importance of the bottom

fauna of Lake Nipigon. Univ. Toronto Stud.: Biol. Ser., Pub. Ont. Fish. Res.

Lab. No. 25.

Baker, F. C. 1916. The relation of mollusks to fish in Oneida Lake. N. Y. State

Col. For. Tech. Pub. No. 4.

Bigelow, N. K. 1924. The food of young suckers (Catostomus commersonii) in

Lake Nipigon. Univ. Toronto Stud.: Biol. Ser., Pub. Ont. Fish. Res. Lab.

No. 21.

Clemens, W. A., J. R. Dymond and N. K. Bigelow. 1924. Food studies of Lake

Nipigon fishes. Univ. Toronto Stud.: Biol. Ser., Pub. Ont. Fish. Res. Lab.

No. 25.

Evermann, B. W. and H. W. Clark. 1920. Lake Maxinkuckee, a physical and

biological survey. Indiana Dept. Conserv. Vol. I and II.

Forbes, S. A. 1880. The food of fishes: Acanthopteri. Bull. Ill. State Lab. Nat.

Hist. 1 (3): 19-70.

Forbes, S. A. 1883. The food of smaller freshwater fishes. Bull. Ill. State Lab.

Nat. Hist. 1 (6): 65-94.

Forbes, S. A. and R. E. Richardson. 1908. The fishes of Illinois. Nat. Hist. Sur¬

vey of Ill. Ichthyology. Vol. in.

Hankinson, T. L. 1908. A biological survey of Walnut Lake, Michigan. Mich.

State Biol. Survey. Rep. 1907.

164 Wisconsin Academy of Sciences, Arts, and Letters .

Hankinson, T. L. 1911. Ecological notes on the fishes of Walnut Lake, Michigan.

Trans. Amer. Fish. Soc. 40:195-206.

Hankinson, T. L. 1914. Results of Shiras expeditions to Whitefish Point, Michigan:

Fishes. Mich. Geol. and Biol. Survey. Pub. 20, Biol. 4.

Hart, J. L. 1931. The food of the whitefish, Coregonus clupeaformis (Mitch.),

in Ontario waters. Contrib. Can. Biol. & Fish. N. S. 6:447-453.

Juday, C. and E. A. Birge. 1930. The Highland Lake District of northeastern

Wisconsin and the Trout Lake limnological laboratory Trans. Wis. Acad.

Sci., Arts & Let. 25:337-352.

Lydell, D. 1904. The habits and culture of black bass. U. S. Fish. Com. Bui.

22:39.44.

Moore, Emmeline. 1920. Plants of importance in pond fish culture. Rept. U. S.

Commr. Fish, for 1919, Append. 4:5-20.

Pearse, A. S. 1915. On the food of the small shore fishes in the waters near Madison,

Wisconsin. Bui. Wis. Nat. Hist. Soc. 13:7-45.

Pearse, A. S. 1918. Food of the shore fishes of certain Wisconsin lakes. Bui. Bur.

Fish. 35:249-292.

Pearse, A. S. 1920. Habits of the yellow perch in Wisconsin lakes. Bui. Bur.

Fish. 36:297-366.

Pearse, A. S. 1921. The distribution and the food of the fishes of three Wisconsin

lakes in summer. Univ. Wis. Stud. Sci. No. 3:1-53.

Rawson, D. S. 1928. Preliminary studies of the bottom fauna of Lake Simco,

Ontario. Univ. Toronto Stud. Biol., Pub. Ont. Fish. Res. Lab. No. 36.

Reighard, J. 1915. An ecological reconnoissance of the fishes of Dougles Lake,

Cheboygan County, Michigan, in mid-summer. Bui. Bur. Fish. 33:215-249.

Tester, A. L. 1932. Food of the small-mouthed black bass (Micro pterus dolomieu)

in some Ontario' waters. Univ. Toronto Stud. Biol., Pub. Ont. Fish. Res. Lab.

No. 46.

Tracy, H. C. 1910. Annotated list of the fishes known to inhabit the waters of

Rhode Island. Fortieth Ann. Rept. Comm, of Inland Waters of Rhode Island,

pp. 35-176.

TABLE I

Percentage composition of food of different species of fish taken in various lakes in 1931 .

Couey — Fish Food Studies 165

166 Wisconsin Academy of Sciences , Arts, and Letters.

Coney — Fish Food Studies 167

168 Wisconsin Academy of Sciences , Arts , and Letters .

Couey — Fish Food Studies

169

Table II

Percentage composition of food of different species of fish taken in various lakes in 1932.

170 Wisconsin Academy of Sciences , Arts, and Letters,

Coney— Fish Food Studies 171

172 Wisconsin Academy of Sciences , Arts , and Letters.

PHOTOSYNTHESIS OF ALGAE AT DIFFERENT DEPTHS

IN SOME LAKES OF NORTHEASTERN WISCONSIN

I. OBSERVATIONS OF 1933

Harold A. Schomer and Chancey Juday

From the Limnological Laboratory of the Wisconsin Geological and Natural

History Survey. Notes and reports No. 59.

Introduction

Observations on the penetration of solar radiation into the

waters of the inland lakes have been carried on more or less con¬

tinuously by the Wisconsin Survey since 1900. In the earlier

years, these studies were more closely related to the problem

of the thermal changes which take place in lake waters and

that of the annual heat budget; they also included the problem

of the manner in which the heat is distributed to different strata

in lakes.

At the same time, the question of the relation of solar radi¬

ation to the process of photosynthesis in the various aquatic

plants at differ en depths was kept in mind. No work was done

on this problem, however, until the summer of 1932, when

observations were made on the photosynthetic activities of two

cultures of green algae and of a few of the large aquatic plants

in three of the northeastern lakes. A preliminary report on

the results obtained in these experiments has recently been pub¬

lished (Schomer 1934). These studies were continued during

the summer of 1933 with the algal cultures and the present re¬

port is based upon the results obtained in this investigation.

Method

The general plan of the investigation was essentially the

same as that of Marshall and Orr (1928). Algal cultures were

placed in bottles and suspended at different depths from a buoy

for a definite period of time and the quantity of oxygen produced

during the interval was determined by a modified Winkler

method. In some cases, hydrogen ion readings and carbon diox¬

ide titrations were made.

173

174 Wisconsin Academy of Sciences, Arts, and Letters.

Apparatus. The glass stoppered bottles had a capacity of

about 150 cc and half of them were painted with a water-proof

black paint so that they could be used as controls in determining

the amount of oxygen used in respiration during the progress of

the experiment. While suspended in the lake, these controls

were also enclosed in black cloth bags in order to prevent the

reflection of light from their surfaces.

The bottles were suspended in coarse mesh galvanized wire

baskets which were large enough to hold four of them, two

samples and two controls; duplicates were run at each depth.

The buoy from which the bottles were suspended, consisted of

two floats connected by a half-inch galvanized iron pipe about

3 m. long. The buoy was anchored in a north-south direction so

that the shadows of the floats would not fall upon the algal cul¬

tures. The surface basket was attached to one of the floats by

means of an adjustable supporting wire.

Cultures. Pure cultures of two species of algae were used

for the experiments, namely Coccomyxa simplex and Chlorella

pyrenoides. Several nutrient solutions containing carbohydrate

compounds were tried, but it was not possible to produce suffi¬

cient quantities of the algae in these media under sterile condi¬

tions; the cultures became contaminated with fungi. Several

mineral nutrient media were then tried and the following one

proved to be very satisfactory for the cultivation of these two

forms :

KHaPO* _ _ __2.72 grams

MgSO* _ _ 4.93 grams

Ca(N03)2 _ 4.72 grams

These salts were dissolved in distilled water and made up to

ten liters. A few drops of FeCh solution were added to each liter.

Abundant growths of these algae were obtained in a few days in

this culture medium; the cultures were kept in large glass cov¬

ered battery jars on an outdoor stand which was shaded by

trees.

Procedure* The algae were removed from one liter of the

culture medium by means of a centrifuge and they were then

added to 4 1. of filtered surface water from Trout Lake; this

water contains a good supply of raw materials necessary for

Schomer & J uday — Photosynthesis of Algae 175

photosynthesis. The mean free carbon dioxide content is 1*3

mg/1 and the bound carbon dioxide 18.8 mg/1 ; the Ca averages

9.6 mg/1, the Mg 6.4 mg/1 and the hydrogen ion pH 7.5.

The algal suspension in the filtered lake water was thor¬

oughly stirred and the bottles were filled ; care was used in clos¬

ing the bottles in order to avoid the inclusion of air bubbles.

Two clear and two black bottles were then placed in each basket

and the baskets were stored in a light-tight box, painted black

on the inside, for transportation to the buoy where they were

suspended; at the end of three hours, the cultures were taken

up and the chemicals were added at once in the boat. In order

to obtain the oxygen content of the samples at the beginning of

the experiment, two control samples were taken along and the

chemicals added to them at the time the cultures were suspended

from the buoy.

The exposures were limited to three hours because gas bub¬

bles were noted in some of the bottles when longer periods were

used; this was true especially in the strata showing optimum

photosynthesis. Samples of the algal suspension were removed

at the time the bottles were filled and preserved for the purpose

of enumerating the organisms.

SOLARIMETEB READINGS

In 1932, observations were made on the amount of solar and

sky radiation delivered to the surface of the lake during the

progress of the experiments on photosynthesis; this was done

by taking solarimeter readings at regular intervals during each

experiment. In 1983 however, a self-registering instrument

was used, so that the total quantity of energy reaching the sur¬

face of the lakes during the experiments was obtained ; this ap¬

paratus was also operated for periods of 12 to 24 hours. Some

of the records are represented in Figures 1, 2 and 3.

Figure 1 shows the results obtained on July 6, 7 and 9, 1983.

On July 6, the sky was only slightly cloudy at times, while

July 7 was a cloudy day. There was an alternation of clouds

and sunshine on July 9 ; the highest reading of the summer was

obtained about noon on this date* In Figure 2, August 2 was

a cloudy day, while August 4 and August 5 had both clouds and

sunshine. August 7 in Figure 3 was characterized by clouds

176 Wisconsin Academy of Sciences , Arts, and Letters .

and sun. On August 23 (Fig. 3), the sky was fairly clear until

about 1:15 p.m., after which there was an alternation of clouds

and sunshine ; the experiment performed on this date was term¬

inated soon after the clouds appeared.

JULY 7

JULY 9

Fig. 1. The curves in this figure show the amount of solar and sky radiation

at Trout Lake on July 6, 7 and 9, 1933, between 4:00 a.m. and 8:00 p.m.

Schomer & Juday — Photosynthesis of Algae 177

The solarlmeter used in making these records was stand¬

ardized against a Callendar pyrheliometer which has been in

continuous use by the Weather Bureau at Madison, Wisconsin,

for more than twenty years. This standardization made it

possible to determine the quantity of radiation delivered to the

Fig. 2. The curves in this figure show the amount of solar and sky radiation

at Trout Lake during a period of eight hours on August 2, 4 and 5, 1933.

178 Wisconsin Academy of Sciences , Arts , and Letters.

lake surface during any period of the day. The amount of this

energy is indicated in terms of the unit employed by the United

States Weather Bureau, namely the gram calorie per square

centimeter per minute; for the sake of brevity, the word “cal¬

orie” is used in the following discussion instead of the complete

expression.

Observations were made with a pyrlimnometer to determine

the percentage transmission of the radiation to the various

strata; from these results, the quantity of energy penetrating

to the various depths has been computed.

For these experiments on photosynthesis, four lakes were

selected which covered the range from very clear, transparent

water, which transmitted the maximum amount of radiation, to

that which has a deep brown color due to the presence of veget¬

able stains derived from peat and bog deposits and which cuts

AUGUST 7

AUGUST 23

Fig. 3. These curves show the amount of solar and sky radiation at Trout

Lake on August 7 and 23, 1933.

Schomer & Juday — Photosynthesis of Algae 179

off the solar energy very rapidly. On the platinum-cobalt scale,

the brown color of these lake waters ranged from zero in Crys¬

tal Lake to 168 in Helmet Lake.

Data Obtained

Crystal Lake

This lake has an area of 30.2 ha. and a maximum depth of 21

m. It has neither an inlet nor an outlet and the shores consist

of sand. The water is very soft and transparent- A maximum

disc reading of 13.6 m. has been obtained on this lake and the

reading at the time of the experiment was 13.5 m. The water did

not have any brown color whatever.

On August 4, 1933, a series of samples of Coccomyxa sim¬

plex was suspended at various depths down to 15 m. between

11:00 a.m. and 2:00 p.m. On August 5, cultures of Chlorella

pyrenoides were used between 11:00 a.m. and 2:00 p.m.; this

series extended to a depth of 20 m. A second series of Coc¬

comyxa cultures was used on August 7 and it extended to a depth

of 20 m- also ; the time interval was 10 :45 a.m. to 1 :45 p.m. The

solarimeter records for these three dates are shown in Figures

2 and 3.

The curves in Figure 4 show the results obtained in these

three experiments; they are based on the quantity of oxygen

produced per million cells of algae at the different depths. They

indicate that the solar energy was too great in the upper water

for maximum photosynthesis at the time of day and under the

weather conditions which prevailed during these experiments.

Thus there was a gradual increase in the oxygen production from

the surface to a depth of 5 m. in two series and to 6 m. in the

third. The maximum yield of oxygen on August 4 was 0.426

mg. per million cells, 0.528 on August 5 and 0.501 on August 7.

(Table 1)- Below these optimum depths, there was a gradual

decline in the oxygen production with increasing depth; at 17

m. it had fallen to the point where the oxygen produced in pho¬

tosynthesis was just equal to the amount consumed in respira¬

tion and decomposition. This depth is known as the compensa¬

tion point. It is indicated by a plus sign in curves B, and C;

series A did not go deep enough to show this point. A certain

amount of oxygen was produced below this depth, but the quan-

180 Wisconsin Academy of Sciences , Arts, and Letters .

tity was less than that consumed in respiration and decompo¬

sition. It may be noted in this connection, however, that three

species of moss thrive on the bottom of Crystal Lake at depths

of 18 to 20 m. (Juday 1934).

Fig. 4. Photosynthesis of algal cultures at different depths in Crystal Lake

during three hour periods on August 4 (A), 5 (C) and 7 (B), 1933. Curves A and

B represent Coccomyxa simplex and curve C, Chlorella pyrenoides. The curves

show the amount of oxygen produced per million cells at the different depths.

The total quantity of solar energy delivered to the surface of

Crystal Lake during the three hour periods of these experi¬

ments was as follows: — August 4, 124.9 cal.; August 5, 134.9

cal.; August 7, 141.4 cal. Pyrlimnometer readings taken on

July 29 showed that 0.9 per cent of the solar energy delivered

to the surface of the lake penetrated to a depth of 17 m., which

was the compensation point. On this basis, 1.12 cal/cm2 reached

17 m. during the three hour period on August 4, 1.21 cal. on

August 5 and 1.27 cal- on August 7. Thus it required an aver¬

age of 1.20 cal/cm2 of solar energy during a period of three

hours in the middle of the day to produce enough oxygen at 17

m. to balance the amount consumed in respiration and decompo¬

sition.

About 14.5 per cent of the solar radiation penetrated to a

depth of 5 m. and 12 per cent to 6 m. ; these are the two depths

Schomer & Juday —Photosynthesis of Algae 181

at which maximum oxygen production was found. In the 5 m.

maxima for Coccomyxa, 18.1 cal/cm2 reached this depth in three

hours on August 4 and 20.5 cal. on August 7. For the Chlorella

experiment on August 5, 16.2 cal. reached 6 m., the point of

maximum oxygen production, in three hours. The radiation at

5 m. consisted of 16 per cent violet, 20 per cent blue, 27 per cent

green, 23 per cent yellow, 10 per cent orange and 4 per cent

red; thus 70 per cent of it fell in the blue-green-yellow part of

the spectrum.

Table 1 shows that the temperature of Crystal Lake ranged

from a maximum of 22.9° C. at the surface to a minimum of

11.3° at the bottom. The temperatures at the depths of optimum

oxygen production were substantially the same as those at the

surface. The differences in temperature at the different depths

apparently had little effect upon respiration, so that the mean

of the entire series of black bottles was used in each case in

making a correction for the oxygen consumed in this process.

Trout Lake

Trout Lake has an area of 1,583 ha. and a maximum depth

of 35 m. The water is usually slightly stained with vegetable

extractives brought in by some of the tributary streams. The

color of the surface water ranges from zero to 14 on the plati¬

num-cobalt scale; it was 14 on August 29, 1933. The water is

much less transparent than that of Crystal Lake ; the disc read¬

ings in Trout Lake vary from 3.3 to 6.5 m., with a mean of 4.5

m. A reading of 5.9 m. was recorded on August 29, 1933.

Figure 5 shows the results obtained on August 23 and 28,

1933. Chlorella was used on the former date and Coccomyxa on

the latter. The cultures were exposed from 10:45 a.m. to 1 :45

p.m. on August 23 and from 11 :25 a.m. to 2 :25 p.m. on August

28. Figure 3 shows the solar energy record for August 23 ; on

this date the sky was clear until about 1:15 p.m., with alter¬

nating clouds and sunshine during the remainder of the after¬

noon. Thus the three hour experimental period extended into

the partly cloudy time only about half an hour. There was an

unusual drouth during the summer and this was accompanied by

many forest fires in the month of August, so that the air con¬

tained varing amounts of smoke during this time. This smoke

182 Wisconsin Academy of Sciences, Arts, and Letters .

reduced the amount of radiant energy that reached the surface

of the lake ; on some days, in fact, the smoke was so dense that

it caused a very material reduction in the radiation.

Fig. 5. Photosynthesis of algal cultures at different depths in Trout Lake during

three hour periods on August 23 (B) and 28 (A), 1933. Curve A represents

Coccomyxa simplex and curve B, Chlorella pyrenoides. The curves show the

amount of oxygen produced per million cells at the different depths.

Curve A of Figure 5 represents the results obtained with

Coccomyxa cultures on August 28. The maximum oxygen pro¬

duction was found at 1 m., namely 0.145 mg. per million cells.

The compensation point fell at 10. m. Curve B indicates the

oxygen yield of Chlorella on August 23. The oxygen produc¬

tion was larger on this date than on August 28; this was ac¬

counted for in part, at least, by a larger amount of solar radia¬

tion on the former date. A total of 190.4 cal. reached the sur¬

face of Trout Lake during the three hour period on August 23

and only 122.1 cal. on August 28.

The maximum production of oxygen was noted at 2 m. on

August 23 and it amounted to 0.225 mg. per million cells. Sub-

Schomer & Juday — Photosynthesis of Algae 183

stantially the same quantity of oxygen was produced at 3 m. ;

in fact, there was very little difference in the yields at 1 m.,

2 m. and 3 m. (Table I). The compensation point for both

series of algae was found at 10 m.

About 28 per cent of the solar energy incident on the sur¬

face of Trout Lake during mid-day penetrates to a depth of

1 m., 17 per cent to 2 m., 11.2 per cent to 3 m. and 1 per cent to

10 m. On this basis, the quantity of solar energy reaching the

various depths during the three hour experimental period was

as follows: —August 23, 53.3 cal/cm2 at 1 m., 32.4 at 2 m., 21.3

at 3 m. and 1.9 at 10 m. ; August 28, 34.2 at 1 m., 20.7 at 2 m.,

13.7 at 3 m. and 1.22 at 10 m.

The maximum oxygen production of Chlorella at 2 m. in

Trout Lake on August 23 was correlated with 32.4 cal., while

that at 6 m. in Crystal Lake on August 5 was correlated with

16.2 cal., or just half as much solar energy. Furthermore, the

oxygen production in Crystal was more than twice as great as

that in Trout, namely 0.528 mg. per million cells in the former

and 0.225 mg. in the latter. A similar difference was noted

in the Coccomyxa cultures. The maximum of 0.145 mg. per

million cells at 1 m. in Trout Lake on August 28 was correlated

with 20.7 cal., while those at 5 m. in Crystal Lake, 0-426 and

0.501 mg., were correlated with 18.1 and 20.5 cal. on August 4

and 7, respectively.

The total energy delivered to the compensation depths in

three hours was not very different, however; it was 1.56 cal.

at 10 m. in Trout Lake and 1.20 cal. at 17 m. in Crystal Lake.

At 1 m- in Trout Lake, the percentages of radiation belong¬

ing to the different parts of the spectrum were as follows: —

violet 9 per cent, blue 17, green 18, yellow 21, orange 20 and

red 15.

Three series of experiments were run in the upper 5 m. of

Trout Lake on August 24, 1933; they began at 7:30 a.m. and

were continued until 4:30 p.m. (Table I). Coccomyxa cultures

were used for the three series. A maximum oxygen production

of 0.221 mg. per million cells was obtained at 2 m. in the first

series in the period from 7 :30 to 10 :30 a.m. In the mid-day

series, 10:30 a.m. to 1:30 p.m., the maximum yield was 0.242

mg. per million cells at 3 m. ; in this series, however, there was

184 Wisconsin Academy of Sciences , Arts , and Letters .

very little difference in the yield between 1 m. and 4 m. The

largest production in the 1:30-4:30 p.m. series was found at

1 m. ; it amounted^ to 0.233 mg. per million cells. None of the

series went deep enough to determine the compensation point.

It is interesting to note that the yield of the surface samples

was smallest in the mid-day series ; it was less than half as much

as that of the afternoon series.

The amount of solar energy delivered to the surface during

each of the experiments was smallest in the forenoon series

and largest in the mid-day series; it amounted to 70.9 cal in

the former and 176.8 cal. in the latter series.

Two series were run on August 26; the first one covered

the period from 8:45 to 11 :45 a.m. and the second from 2 :00

to 5:00 p.m. The largest production of oxygen in the forenoon

series was 0.300 mg. per million cells at 25 m. and in the after¬

noon series 0.257 mg. at 3 m.

Mud Lake

This is a small body of water with an area of only 5.48 ha-

and a maximum depth of 15.7 m. It has neither an inlet nor an

outlet, so that the water is soft; the specific conductance of

the surface water averages about 16 X 10 6. The shores are

somewhat boggy in places and the water receives a certain

amount of brown vegetable stain from these bog areas ; as a result

the water has a color of 33 on the platinum-cobalt scale. The

transparency is rather low; the disc reading was 1.9 m. on

August 14, 1933.

A series of Chlorella cultures was suspended in Mud Lake on

August 15, 1933, from 11 :45 a.m. to 2 :45 p.m. and one of Coe-

comyxa culture on August 19, from 11:10 a.m. to 2:10 p.m.

Both species of algae yielded substantially the same amounts

of oxygen in the upper 2 m. as shown in the curves of Figure 6,

but Chlorella gave a somewhat larger yield between 2 m. and

10 m. The optimum production was found at 0.5 m. ; the amount

produced at this depth was 0.329 mg. per milllion cells on Aug¬

ust 15 and 0.324 mg. on August 19. (Table I). The com¬

pensation point fell at the same depth for both series, namely

4.5 m.

Schooner & Juday — Photosynthesis of Algae

185

Fig. 6. Photosynthsis of algal cultures at different depths in Mud Lake during

three hour periods on August IS (B) and 19 (A), 1933. Curve A represents

Coccomyxa simplex and curve B, Chlorella pyrenoides. The curves show the amount

of oxygen produced per million cells at the different depths.

Pyrlimnometer readings taken on August 14 showed that

22-5 per cent of the radiation incident on the surface of Mud

Lake penetrated to a depth of 0.5 m., which was the depth of

maximum oxygen production; only 13 per cent of this energy

was left at 1 m., 3.7 per cent at 2 m. and 0.4 per cent at 4.5 m.

The total radiation incident on the surface during the three hour

period on August 15 was 191.0 cal/cm2 and 207.2 cal. on August

19. On August 15, therefore, about 43.0 cal. of this energy

reached a depth 0.5 m. and only 0-76 cal. penetrated to 4.5 m. ;

on August 19, the respective amounts were 46.6 cal. at 0.5 m.

and 0.83 cal. at 4.5 m.

Helmet Lake

This is a typical bog lakelet; it has an area of 3.0 ha. and a

maximum depth of 10.4 m. There is no inlet or outlet and the

water is soft; the specific conductance was 19 X 10*6 on August

8, 1933. It has boggy shores and the water is deeply stained

with vegetable extractives derived from the peat. On August

186 Wisconsin Academy of Sciences , Arts, and Letters .

11, the color of the surface water was 168 on the platinum-co¬

balt scale. In some years the water is more highly colored; a

maximum reading of 268 was obtained in 1980. The transpar¬

ency is low also; the disc reading on August 8, 1938, was 1.5

m. Readings as low as 0.8 m. have been obtained in other years.

Five series of cultures were used in Helmet Lake and the

results for three of them are shown in the curves of Figure 7.

On August 10, the Coccomyxa cultures were exposed from 11 :25

a.m. to 2:25 p.m. On August 11, Chlorella cultures were used

from 10 :45 a.m. to 1 :45 p.m. and Coccomyxa again from 11 :25

a.m- to 2 :25 p.m. on August 14. The curves in Figure 7 show

that there was a marked difference in the quantity of oxygen

produced in these three experiments.

0.1 0.2 0.3 0.4

Fig. 7. Photosynthesis of algal cultures in Helmet Lake during three hour

periods on August 10 (A), 11 (B) and 14 (C), 1933. Curves A and C represent

Coccomyxa simplex and curve B Chlorella pyrenoides. The curves show the amount

of oxygen produced per million cells at the different depths.

On August 10, the sky was overcast and it rained while the

experiment was in progress. The results for this series are

shown in curve A. As might be expected under these weather

conditions, the maximum production of oxygen was found in

the surface sample; it amounted to 0.298 mg. per million cells

for the three hour period. There was a rapid decline in oxygen

Schomer & Juday — Photosynthesis of Algae 187

production below the surface ; by far the greater part of the de¬

crease was noted between 0.25 and 0.75 m. The compensation

point was found at 1 m. The solarimeter was not in operation

on this day, so that the amount of radiation reaching the sur¬

face of the lakelet was not determined.

The Chlorella culture gave a maximum yield of oxygen at

a depth of 0.25 m. on August 11 ; it was 0.254 mg. of oxygen per

million cells for the three hour period- (See curve B in Figure

7). Below this depth the production decreased rapidly to the

compensation point at 1 m. The radiation delivered to the sur¬

face of the lakelet during this experiment amounted to 191.1

cal/cm2 and 14 per cent penetrated to a depth of 0.25 m., the

depth of maximum photosynthesis; this was equivalent to 26.7

cal. in three hours. The maximum oxygen yield in this series

was somewhat smaller than that of the surface sample on

August 10.

The optimum oxygen production on August 14 was obtained

at 0.25 m., but it was larger than in the other two series, namely

0.413 mg. per million cells. (Curve C in Figure 7). The solar

energy reaching the surface of Helmet during the experimental

period on this day amounted to 236.3 cal- and 33.1 cal. penetrated

to 0.25 m. This is approximately 25 per cent more solar energy

than reached this depth on August 11 ; the oxygen yield, on the

other hand, was about 60 per cent larger on August 14 than on

August 11.

The compensation point was situated at a depth of 1 m. in

the three Helmet Lake experiments. The amount of solar energy

reaching this depth in three hours was 2.86 cal. on August 11

and 3.54 cal. on August 14. These quantities were considerably

larger than those found at the compensation points in the other

three lakes ; in the latter, the amounts ranged from a minimum

of 0.76 cal. in one series of Mud Lake to a maximum of 1.90

cal. in a Trout Lake series. In such highly colored water as

that of Helmet Lake, more than 80 per cent of the energy pene¬

trating to a depth of 1 m- falls in the orange-red part of the

spectrum and apparently it is not as effective in the process of

photosynthesis.

188 Wisconsin Academy of Sciences , Arts , and Letters.

Utilization of Solar Radiation in Photosynthesis

The oxygen production gives a basis for the computation of

the percentage of energy utilized by the algae in the process of

photosynthesis; the oxygen yield per square centimeter of cell

surface at the different depths is readily converted into terms

of glucose with a combustion value of 3760 calories per gram.

In Crystal Lake, the percentage of energy utilized by the

algae gradually increased from about 0.5 per cent at the surface

to a maximum of 9.2 per cent of that penetrating to a depth of

10 m. It ranged from 0.05 per cent at the surface in Trout Lake

to a maximum of 4.3 per cent at 7 m. in the Chlorella series and

from 0.09 per cent at the surface to 6.2 per cent at 10 m. in the

Coccomyxa series.

The highest percentage of utilization was found in the highly

colored water of Helmet Lake. In the Chlorella series of Aug¬

ust 11, it ranged from 0.16 per cent at the surface to 9.5 per

cent at 1 m. and in the Coccomyxa series of August 14, from 0.23

per cent at the surface to 11.3 per cent at 1 m. In the Chlorella

series, the utilization declined to 4.6 per cent at 2 m. and in the

Coccomyxa series of August 14, from 0.23 per cent at the surface

to 11.3 per cent at 1 m. In the Chlorella series, the utilization

declined to 4.6 per cent at 2 m. and in the Coccomyxa series to 6.3

per cent at 2 m.

Results of Other Investigators

Gaarder and Gran (1927) used marine plankton rich in

diatoms in some photosynthesis experiments at various depths

in the latter part of March ; they obtained substantially the same

results at 2 m. as at the surface and the 5 m. sample was only

5-10 per cent below the maximum. The compensation point was

reached at 10 m.

Marshall and Orr (1928) made a large number of experiments

on the photosynthesis of diatom cultures suspended at different

depths in the sea at the Millport Marine Station. Since the

methods employed in the Wisconsin investigations were sub¬

stantially the same as those of these investigators, some direct

comparisons of results can be made. Marshall and Orr found

a maximum oxygen production of 0.393 mg. per million diatom

cells at 6 m. on June 10, 1927, from 12:00 to 2:57 p.m. and a

Schomer & Juday— Photosynthesis of Algae 189

maximum of 0.404 mg. was noted at 2 m. on the same date in

the 3:00-5:55 p.m. series. A somewhat larger maximum was

obtained at 2 m. in a 9:00-12:00 a.m. series on June 28, namely

0.422 mg. per million diatom cells. The light intensity was much

greater, however, on June 10 than on June 28; on the former

date, 390.4 mg. of oxalic acid were decomposed during the three

hours of the experiment and only 119.8 mg. in the experimental

period on the latter date.

In comparison with this, a maximum oxygen production

of 0.426 mg. per million Coccomymxa cells was obtained at 5 m.

in Crystal Lake from 11:00 a.m. to 2 : 00 p.m. on August 4, 1933

and one of 0.528 mg. per million cells of Chlorella at 6 m. during

the same period on August 5. The total quantity of solar energy

delivered to the surface of the lake was much the same on the

two days, namely 124.9 cal/cm2 during the three hours of Aug¬

ust 4 and 134.9 cal/cm2 during that of August 5.

Marshall and Orr found maximum oxygen production at

the surface on dark days, but at depths of 2 m. to 6 m. on bright

days, especially in summer. Similar results were obtained on

Crystal and Trout lakes, but the maximum yield was found at

0.5 m. and 0.25 m. in the colored waters of Mud and Helmet

lakes. They also found that the light intensity was too great on

bright days in winter for maximum oxygen production at the

surface. The compensation depths in their experiments fell in

the 20-30 m. stratum in mid-summer. In the Wisconsin lakes

it ranged from a maximum of 17 m. in Crystal Lake to a min¬

imum of 1 m. in Helmet Lake.

Nielsen (1933) experimented with natural plankton and

found that the oxygen production was substantially the same at

0.2 m. and at 2 m. The compensation point was reached at a

depth of 7 m. in his experiments. This is not as deep as the

compensation points in Crystal and Trout lakes, but deeper than

those of Mud and Helmet lakes.

Kurasige (1932) studied the photosynthesis of natural plank¬

ton consisting chiefly of Melosira and Synedra; the experiment

was performed out of doors in the latter part of March, using a

40 1. aquarium. The diatoms produced 0.515 mg. of oxygen per

million cells per day at a temperature of 8-16° C-, when the

total radiation was about 500 cal/cm2. He found a linear rela-

190 Wisconsin Academy of Sciences , Arts, and Letters .

tion between oxygen production and the amount of solar radia¬

tion. The oxygen production decreased between 11:00 a.m. and

3:00 p.m., thus showing that the sunlight was too intense for

optimum photosynthesis during this period.

Summary

1. The photosynthesis of two species of algae at different

depths in four lakes was determined by measuring the quantity

of oxygen produced.

2. Maximum oxygen production was found at the surface on

dark, cloudy days, but at a certain distance below the surface

on bright, clear days.

3. The depth of the maximum production on clear days de¬

pended upon the transparency and color of the water.

4. The depth of the compensation point, where the quantity

of oxygen produced by photosynthesis just balanced that con¬

sumed in respiration, also varied with the transparency and color

of the water.

5. The amount of solar energy penetrating to the various

depths was determined.

6. The percentage of utilization of solar radiation by the

algae increased with increasing depth, reaching a maximum

where the total radiation varied from 1.2 to 8 cal/cm2 in a three

hour period.

Schomer & Juday — Photosynthesis of Algae

191

Literature

Birge E. A. and Juday C. 1929. Transmission of solar radiation by the waters of

inland lakes. Trans. Wis. Acad. Sci. Arts & Let. 24:509-580. Second report.

1930. Ibid. 25:285-335. Third report. 1931. Ibid. 26:383-425. Fourth

report. 1932. Ibid. 27:523-562.

Gaarder T. and Gran H. H. 1927. Production of plankton in Oslo Fjord. Cons.

Perm. Internat. Explo. de la Mer. Rapports et Procfcs-Verbaux des Reunions.

42:1-48.

Juday C. The depth distribution of some aquatic plants. Ecology 15: — .

Kurasige, H. 1932. Some experimental observations on the diurnal change of pH and

oxygen production by aquatic plants in relation to the solar radiation. Geophys.

Mag. 5:343-359.

Marshall, S. M. and Orr, A. P. 1928. The photosynthesis of diatom cultures in

the sea. Jour. Mar. Biol. Assoc. United Kingdom, N. S. 15:231-360.

Nielsen, E. S. 1933. Einleitende Untersuchungen iiber die Stoff -production des

Planktons. Med. Kom. Danmarks Fisk, og Havunders. Serie Plankton, Bd. II.

Reviewed in Nature 132:572.

Schomer, H. A. 1934. Photosynthesis of water plants at various depths in the

lakes of northeastern Wisconsin. Ecology 15:217-218.

192 Wisconsin Academy of Sciences, Arts, and Letters .

Table I

Several series of experiments on photosynthesis are given for 1933. Solar

energy is expressed in gram calories per square centimeter for the three hour period in

each case and the oxygen is indicated in milligrams per million cells for the three

hour interval.

Schromer & Juday — -Photosynthesis of Algae

193

(Table I Continued)

THE COURSE AND SIGNIFICANCE OF

SEXUAL DIFFERENTIATION

Charles E. Allen

In the most nearly primitive condition that we can now en¬

visage with certainty living matter existed in the form of cells,

isolated or temporarily in contact; many organisms still extant

are in this respect relatively primitive. Cells increased in num¬

ber by division — the simplest and doubtless the original method

of division being that of a single cell into two similar daughter

cells. In case the parent cell was differentiated into cytoplasm

and nucleus — as are cells of the great majority of present-day

types — its division was necessarily preceded by a division of the

nucleus.

Cell division was, therefore, so far as can now be determined,

the original method of reproduction. It remains the basic re¬

productive process in all living organisms. For a cell aggregate

or a multicellular organism consists of cells owing their origin

ultimately to division of the cells of the parent organism or

organisms, and so on indefinitely backward.

In contrast to the increase in number of cells by division, a

reverse process is possible— the union of two or more cells into

one. As illustrated, for example, in some flagellates and rhizo-

pods, the union of two cells without walls which have come in

contact is superficially very like the union of two drops of water

when they touch into one drop. Actually, of course, the union

of protoplasts must be a much more complex procedure than

this; but the similarity is striking and probably not without

significance. If the uniting cells are delimited by walls, as is

true of the anastomosing cells of some fungi, the additional

step is involved of a dissolution of the walls in the region of

contact.

The possibility of cell union, manifested in various widely

separated groups of organisms, offers a suggestion as to how a

new evolutionary departure may have occurred. This departure

involved the union not only of two cells but also of their nuclei.

Cells which thus united in pairs were gametes ; the single-nucle¬

ate cell formed by their union was a zygote. Whether or not the

195

196 Wisconsin Academy of Sciences , Arts, and Letters.

new step first occurred, as it were, by chance, a union of nuclei

following upon a casual union of cells, can not now be said. But

it is certain that, once brought about, gametic union opened the

way to most far-reaching consequences. Its central importance

lay in the facts that it brought two distinct and usually different

sets of hereditary substances into a new combination, and that

it necessitated sooner or later a reshuffling of those substances

into still other new and varied groupings. By these means novel

possibilities of hereditary variation were introduced, and new

raw material was provided for the evolutionary process.

However it first came about, and whether once or, as is most

probable, many times in different lines, the union of gametes

became an established habit — a habit fixed, it may be imagined,

in consequence of the advantages enjoyed by those organisms

which had developed the new possibility. It became and re¬

mained a definite part of the life cycle, not only of many rela¬

tively simple organisms, but of those lines which were to lead to

the more complex or “higher” groups of animals and plants.

True, gametic union has dropped out of the life cycles of indi¬

vidual species in various of these higher groups ; but such union

is still of nearly universal occurrence save in a very few lowly

assemblages.

Many one-celled and simple colonial plants still living retain

what must be considered the original type of gametic union, in

which the gametes are, to all appearance, substantially alike. In

some instances these like gametes are ordinary vegetative cells

that, under particular conditions, assume the gametic function;

in other cases they are smaller, having been produced by a divi¬

sion or a series of divisions of a vegetative cell.

But among simple plants are others, often closely related to

some of those just mentioned, which produce two sets of visibly

different gametes. The differentiation of two kinds of gametes

constituted another significant innovation. Like gametic union,

gametic differentiation became a habit fixed in the genetic con¬

stitution of the species concerned. We may not explain in causal

terms the occurrence of this type of differentiation; a possible

guess is that it expressed a tendency inherent in the constitution

of living matter which, once a certain degree of evolutionary

development was reached, found expression. But while the caus¬

ative explanation escapes us, it is easy to perceive the advantage

Allen — Sexual Differentiation

197

that a species derived from the differentiation of two classes of

gametes. This advantage followed from the fact that, gametic

union being an established habit and its periodic occurrence a

matter of vital import to the species, the gametes were burdened

with two distinct functions: one to come into contact, the

other to provide for the zygote the food needed by it for its own

activities, including division or “germination,” and by the cells

to be formed when it divides.

The ancestral lines of nlost animals and plants converge in

the flagellates-— organisms which during at least a considerable

part of their existence are single motile cells. Therefore, mo¬

tility was an original property of gametes, if and when gametic

union was established at the flagellate level of evolutionary de¬

velopment; or, if at a somewhat higher level, the production of

motile gametes involved that of cells representing the persist¬

ence of a not very distant ancestral condition. In general, there¬

fore, the coming together of gametes was provided for by their

motility — together, of course, with the development of the power

of response by movement toward the source of a stimulus eman¬

ating from another gamete of the same species. The provision

of food to be stored in and carried by the gametes also implied

no new development; merely some extension of a process of

food-manufacture already in possession and essential to the

existence of the species.

But, even at a low level of development, these two functions

necessary to the success of gametic union were measurably in

conflict. The storage of food within a gamete tended to an in¬

crease in size, rendering it less suited to active movement. The

difficulty was increased if, as in many algae and fungi, the zy¬

gote was to pass into an extended dormant period during which

its continued existence would be dependent upon stored food;

or if, as in the lines that led to “higher” plants and animals, the

germination of the zygote was to be followed by an embryonic

phase in the development of a new generation.

Even in many comparatively simple organisms, then, the in¬

consistency between two duties to be performed by the gametes

made itself felt. The inconsistency was avoided, as many diffi¬

culties in more familiar stages of development have been

avoided, by a division of labor. One gamete, assuming the func¬

tion of food-carriage, became larger, less motile, and ultimately

198 Wisconsin Academy of Sciences , Arts , and Letters .

quite non-motile. It is henceforth distinguished as the female

gamete. The other, the male gamete, remaining small or even

becoming smaller, carried only the food supply needed in its own

brief existence, and continued motile or even became capable of

greater activity.

It was said that gametic union probably became established

independently in different lines as a regularly recurrent process.

It is certain that gametic differentiation has occurred independ¬

ently in many diverse lines of animals and plants. If the illus¬

trations here cited of this differentiation are taken from the

plant kingdom, it is because many plants, particularly algae,

still survive which illustrate steps in the increasing differentia¬

tion of the gametes ; whereas among animals, even including

protozoa, the early chapters of this history are but sparsely rep¬

resented.

In selecting species illustrative of the sequence in question,

a caution necessary in dealing with most evolutionary series

must be kept in mind ; namely, that a species now living which

represents stage A is not likely to be directly ancestral to an¬

other contemporary species representing stage B . Rather, the

former species presents, with reference to a particular evolu¬

tionary process, a condition essentially similar to one through

which, it may reasonably be concluded, passed the ancestors of

the second species. The relationship between such living spe¬

cies, if any, is ordinarily collateral, not lineal.

With the progress of sexual differentiation the female and

male gametes came to differ both in structure (including size)

and in behavior. Only one of these differences may be apparent

at a very early stage in differentiation. Thus the gametes of

Ectocarpus, a widespread genus of brown seaweeds, are alike

in structure and size but different in behavior. In Giffordia,

however, a close relative, the gametes have come to differ in

size as well as in behavior.

Sex — the occurrence of sexual differentiation — thus arose as

an adaptation to needs that resulted from the adoption of the

habit of gametic union. Sex is not a primitive characteristic of

living matter. It constitutes an adaptation that occurred at pe¬

riods long subsequent to the first appearance of living matter

upon our planet. This adaptation was itself the beginning of a

long developmental history, destined to extend far beyond the

Allen — Sexual Differentiation

199

gametes in which it first appeared, and to affect in different

ways and in varying degrees the structures and functions of

the organisms producing the gametes.

The appearance of, and the progressive increase in, differ¬

ences between female and male gametes, may be considered the

first chapter of sexual differentiation; the female becoming

many times, often thousands of times, as large as the male and

quite passive; the male gamete, in most groups, remaining ex¬

tremely active* In this latter respect, however, some interest¬

ing divergences appear; as in the red algae, whose male gam¬

etes, while very small, are non-motile; their movement to the

neighborhood of the female gametes is effected by currents of

the water in which the plants are submerged.

By a second series of steps, there came about a differentia¬

tion of the organs in which the gametes are produced. Distinct

female and male organs are to be found in many algae; more

highly specialized structures of these classes appear in liver¬

worts, mosses, and ferns.

In the third phase of this history, differentiation extended to

structures other than the organs in which gametes were form¬

ed; particularly, as in mosses, to such structures as leaves in

the immediate neighborhood of the gamete-bearing organs, or

the branches on which these organs are borne. The extreme of

this type of differentiation is found in those species in which

female and male gametes are formed by separate organisms

(whether plant or animal). This latter degree of sexual sep¬

aration, also, has been brought about independently in different

lines; and it is notable that in various of the larger groups of

plants some species retain the power of producing both kinds of

gametes on the same plant, whereas in other species, often with¬

in the same genus, the possibilities of such production are more

or less sharply separated; in the latter species some individual

plants are male, others female. In certain cases the female and

male plants have become greatly different ; the male plant being

usually much smaller than the female, as well as more or less

divergent in structure. The climax in this direction is illus¬

trated by some species of Oedogonium among the green algae,

and by those mosses which have dwarf male plants. Here sex¬

ual differentiation, beginning with the gametes, has progressed

until it involves the whole organism.

200 Wisconsin Academy of Sciences , Arts , cmd Letters.

In a fourth series of steps a tendency appears for the gam¬

ete-bearing plant to take over in part the particular functions

first performed by the gametes. Those functions, as we have

seen, were to bring the gametes together, and to provide food

for the use of the zygote and for the launching of the new gen¬

eration which will come from the zygote. As this new genera¬

tion developed a longer and longer embryonic stage, its need of

an initial food supply increased. With this need was connected

another, also of increasing importance; that, namely, of shelter

for the embryo, originally enclosed only within the wall of the

zygote. Liverworts and mosses supply relatively simple illus¬

trations of the ways by which the gametes are in a measure re¬

lieved of their duties. The male gamete-bearing organs are so

constructed that by the dissolution of some of their cells the

male gametes are freed; and water-absorbing substances are

secreted whose swelling forces out the gametes. In some liver¬

worts, an explosive discharge throws the male gametes outward

to a considerable distance. On the other hand, the female organ

provides a pathway through which the male gamete may swim

in order to reach the female gamete, and a substance to whose

stimulating effect the male gamete responds by a directed move¬

ment. The female organ is partly imbedded in tissues of the

parent plant, fitted to provide nutrition. The female parent,

too, provides special sheltering structures for the embryo. All

these devices, assisting in the performance of duties originally

incumbent upon the gametes alone, constitute part of the story

of sexual differentiation.

Still further developments in the same direction appear in

seed plants. The pollen grain, a young male plant, is itself

transported by wind or insects to the neighborhood of the fe¬

male plant. The pollen tube, an outgrowth of the same much-

reduced male plant, provides a means by which the male gamete

may reach the female. So effective are these means of trans¬

port that the male gamete has lost most of its power of motility,

formerly its distinctive characteristic. The female plant, like¬

wise reduced, consists (in the gymnosperms) chiefly of tissue

destined to nourish the embryo.

But most remarkable among the sexual adaptations of seed

plants, constituting the fifth, and thus far the final, chapter of

the story, are those by means of which certain functions, in-

Allen — Sexual Differentiation

201

cumbent first upon the gametes, then taken over by the parents

— that is, the gamete-producing plants — have been in consider¬

able measure assumed by the grandparents. The paternal

grandparent assists, by the structure of its stamens, in the dis¬

tribution of the potential fathers — a distribution in which the

assistance of wind or of insects (occasionally of other agencies)

has been commandeered. The maternal grandparent provides a

place wherein the young father-to-be may land and may com¬

plete development; a canal through which the paternal pollen

tube may grow ; and structures which will surround the mother

and its enclosed embryo, supplying to the latter, through periods

of growth and dormancy, both nutriment and shelter.

As has been said, a substantially similar story could be told

of the course of sexual differentiation in animals, save for the

fact that the earlier phases of this process are but poorly repre¬

sented in surviving species; whereas the story can be recon¬

structed virtually in its entirety from the observation of plants

still living. It does not appear, however, that in any animal

species the grandparents have ever become so essential to the

production and care of the grandchildren as are grandparents

among the seed plants.

The story of sexual differentiation, as here outlined, like

most stories of biological import, has been learned backward.

First to be observed were sexual differences manifested by the

higher animals; later, not clearly recognized until the closing

years of the seventeenth century, corresponding differences in

the higher plants. Only a few years before had the male gam¬

etes of an animal first been seen. Not until the second half of

the nineteenth century was the process of gametic union under¬

stood in its major outlines; and much of what has been here

discussed proceeds from knowledge acquired but yesterday.

Such a reversed working out of a problem results inevitably in

the persistence of concepts based upon an incomplete knowledge

of conditions in more complex organisms, whereas ideally those

conditions should be explained in terms of concepts derived from

simpler forms. Most emphatically is the latter statement true

of any notion with an evolutionary bearing.

One outcome of the growth of knowledge in reverse has been

the notion of sex as something fundamentally characteristic of

living matter. Really, sex is neither fundamental nor primitive.

202 Wisconsin Academy of Sciences , Arts, and Letters.

It represents an adaptation to certain needs — or a means of

solution of certain problems — which appeared at a particular

and relatively advanced stage of evolutionary progress.

Another consequence of our having learned the story back¬

ward rather than forward is the still frequent and absurd ap¬

plication, particularly in textbooks, of the term “sexual repro-

duction,, to gametic union. For, although in a multicellular

organism the union of gametes is a step preparatory to repro¬

duction of that organism, gametic union itself is never repro¬

duction. It is the exact reverse. Yet more absurd is it to call

the union of like gametes “sexual reproduction.’' Since such a

union represents a step that preceded in evolution the appear¬

ance of sexual differentiation, it is no more “sexual” than it is

“reproduction.” It is easier for scientific, as well as for some

other classes of writers, to use old-established but inaccurate

phrases than it is to criticize and make precise their terminology

so that they may, within the limitations imposed by a very im¬

perfect language, say just what they mean.

WILD LIFE RESEARCH IN WISCONSIN

Aldo Leopold

University of Wisconsin

History

Wisconsin has always been a stronghold for ornithology. In

the 1840s, before the state was half settled, the two Kumliens

( 1 ) at Delavan, and Dr. P. R. Hoy at Racine, had started their

painstaking inventories of our avian species.

Then followed the era of “economic ornithology,” the inven¬

tory not of bird species but of bird foods. In 1873 F. H. King

(2) started exploring the economic relations of Wisconsin birds,

a field which was later developed nationally by Judd, Fisher,

Beal, McAtee, and other naturalists of the U. S. Biological Sur¬

vey. King's undeviating rigor in applying the economic yard¬

stick tells us much, not only about birds, but about the natural

philosophy of his generation. With all the circumspection of a

cautious radical expecting short shrift from his contemporaries,

King suggests “the preservation of animals to some extent de¬

trimental , i.e., to some extent at cross purposes with the sacred

economy of Homo sapiens. Apparently it was dangerous in those

days to assert flatly that a beautiful animal is his own justifica¬

tion.

King discovers, evidently with surprise, that an actual count

showed more birds per square mile near Ithaca, New York, than

in the more recently settled regions of Jefferson County, Wiscon¬

sin. He shared in, or at least had to deal with, the then uni¬

versal assumption that wild life must disappear with the ad¬

vance of settlement. This should be of special interest to the

modern game manager, whose life-work rests squarely on the

opposite premise.

At the turn of the century it became fashionable not only to

love birds, but publicly to avow the fact. The amateur move¬

ment, as expressed in Bird Clubs and Audubon Societies, was

strong in this state. Bird-lovers lacked both the botany and

the entomology necessary to pursue economic studies, and of

course also the dead specimens for study. Their energies turned

203

204 Wisconsin Academy of Sciences, Arts, and Letters .

to expanding the species inventory, to migration records, and

latterly to banding. It may not be generally known that one of

the pioneers in large-scale banding was Dr. L. J. Cole ( 3 ) of

Wisconsin. Desultory banding had, of course, been practiced

since the Middle Ages (4).

Wild life research in the modern sense began in this state

when Herbert L. Stoddard, an “incorrigible” young naturalist

from Sauk City, was retained by the Milwaukee Public Museum

as taxidermist and collector in 1921. Sometime during that ap¬

prenticeship he must have conceived what is today the founda¬

tional theorem of wild life management, namely:

A species can he decimated by throwing its environment out of bal¬

ance. Conversely, it can be restored by restoring the balanced assortment

of environmental features required for its welfare.

Stoddard first applied this theorem to his now famous Geor¬

gia quail investigation (5). The idea, however, must have been

born during his work here. It is significant that its birthplace

was in the mind of a man who had never seen the inside of a

High School. A whole new profession is now engaged in follow¬

ing down its ramifications.

Stoddard cast his intellectual pebble in Georgia in 1924. By

1928 the outermost ripples had travelled back to the state of his

nativity. In that year two projects, patterned after the Georgia

model, were started here. Alfred 0. Gross began the Wisconsin

prairie chicken investigation (6), under the auspices of the Re¬

search Bureau of the State Conservation Commission. Paul L.

Errington, under a grant to the University from the ammuni¬

tion industry, started the Wisconsin quail investigation (7).

The Quail Investigation

It is now three years since the completion of Errington’s

work, and perhaps not too early to attempt a summary of his

discoveries and an appraisal of their significance. This may

seem unbecoming in one who enjoyed daily association with his

work, but those who understand the degree to which he was

“his own man” may find it not improper.

Errington’s most valuable findings relate to predation and

its interrelations with food and cover. It has long been known

that fitness determines survival, but we have only dimly realized

that environment determines fitness. Errington worked out, for

Leopold-Wild Life Research

205

Wisconsin quail, the details of just how and why. He qualified

the usual supposition that species have certain inherent apti¬

tudes for capture or escape, by showing that those aptitudes are

operative only within certain narrow limits of circumstance.

Thus the Cooper’s hawk catches quail, but given a suitable vari¬

ety of food and cover, the covey promptly changes its habits and

becomes hawk-proof. On the other hand, if without alternative

foods, it is annihilated either by the hawk or by starvation.

A corollary of this is that predation falls heaviest on coveys

occupying marginal ranges. These may be wiped out while near¬

by coveys with good food and cover go unscathed. Predation is

never uniformly distributed among population units.

All predation has been considered as selective of the unfit.

Errington discovered wide variations in selectivity. Certain pre¬

dators catch quail by chance, i.e., non-selectively ; others catch

only misfits, and are highly selective. The predators of high

“economic” value, such as the Buteo hawks, are the most se¬

lective.

To get the data which yielded these conclusions, Errington

invented new technique for field studies, supplementing the

techniques previously devised by Stoddard. For example, by

tethering young hawks and owls and forcing them to regurgi¬

tate the food brought them by parents, he obtained their daily

menu without recourse to killing, and secured a continuity im¬

possible under older methods. By feeding captive raptors vari¬

ous animals and then collecting their pellets, he found the rate

of pellet formation and the degree to which the smaller bones

are digested, thus allowing more and sounder conclusions to be

drawn from the pellets of wild birds.

Stoddard and Errington jointly are probably responsible for

the discovery of the food sequence, a concept unknown to the

economic ornithologists with their composite cross-sections of

stomachs gathered at diverse times and places. The daily menu

of quail in winter was found to follow a sequence representing

a descending scale of palatability. Winter survival was found

to be a question of how low on the scale the last blizzard came.

That low-scale foods do not sustain weight and fitness was ex¬

perimentally verified.

Errington now promulgates, for Wisconsin quail, the general

theorem (8) that predation varies with the degree to which a

206 Wisconsin Academy of Sciences, Arts, and Letters .

population falls below or rises above carrying capacity. A range

when over-stocked loses heavily, apparently regardless of the

number or kind of predators or buffers. The same range when

under-stocked loses lightly, regardless of the number or kind of

predators or buffers. That is to say, quail populations on any

given range fluctuate about a norm representing carrying ca¬

pacity. It follows that enhanced carrying capacity, i.e., food

and cover improvements, is the only effective predator-control,

also that surplus over capacity had as well be used by hunters

as by natural enemies.

In general, Errington created a coherent, though necessarily

tentative, theory of predation which so far applies only to Wis¬

consin quail, but which may ultimately have far-flung conse¬

quences to ecological science. It is a timely contribution, be¬

cause predation continues to be the prime cause of dissension

between gunpowder and field glass hunters.

Research Sequence

Wild life management research projects are so far of several

types, which occur in a rather definite sequence. By projecting

this sequence a little into the future, we can gain a rough idea

of the volume and kind of work which lies ahead. For any one

species the sequence is :

1. Survey of the species, to find the most important chances for im¬

proving its environment and thus raising population levels.

2. Tests of the findings on actual range, to see if the higher levels

are obtained.

3. Ecological exploration of key plants.

4. Physiological and pathological studies.

5. Economic vehicles for encouraging management.

Species surveys have been made by Errington of Wisconsin

quail and by Gross and Schmidt of prairie chickens. Deer, wa¬

terfowl, shorebirds, fur animals, and cottontails remain virgin

fields as yet untouched. Michigan surveys of pheasants by Wight

(9) and of Hungarian partridge by Yeatter (9a) will probably

suffice for this state; likewise Minnesota's survey of ruffed

grouse by King.

Errington’s findings on quail are now being tested on the

Coon Valley Erosion Control Project by Holt. The findings of

Schmidt and Gross on prairie chickens are being tested on the

Leopold — Wild Life Research

207

central Wisconsin Game Area. Some less thorough tests of

Wight's, Yeatter's, and Errington's findings are being conduct¬

ed, in cooperation with the University, by volunteer farm

groups at Faville Grove in Jefferson County, Monroe Center in

Adams County, and Riley in Dane County.

Tests nearly always disclose deficient knowledge of some key

plant, but this type of research is as yet lacking in Wisconsin.

For example, we need to learn how, by ecological manipulation,

to control the supply of wild rice, wild celery, trefoil, ragweed,

and false climbing buckwheat. Stoddard induces a growth of

legumes in Georgia by plowing furrows in sod, or by controlled

burning. In Scotland the grouse crop — some 2 million birds

per year — rests squarely on the ecological manipulation of

heather (10).

Physiological and pathological studies have begun. A cycle

study was recently inaugurated by Grange (10a) and Wing

(10b) under a grant of funds to the University by a group of

sportsmen. Wisconsin also enjoys close cooperative relations

with Dr. R. G. Green's study of cyclic diseases at the University

of Minnesota. There is crying need for a study of vitamin and

mineral nutrition as affecting disease and reproduction in game

birds.

To accomplish conservation, game research must not stop

with its “layette" of biological facts. There remains the prob¬

lem of economic adjustments to encourage their use by land-

owners. Some current aspects of conservation economics have

been treated in a separate paper (if).

Opportunities

Wild life research offers a new field for service, not only to

the larger universities and agricultural colleges, but also to col¬

leges, agricultural high school teachers, county agents, forest

officers, game officials, and private naturalists throughout the

state. The first criterion of vitality in such a program is its

degree of dispersion among many investigators of diverse points

of view. There are so far ten times more challenging problems

than investigators to attack them.

It may also be proper to ask why any scientific institution

in this state, confronted by so many unanswered questions of

vital import to our future, should have to seek outlets for its

208 Wisconsin Academy of Sciences, Arts, and Letters.

brains and energy in expeditions to foreign lands. The era of

geographical exploration of the earth is about over, but the era

of ecological exploration of our own dooryards has just begun.

Wild life research is one of many virgin fields of inquiry in

which any persistent investigator may contribute not only to

science, but to the permanence of the organic resources on which

civilization is dependent.

References

1. Kumlien, L. and Hollister, N. “The Birds of Wisconsin.” Wis. Nat.

Hist. Soc., 1903.

2. King, F. H. ‘'‘Economic Relations of Wisconsin Birds.” Geology of

Wisconsin, Vol. 1, 1873-1879, pp. 441-610.

3. Cole, L. J. “The Early History of Bird Banding in America.” Wil¬

son Bulletin, June, 1922, pp. 108-115.

4. Haskins, C. E. “Studies in the History of Mediaeval Science.” Har¬

vard Press, Cambridge, 1927.

5. Stoddard, Herbert L. “The Bobwhite Quail, Its Habits, Preservation,

and Increase.” Chas. Scribner’s Sons, New York, 1931.

6. Gross, A. O. “Progress Report of the Wisconsin Prairie Chicken In¬

vestigation.” Wisconsin Conservation Commission, 1930.

7. Errington, Paul L. Numerous papers in American Game, Wilson Bul¬

letin, The Condor, Trans. Wis. Acad. Sciences, 1928-1933.

8. Errington, Paul L. “Vulnerability of Bob-white Populations to Pre¬

dation.” Ecology, Vol. XV, No. 2, April, 1934, pp. 110-127.

9. Wight, H. M. “Suggestions for Pheasant Management in Southern

Michigan.” Michigan Department of Conservation, August, 1933.

9a. Yeatter, R. E. “The Hungarian Partridge in the Great Lakes Re¬

gion.” Bulletin No. 5, Michigan School of Forestry and Con¬

servation, December, 1934.

10. Leopold, Aldo, and Ball, J. N. “British and American Grouse Man¬

agement.” American Game, July-August, September-October,

1931.

10a. Grange, Wallace. “A Study of the Wisconsin Game Cycle from Low

to High and the Beginning of the Second Low, 1928 to 1934.”

MSS. report, 1934.

10b. Wing, L. W. “Cycles of Migration.” Wilson Bulletin, Vol. XLVI,

September, 1934, pp. 150-156.

11. Leopold, Aldo. “Conservation Economics.” Jour. Forestry, Vol.

XXXII, No. 5, May, 1934.

MAPLE SUGAR: A BIBLIOGRAPHY OF EARLY RECORDS

H. A. SCHUETTE AND SYBIL C. SCHUETTE

University of Wisconsin , Madison, and Kellogg Public Library ,

Green Bay

This bibliography of references to maple sap and its prod¬

ucts is presented as part of a program of research on the history

of foods. In this instance interest centers around two typically

American foods obtainable from the “wounded maple'5 and in

the past referred to as Canada, American or Indian sugar and

American “melasses” or “syrrup” of maple, respectively. Its

first item, an excerpt from the journal of a Jesuit missionary,

marks it as beginning with a period (1634) which is coincident

with the coming of the first white man to Wisconsin.

It is not a bibliography in the sense that it is an exhaustive

compilation of all the titles or relevant comments and observa¬

tions which have appeared on this subject during the two-and-

a-quarter century span which opened with a significant date in

the history of the Northwest Territory and closed, approxi¬

mately, during Civil War days. In this respect it falls short of

that measure of completeness. Except for a few items, it is the

result of a search of a selected group of books of travel and

natural history, diaries, journals, narratives and miscellaneous

communications in periodicals, many of them long out of print,

in the collection of the State Historical Society of Wisconsin.

It is deemed to be sufficiently comprehensive to supply the stu¬

dent of this field of history with source material which is not

only instructive and interesting, but, at times, even amusing.

1. Le Juene, Paul 1634

Jesuit Relations and Allied Documents. Travels and

Explorations of the Jesuit Missionaries in New France.

1610-1791. R. G. Thwaites, Editor. The Burrows Broth¬

ers Company, Cleveland, 1901, Vol. VI, p. 273.

(The sugar maple a source of food for the Indian)

“When they are pressed by famine, they eat the shav¬

ings or bark of a certain tree, which they call Michtan ,

which they split in the Spring to get from it a juice, sweet

209

210 Wisconsin Academy of Sciences, Arts, and Letters.

as honey or as sugar; I have been told of this by several,

but they do not enjoy much of it, so scanty is the flow.”

2. Denys, Nicolas 1672

Histoire naturelle des Peuples, des Animaux, des Arbres

& Plantes de PAmerique Septentrionale, & de ses divers

Climats. Paris, 1672, Vol. II. Chap. XX. W. F. Ganong,

The Description and Natural History of the Coasts of

North America (Acadia). Toronto, 1908, p. 380-1.

(Description of the tapping of maple trees whose

sap is a favorite drink of both Indians and French)

“That tree has sap different from that of all others.

There is made from it a beverage very pleasing to drink,

of the colour of Spanish wine but not so good. It has a

sweetness which renders it of very good taste; it does not

inconvenience the stomach. It passes as promptly as the

waters of Pouque. I believe that it would be good for those

who have the stone. To obtain it in the spring and autumn,

when the tree is in sap, a gash is made about half a foot

deep, a little hollowed in the middle to receive the water.

This gash has a height of about a foot, and almost the same

breadth. Below the gash, five or six inches, there is made

a hole with a drill or gimlet which penetrates to the middle

of the gash where the water collects. There is inserted a

quill, or two end to end if one is not long enough, of which

the lower extremity leads to some vessel to receive the wa¬

ter. In two or three hours it will yield three or fours pots

of the liquid. This is the drink of the Indians, and even of

the French, who are fond of it.”

3. Nouvel, Henri 1673

Jesuit Relations. Vol. LVI, p. 101.

(Sap is called “maple water”)

“About the same time, I made various excursions on the

ice in quest of stray sheep, — finding five children, to Bap¬

tize, and a sick young man, for whose salvation Providence

was more watchful than I. For, having inadvertently bap¬

tized him, not with natural water, but with a certain liquor

that runs from the trees toward the end of Winter, and

which is known as “Maplewater”, which I took for natural

water, I discovered my mistake when, wishing to give this

Schuette & Schuette— Maple Sugar Bibliography 211

patient a dose of Theriae, I asked for some maple-water, —

which, being naturally sweet, is more suitable for such a

purpose. I was given some of the same liquor that I had

used in baptizing him, and was thus obliged to repair that

error,— happily a little before his death.”

4. Dalmas, Antoine 1674

Ibid., Yol. LVIII, p. 121.

(Red maples)

This is a report to the superior Dablon, of an expedition

of observation around Isle Jesus, near Montreal made in

September, 1674.

“We visited the country; the land is very stony, but has

many walnut-trees, beeches, aspens, and red maples, which

are numerous along these Shores.”

5. Nouvel, Henri 1675

Ibid., Vol. LX, p. 217.

(Comment on lofty oaks and maples)

Pertinent to the region near Lake Erie.

6. Thornton, _ 1684

Phil Mag., 1, 322-3 (1798).

Of an Attempt to make the Maple Sugar above an hun¬

dred Years ago. Dr. Robinson to Mr. Ray.

“It appears, by the following correspondence between

Dr. Robinson and Mr. Ray, that the property of the Ameri¬

can maple of yielding a saccharin juice was known above

a century ago, and that attempts were even made to pro¬

duce sugar from it:

London, March 10, 1684.

“Dear Sir,

“I have enclosed you some sugar of the first boiling got

from the juice of the wounded maple: Mr. Ashton, Secre¬

tary to The Royal Society, presented it to me. Twas sent

from Canada, where the natives prepare it from the said

juice; eight pints yielding commonly a pound of sugar. The

Indians have practiced it time out of mind ; the French be¬

gin to refine it ; and to turn it to much advantage. If you

have any of these trees by you, could you not make the trial

proceeding as with the sugar cane?

212 Wisconsin Academy of Sciences, Arts, and Letters.

Answer to Dr. Robinson:

Black Notley, April 1, 1684.

“Yours of the 10th instant I received, and therein an

enclosed specimen of the Canada sugar, a thing to me

strange and before unheard of. It were well worth the

experiment you mention. I therefore engaged a friend and

neighbour of mine, an ingenious apothecary, whom I em¬

ployed yesterday to boil the juice of the greater maple, a

tree which grows freely half a mile off from my residence.

Having made an extract, he found a whitish substance, like

to brown sugar, and tasting very sweet, immersed in a sub¬

stance of the color and consistency of molasses. Upon cur¬

ing, I have no doubt it will make perfect sugar. When it

is cured, I will give you a farther account of it.”

7. _ 1685

Phil. Trans., 15, 988.

( An account of a sort of Sugar made of the

Juice of the Maple, In Canada)

“The Savages of Canada, in the time that the Sap rises,

in the Maple, make an Incision in the Tree, by which it runs

out ; and after they have evaporated 8 pounds of the liquor,

there remains one pound as sweet, and as much Sugar, as

that which is got out of the Canes ; Part of the same Sugar,

is sent to be refined at Roven.

“The Savages have practiced this Art, longer than any

now living among them can remember.

“There is made with this Sugar a very good Syrup of

Maiden Hair, and other Capillary Plants, which is used in

France .”

8. Joutel, Henri 1688

Journal historique du dernier Voyage que feu M. de la

Sale fit dans la Golfe de Mexique pour trouver Tembouch-

ure, & le cours de la Riviere de Missicipi. Paris, 1713,

p. 352; in translation, A Journal of the Last Voyage per¬

form'd by Monsr. de la Sale, to the Gulph of Mexico to find

out the Mouth of the Missisipi River, London, 1714, p. 179.

See also Pierre Margry, Decouvertes et Etablissements des

Frangais dans l'ouest et dans le sud du l'Amerique Septen-

trionale (1614-1754). Paris, 1878, Pt. Ill, p. 510.

Schuette & Schuette — Maple Sugar Bibliography 213

(Sweet water from a tree)

‘‘The bad Weather oblig'd us to stay in that Place, till

April. That Time of Rest was advantageous for the Heal¬

ing my Foot ; and there being but very little Game in that

Place, we had Nothing but our Meal or Indian Wheat to

feed on; yet we discover'd a Kind of Manna , which was a

great help to us. It was a Sort of Trees, resembling our

Maple, in which we made Incisions, whence flow'd a sweet

Liquor, and in it we boil'd our Indian wheat which made it

delicious, sweet and of a very agreeable Relish.

“There being no Sugar-Canes in that Country, those

Trees supply'd that Liquor, which being boil'd up and evap¬

orated, turn'd into a Kind of Sugar somewhat brownish,

but very good."

9. LeClerq, Chrestien 1691

Nouvelle relation de la Gaspesie, qui contient les Moeurs

& la Religion des Sauvages Gaspesiens Porte-Croix, adora-

teurs du Soleil, & d'autres Peuples de 1'Amerique Septen-

trionale, dite la Canada. Paris. 1691, p. 124; W. F. Ga-

nong, New Relation of Gaspesia with the Customs and

religion of the Gaspesian Indians. Toronto, 1910, p. 122-3.

(Water of the maple)

“As to the water of the maple, which is the sap of that

same tree, it is equally delicious to French and Indians, who

take their fill of it in spring. It is true also that it is very

pleasing and abundant in Gaspesia, for through a very little

opening which is made with an axe in a maple, ten to a

dozen half-gallons may run out. A thing which has seemed

to me very remarkable in the maple water is this, that if,

by virtue of boiling, it is reduced to a third, it becomes a

real syrup, which hardens to something like sugar, and

takes on a reddish colour. It is formed into little loaves

which are sent to France as a curiosity, and which in ac

tual use serve very often as a substitute for French sugar.

I have several times mixed it with brandy, cloves & cinna¬

mon, and this makes a kind of very agreeable rossolis. The

observation is worthy of note that there must be snow at

foot of the tree in order that it shall let its sweet water

run; and it refuses to yield this liquid when the snow ap¬

pears no more upon the ground."

214 Wisconsin Academy of Sciences , Arts, and Letters.

10. de Lahontan, Louis baron 1703

New Voyages to North- America. London, 1708, Vol.

I, (a) p. 106, (b) p. 249; Vol. II, (c) p. 15.

a. (Maple syrrup at Green Bay)

“For Drink they gave me a very pleasant Liquor, which

was nothing but a Syrrup of Maple beat up with water;

if

b. (The mapple-tree)

“It yields a Sap, which has a much pleasanter taste than

the best Limonade or Cherry-water, and makes the whol-

somest drink in the World. This Liquor is drawn by cutting

the Tree two Inches deep in the Wood, the cut being run

sloping to length of ten or twelve Inches. . . Of this Sap

they make Sugar and Syrup, which is so valuable, that there

can’t be a better remedy for fortifying the Stomach. ’Tis

but few of the Inhabitants that have the patience to make

Mapple-Water, for as common and used things are always

slighted, so there’s scarce any body but Children that give

themselves the trouble of gashing these Trees. . . .”

c. (Mapletree-Water)

“I remember one Day in a Village of the Outaouas at

Missilimakinac a Slave brought into the Cottage where I

was, a sort of Vessel made of a thick piece of soft wood,

which he had borrowed on purpose, in which he pretended

to preserve Mapletree- Water.”

11. Dudley, Paul 1720

Phil . Trans., 31, 27-8.

An Account of the Method of making Sugar from the

Juice of the Maple Tree in New England

Directions are given for making maple sugar of the

juice of the upland maple that is maple trees that grow

upon the highlands.

“Some put in a little Beef Sewet, as big as a Walnut,

when they take it off the Fire, to make it turn the better

to Sugar, and to prevent its candying, but it will do with¬

out . our Physicians look upon it not only to be as

good for common use as the West India sugar, but to exceed

all other for its Medicinal Virtue.”

Schuette & Schuette — Maple Sugar Bibliography 215

12. Charlevoix, P. F. X., de 1721

Journal d’un Voyage fait par ordre du Roi dans l’Amer-

ique Septentrionale. Paris, 1744, Vol. Ill, p. 121; Anon.,

Journal of a voyage to North- America, London, 1761, Vol.

I, p. 191-4; Louise Phelps Kellogg edition, Chicago, 1923,

Vol. I, 176-9.

(Juice of the Maple)

“I was regaled here with the juice of the maple; this is

the season of its flowing. It is extremely delicious, has a

most pleasing coolness, and is exceedingly wholesome; the

manner of its extracting it is very simple.”

13. Rasies, Sebastien 1722

Jesuit Relations, Vol. LXVII. p. 93.

Lettre du Pere Sebastien Rasies, Missionnaire de la Cam-

pagnie de Jesus dans la nouvelle France, a M. son neveu

Father Rasies is among the Abnakis Indians in Lower

Canada and writes on October 15 from Nanrantsouak as

follows :

“. . . . my only nourishment is pounded Indian corn, of

which I make every day a sort of broth; that I cook in

water. The only improvement that I can supply for it is,

to mix with it a little sugar, to relieve its insipidity. There

is no lack of sugar in these forests. In the spring the maple-

trees contain a fluid resembling that which the canes of the

islands contain. The women busy themselves in receiving

it into vessels of bark, when it trickles from these trees;

they boil it, and obtain from it a fairly good sugar. The

first which is obtained is always the best.”

14. Beverley, Robert 1722

The History of Virginia. London, 2 ed. 1722, p. 118.

(The natural Product , and Conveniences of Virginia)

“The Honey and Sugar-Trees are likewise spontaneous,

near the Heads of the Rivers. The Honey-Tree bears a

thick swelling Pod, full of Honey, appearing at a Distance

like the bending Pod of a Bean or Pea; it is very like the

Carob Tree in the Herbals. The Sugar-Tree yields a kind

of Sap or Juice, which by boiling is made into Sugar. This

juice is drawn out, by wounding the Trunk of the Tree, and

placing a Receiver under the Wound, It is said, that the

216 Wisconsin Academy of Sciences , Arts, and Letters .

Indians made on Pound of Sugar, out of eight Pounds of the

Liquor. Some of this Sugar I examined very carefully. It

was bright and moist, with a large full Grain; the Sweet¬

ness of it being like that of good Muscovada.

‘‘Though this Discovery has not been made by the Eng¬

lish above 28 or 30 Years; yet it has been known among

the Indians before the English settled there. It was found

out by the English after this Manner. The soldiers which

were kept on the Land Frontiers, to clear them of the In¬

dians, taking their Range through a Piece of low Ground,

about forty Miles above the then inhabited Parts of Patoiv-

meck River, and resting themselves in the Woods of those

low Grounds, observed an inspissate Juice, like Molasses,

distilling from the Tree. The Heat of the Sun had candied

some of this Juice, which gave the Men a Curiosity to taste

it. They found it sweet, and by this Process of Nature,

learn’d to improve it into Sugar. Biut the Christian Inhabi¬

tants are now settled where many of these Trees grow, but

it hath not yet been tried, whether for Quantity, or Quality

it may be worth while to cultivate this Discovery.

“Thus the Canada Indians made Sugar of the Sap of a

Tree. And Peter Martyr mentions a Tree that yields the

like Sap, but without any Description. The Eleomeli of the

Ancients, a sweet Juice like Honey, is said to be got by

wounding the Olive Tree: and the East-Indians extract a

Sort of Sugar, they call Jagra, from the Juice or potable

Liquor, that flows from the Coco-Tree: The whole Process

of Boiling, Graining and Refining of which, is accurately

set down by the Authors of Hortus Malabaricus.”

15. Lafitau, Jos. F. 1724

Moeurs des sauvages Ameriquains, comparees aux

meours des premiers temps. Paris, 1724, Vol. II, p. 154.

(Maple syrup)

“Au mois de Mars, lorsque le Soleil a pris un peu de

force, & que les Arbres commencent a entrer en seve, elles

font des incisions transversales avec la hache sur le tronc

de ces arbres, d’ou il coule en abundance une eau, qu’elles

regoivent dans de grands vaisseaux d’ecorce; elles sont en-

suite bouillir cette eau sur le feu, qui en consume tout le

phlegme, & qui epaissit le reste en consistence de syrop, ou

Schuette & Schuette — Maple Sugar Bibliography 217

meme de pain de sucre, selon le degre & la quantite de cha-

leur qu’ils veulent lui donner. II n’y a point a cela d’autre

mystere. Ce sucre est tres pectoral, admirable pour les

medicamens ; mais quoqu’il soit plus sain que celui des Can¬

nes, il n’en a point Fagrement, ni la delicatesse, & a presque

tou jours un petit gout de brule, Les Francois le travaillent

mieux que les Sauvagesses de qui ils ont appris a le faire;

mais ils n’ont pu encore venir a bout de la blanchir, & de la

raffiner.”

16. Lemery, L. 1745

A Treatise of All Sorts of Foods, Both Animal and Veg¬

etable : also of Drinkables. Translated by D. Hay, London,

3 ed. 1745, p. 351.

(The sap of the maple)

“The Body, Branches, and Root of the Maple, yields a

sweet and pleasant Sap ; this Liquor, Mr. Ray says, is more

abounding in cold and rainy Weather, than in any other,

While the Birch, on the contrary, yield more in hot and dry

Weather.”

17. Kalm, Peter 1748

Travels into North America. J. R. Forster trans. Lon¬

don, 1772, Vol. I, 2 ed., p. 132.

(Treacle and Sugar)

“When the tree is felled early in spring, a sweet juice

runs out of it, like that which runs out of our birches. This

juice they do not make any use of here; but in Canada they

make both treacle and sugar of it.”

18. _ _ 1765

Gentlemen's Mag., 35, 439.

(An American discovery)

“The Americans have discovered a method of making

sugar from a liquor procured by boring the maple tree.

They say that more than 30 gallons have been procured

from one tree, which being manufactured after the manner

of the syrup proceeding from the sugar canes, produces a

sugar equal in goodness to that of Jamaica; and that the

molasses extracted from the pressure of the liquor, is very

little inferior to our West India molasses”.

238 Wisconsin Academy of Sciences, Arts, and Letters .

19. Bossu, N. 1771

Travels through that Part of North America formerly

called Louisiana. Translated from the French by J. R. Lon¬

don, 1771, Vol. I, 188-9.

(Sagamite sweetened with syrwp of the maple tree)

° ‘After the first ceremonies were over, they brought me

a calebash full of the vegetable juice of the maple tree. The

Indians extract it in January, make a hole at the bottom of

it, and apply a little tube to that. At the first thaw, they

get a little barrel full of this juice, which they boil to a

syrup; and being boiled over again, it changes into a red¬

dish sugar, looking like Calabrian manna ; the apothecaries

justly prefer it to the sugar which is made of sugar canes.

The French who are settled at the Illinois have learnt from

the Indians to make this syrup, which is an exceeding good

remedy for colds, and rheumatisms . . . they likewise

brought a dish of boiled gruel, of maize flour, called Saga¬

mite, sweetened with syrup of the maple tree ; it is an Indi¬

an dish which is tolerably good and refreshing.”

20. Adair, James 1775

The History of the American Indians. London, 1775.

p. 416.

(Indians make sugar.)

“Several of the Indians produce sugar out of the sweet

maple-tree, by making an incision, draining the juice, and

boiling it to a proper consistence.”

21. Carver, Jonathan 1778

Travels through the Interior Parts of North- America in

the Years 1766, 1767, and 1768. London, 1778, (a) p. 262,

(b) p. 282, (c) p. 496.

a. (Sugar not a sweetening agent)

“And even when they consume the sugar which they

have extracted from the maple tree, they use it not to ren¬

der some other food palatable, but generally eat it by itself.”

b . (Present of sugar)

“In the morning before I continued my route, several of

their wives brought me a present of some sugar, for whom

I found a few ribands.”

Schuette & Schuette — Maple Sugar Bibliography 219

c. (Two sorts of maple trees)

“Of this tree there are two sorts, the hard and the soft,

both of which yield a luscious juice, from which the Indians

by boiling make very good sugar. The sap of the former is

much richer and sweeter than the latter, but the soft pro¬

duces the greater quantity. . .

22. Bliss, Eugene F. 1782

Diary of David Zeisberger, a Moravian Missionary

among the Indians of Ohio. Trans, and ed. Cincinnati,

1885, Vol. I, p. 63, 66, 137, 186, 224, 324; Vol. II, p. 95,

305, 311, 347, 384.

(Sugar making in Ohio)

Sugar making was practiced by the Indians of Ohio in

the period 1782-1797.

23. Belknap, Jeremy 1784

Coll. Mass. Hist. Soc., [5] 3, 181 (1877).

(White Mountain Tour)

“Great quantities of maple sugar are made here. Mr.

W. has a set of vats to contain the sap, and a boiling house.

They commonly make enough for a year’s store. _ Our

bill of fare this day was ham, tongues, dried beef, trouts,

and a sauce composed of raspberries, cream, and maple

sugar.”

24. Hollingsworth, S. 1786

An account of the Present State of Nova Scotia. Edin¬

burgh, 1786, p. 21-2.

(Natural Productions)

— none is more useful to the inhabitants, than a

species of maple, distinguished by the name of the sugar

tree, as affording a considerable quantity of that ingred¬

ient; the juice flows fast into a vessel placed below

to receive it, and decreases in quantity as the sun declines

toward evening.

“The sugar, when cold, is of a reddish brown colour,

somewhat transparent, and very pleasant to the taste. It

can only, however, be considered as of use to the inhabi¬

tants within in the province; and they have not failed to

ascribe to it several virtues, either real or imaginary, as a

medicine.”

220

Wisconsin Academy of Sciences, Arts, and Letters .

25. _ 1788

Amer. Museum TJniv. Mag., 4, 349-50.

(Advantages of the culture of the sugar maple-tree)

Directions are given for making maple sugar, maple

molasses, maple beer, maple wine and maple vinegar.

26. Schopf, Johann David 1788

Reise durch einige der mittlern und sudlichen verein-

igten nordamerikanischen Staaten, nach Ost-Florida und

den Bahama-Inseln unternommen in den Jahren 1783 und

1784. Erlangen, 1788, Vol. I, p. 417 ; Alfred J. Morrison,

Travels in the Confederation (1783-1784). Philadelphia,

1911, Vol. I, 271-3.

(Maple sugar is cheaper than ordinary sugar.)

‘‘The sugar-maple is largely used by the people of these

parts, because the carriage makes the customary sugar too

dear for them. ... It is brown, to be sure, and somewhat

dirty and viscous, but by repeated refinings can be made

good and agreeable.,,

27. Loskiel, George Heinrich 1789

Geschichte der Mission der evangelischen Briider unter

den Indianern in Nordamerika. Barby and Leipzig, 1789,

p. 92-4 ; C. I. LaTrobe, History of the Mission of the United

Brethren. London, 1794, Pt. I, p. 72-3.

(Maple tree is much esteemed.)

“. . . near the Muskingum, sugar is boiled both in spring,

autumn, and winter, in case of need. . . . This is used by

the Indians either to sweeten their victuals, or in the place

of bread: and it is thought more wholesome, and sweeter

than our common brown sugar.”

28. _ 1789

Amer. Museum Univ. Mag., 6, 98-101.

Observations on manufacturing sugar

from the sap of the maple tree

Directions for making include use of wooden vessels that

“will not give the liquor a bad taste”. Too small a grain is

due to (1) makers have not used “lime or lye, or anything

else, to make it granulate; (2) sugar boiled too much.

Author suggests “a heaped spoonful of slacked lime — for

about six gallons of sap”. If quantity of lime too small.

Schuette & Schuette — Maple Sugar Bibliography 221

“the sugar will not be sufficiently grained; if too much, it

will give the sugar a reddish cast.”

29. _ 1790

Ibid., 7, 803-4

On maple sugar

“The manufactory of maple sugar opens a wide pros¬

pect of wealth to the united states. . . . Hence 250,336

acres of maple land will be sufficient to supply the whole

united states.”

30. A Society of Gentlemen 1790

Univ. Asylum Columbian Mag., 5, 106-9, 153-6; Nova

Scotia Mag., 3, 249-54; Annual Register 1791, 33 (Useful

Projects), 93.

Remarks on the Manufacturing of Maple Sugar

“A communication of such observations and directions

on manufacturing the Maple-Sugar as will be most useful

to those who, from situation, interest or patriotism, may be

induced to engage in and carry on this business.”

The States of New York and Pennsylvania have “a suf¬

ficient number of this kind of tree ... to supply the whole

of the United States, with this article.”

31. 1790

Ibid., 133.

Maple-sugar. Extract of a letter from Mr.

William Cooper, at Cooper' s-T own, Pennsylvania

“Those who think it more profit to clear them off the

ground, to make way for wheat or pasture, will please to

attend to the following receipt, taken from a farmer who

had saved four acres, exposed to the North-West, and then

recollect what employment is more profitable.

‘Received, Coopers' town, April 30, 1790, of William

Cooper, sixteen pounds, for six hundred and forty pounds

of sugar, at six pence per pound; made every pound with

my own hands, without any assistance, in less than four

weeks ; besides attending to the other business of my farm,

as providing firewood, taking care of cattle, etc.

John Nicholls

‘Witness,

‘Richard R. Smith'

222

Wisconsin Academy of Sciences, Arts, and Letters.

32. _ 1790

Ibid., 5, 203.

Shipment of Maple-Sugar

. . . “It has moreover other things in its favour, to rec¬

ommend it in preference to the sugar which is imported

from the West-India Islands. It is made by the hands of

freemen, and at a season of the year when not a single

insect exists to mix with and pollute it ; whereas the West-

India sugar is the product of the unwilling labour of negro

slaves, and made in a climate and in a season of the year,

in which insects of all kinds abound, all of whom feed upon

and mix with the sugar, so that the best India sugar may

be looked upon as a composition consisting of the juice of

the cane — and of the juices or excretions of ants, pismires,

cockroaches, borers, fleas, mosquitoes, spiders, bugs, grass¬

hoppers, flies, lizards; and twenty other West India in¬

sects. To these ingredients is added, the sweat of the ne¬

groes, and when they are angry, nobody knows what else.”

33. Lincklaen, John 1791

Travels in the Years 1791 and 1792 in Pennsylvania,

Mew York and Vermont, Journals of John Lincklaen, Agent

of the Holland Land Company. Helen Lincklaen Fairchild.

Mew York and London, 1897, p. 88-9.

(Vermont Journal — September 1791)

“There is in the whole State a considerable number of

Mapple Trees, but the people do not seem to be persuaded

of advantages they might gain from this tree. — Finally

the chief reason for not making sugar is that they have no

home market, and that the price of transportation by land

is too dear _ 77

34. Belknap, Jeremy 1792

The History of M ew-Hampshire. Boston, 1792, Voi. Ill,

2 ed., p. 113-6.

Forest-trees and other Vegetable productions

Descriptive of manufacture of maple sugar.

35. lmlay, Gilbert 1792

A Topographical Description of the Western Territory

of Morth America. London, 1792, p. 117-8.

Schuette & Schuette — Maple Sugar Bibliography 223

(Maple tree is productive of the finest sugars

under care and management.)

“The perfection to which we have brought our sugars

has induced many people in the upper parts of the States

of New York and .Pennsylvania to make a business of it

during the season of the juice running; and considerable

quantities have been sent to the markets of Philadelphia

and York, not inferior to the best clayed, French, and Span¬

ish sugars.77

36. Kush, .Benjamin 1793

Trans. Amer. Phil. Soc., 3, (a) 69, (b) 74.

An account of the Sugar Maple-tree of the United

States, and of the methods of obtaining Sugar from it, to¬

gether with observations upon the advantages both public

and private of this Sugar.

a. (Concentration of sap by freezing)

By freezing the sap, “one-half of a given quantity of

sap reduced in this way, is better than one third of the

same quantity reduced by boiling.77

b. (Maple sugar and corn mixture)

“They mix a certain quantity of maple sugar, with an

equal quantity of Indian corn, dried and powdered, in its

milky state. This mixture is packed in little baskets, which

are frequently wetted in travelling, without injuring the

sugar. A few spoonfulls of it mixed with half a pint of

spring water, afford them a pleasant and strengthening

meal77.

37. Wansey, Henry 1794

The Journal of an Excursion to the United States of

of North America in the Summer of 1794. Salisbury, 1796,

p. 63.

(Journey from Boston to New York)

“After passing Middleton, I saw the first maple sugar

tree; __ _ many afterwards that had been tapped. There

are many other kinds of maple trees ; the black, the white,

and the red do not produce the saccharine liquor77.

38. Graham, J. A. 1797

A Descriptive Sketch of the Present State of Vermont.

London, 1797, (a) p. 57, (b) 156-7.

224 Wisconsin Academy of Sciences, Arts, and Letters.

a. (Maple sugar at Sandgate)

“The making of sugar from the sap of the maple tree,

and of pot and pearl ashes, has afforded them great assist¬

ance, . . .

b. (Indian method of concentrating sap)

“The method pursued by the Aborigines in making this

article was as follows: Large troughs were made of the

Fine Tree, sufficient to contain a thousand gallons or up¬

wards; the young Indians collected the sap into these

troughs, the women in the mean time (for the men con¬

sider every thing but war and hunting as beneath their

dignity) made large fires for heating the stones necessary

for the process ; when these were fit for their purpose, they

plunged them into the sap in the troughs, and continued

the operation till they had boiled the sugar down to the

consistence they wished.77

39. _ 1797

Amer. Univ. Mag., 2, 221-4.

An Account of the Sugar-Maple Tree

“There are three modes of reducing the sap to sugar:

by evaporation, by freezing, and by boiling; of which the

latter is most general, as being the most expeditious .

It affords a most agreeable melasses, and an excellent vin¬

egar.77

40. Allen, Ira 1798

Natural and Political History of the State of Vermont.

London, 1798, (a) p. 9, (b) p. 277 ; Coll. Vt. Hist. Soc., 1,

(a) 335, (b) 484 (1870).

a. Sugar Maple

“. . . other species of useful timber, amongst which is

the sugar maple, from which the farmers often make more

sugar than serves for the usual consumption of their fami¬

lies, by the use of their kitchen utensils ; . . .7\

b. Maple sugar much used

“Maple sugar forms a great article of domestic con¬

sumption, the material is plenty, the preparation easy, the

taste agreeable, it seldom cloys the stomach, it is an excel¬

lent antiscorbutic, and so innocent, that it may be taken in

almost any quantity by infants.77

Schuette & Schuette — Maple Sugar Bibliography 225

41. Kush, Benjamin 1798

Phil Mag., 1, 182-91.

An Account of the Sugar Maple of the United States

Descriptive, with a plea for use of this sugar in place

of that made by slave labor.

42. Drake, Samuel G. 1799

Tragedies of the Wilderness; or True and Authentic

Narratives of Captives. Boston, 1841, (a) p. 197, (b) p.

215.

(Sugar tubs of elm bark)

“In this month we began to make sugar. As some of

the elm bark will strip at this season, the squaws, after

finding a tree that would do, cut it down, and with a

crooked stick, broad and sharp at the end, took the bark off

the tree, and of this bark made vessels in a curious manner,

that would hold about two gallons each: they made above

one hundred of these kind of vessels. In the sugar tree

they cut a notch, sloping down, and at the end of the notch

stuck in a tomahawk; in the place where they stuck the

tomahawk they drove a long chip, in order to carry the

water out from the tree, and under this they set their ves¬

sel to receive it. As sugar trees were plenty and large here,

they seldom or never notched a tree that was not two or

three feet over. They also made bark vessels for carrying

the water, that would hold about four gallons each. They

had two brass kettles, that held about fifteen gallons each,

and other smaller kettles in which they boiled the water,

but as they could not at times boil away the water as fast

as it was collected, they made vessels of bark, that would

hold about one hundred gallons each for retaining the wa¬

ter ; and though the sugar trees did not run every day, they

had always a sufficient quantity to keep them boiling during

the whole sugar season.

“The way we commonly used our sugar while encamped

was by putting it in bear's fat until the fat was almost as

sweet as the sugar itself, and in this we dipped our roasted

venison".

(Concentration of maple sap by freezing)

“Shortly after we came to this pace the squaws began

to make sugar. We had no large kettles with us this year,

226 Wisconsin Academy of Sciences, Arts, and Letters .

and they made the frost, in some measure, supply the place

of lire in making sugar. Their large bark vessels, for hold¬

ing the stock water, they made broad and shallow ; and as

the weather is very cold here, it frequently freezes at night

in sugar time; and the ice they break and cast out of the

vessels. 1 asked them if they were not throwing away the

sugar. They said no ; it was water they were casting away ;

sugar did not freeze, and there was scarcely any in that ice.

They said 1 might try the experiment, and boil some of it,

and see what 1 would get. 1 never did try it ; but I observed

that, after several times freezing, the water that remained

in the vessel changed its color, and became very sweet.”

43. Williams, Samuel 1809

Natural and Civil History of Vermont. Burlington, Vt.,

1809, Vol. 11, 2 ed., p. 363-4.

(Manufactures in Vermont)

“The manufacture of maple sugar is also an article of

great importance to the state. Perhaps two thirds of the

families are engaged in this business in the spring, and

they make more sugar than is used among the people. Con¬

siderable quantities are carried to the shop keepers ; which

always find a ready sale, and good pay. The business is now

carried on, under the greatest disadvantages: Without

proper conveniences, instruments, or works; solely by the

exertions of private families, in the woods, and without any

other conveniences than one or two iron kettles, the largest

of which will not hold more than four or five pailfulls. . . .

This manufacture is capable of great improvements. . . .

And it might become an article of much importance, in the

commerce of the country.”

44. Henry, Alexander 1809

Travels and Adventures in Canada and the Indian Ter¬

ritories between the Years 1760 and 1776. New York, 1809,

(a) p. 69, (b) p. 70, (c) p, 143.

a. (Sap flow)

“When, in the morning, there is a clear sun, and the

night has left ice of the thickness of a dollar, the greatest

quantity is produced.”

Schuette & Schuette — Maple Sugar Bibliography 227

b . (Sugar diet of the Ojibwas)

“. . . we hunted and fished, yet sugar was our principal

food during the whole month of April. 1 have known In¬

dians to live wholly upon the same and become fat”.

c. (Squaws make the sugar.)

“. . . we turned our attention to sugar making, the man¬

agement of which — belong to the women, the men cutting

wood for the fires, and hunting and fishing'7.

45. _ 1814

Coll. Mass. Hist. Soc., [2] 3, 114 (1846).

Note on New Holderness, N. H.

“The prevailing wood is oak, but there is a good deal of

other wood, particularly of pine, beach, and maple. From

the sap of the black, or sugar maple, (acer saccharinum) a

considerable quantity of sugar is made.77

46. Dwight, Timothy 1821

Travels in JSlew-England and New- York. New-Haven,

1821, Vol. 1, p. 40.

(Descriptive)

“The sap of this tree is a very pleasant drink ; and the

sirup is by many persons preferred to honey.77

47. Hunter, John D. 1823

Manners and Customs of Several Indian Tribes Located

West of the Mississippi. Fhiladelphia, 1823, p. 315.

(The sugar month of the Indians)

. . and the thirteenth month is the sugar month be¬

cause in it they manufacture their sugar, from the maple

and box elder trees.77

48. Hunter, John D. 1824

Memoirs of a Captivity among the Indians of North

America, from Childhood to the Age of Nineteen. London.

1824, 3 ed. p. 290.

(Indian’s fondness for maple sugar)

“in districts of country where the sugar maple abounds,

the Indians prepare considerable quantities of sugar by

simply concentrating the juices of the tree by boiling, till it

acquires a sufficient consistency to crystallize on cooling.

But, as the are extravagently fond of it, very little is pre¬

served beyond the sugar-making season. The men tap the

228 Wisconsin Academy of Sciences , Arts, and Letters.

the trees, attach spigots to them, make the sap troughs;

and sometimes, at this frolicking season, assist the squaws

in collecting sap.77

49. Keating, William H. 1824

.Narrative of an Expedition to the Source of St. Peter's

River, Lake Winnepeek, Lake of the Woods, &c. Philadel¬

phia. 1824. Vol. 1, p. 114.

(Description of rude process practiced by Indians)

“We are informed, that they profess to have been well

acquainted with the art of making maple sugar previous to

their intercourse with white men. Our interpreter states

that having once expressed his doubts on the subject in the

presence of Jose Renard, a Kickapoo chief, the latter an¬

swered him immediately with a smile, ‘Can it be that thou

art so simple as to ask me such a question, seeing that the

Master of Life has imparted to us an instinct which enables

us to substitute stone hatchets and knives for those made of

steel by the whites; wherefore should we not have known

as well as they how to manufacture sugar ? He has made us

all, that we should enjoy life; he has placed before us all

the requisites for the support of existence, food, water, fire,

trees, etc. ; wherefore then should he have withheld from us

the art of excavating the trees in order to make troughs of

them, of placing sap in these, of heating the stones and

throwing them into the sap so as to cause it to boil, and by

by this means reducing it into sugar.7 77

50. James, Edwin 1830

A Narrative of the Captivity and Adventures of John

Tanner. New York, 1830, (a) p. 294, (b) p. 321.

a. ( Objibwa names for the sugar maple and

the river maple trees)

Nin-au-tik = sugar maple (our own tree). She-she-

gum-maw-wis = river maple (sap flows fast).

b. (Menominee name for sugar moon)

Sho-bo-maw-kun ka-zho = sugar moon.

51. Bouchette, Joseph 1832

The British Dominions in North America. London,

1832, Vol. 1, p. 371-2.

Schuette & Schuette— Maple Sugar Bibliography 229

Manufactures — Maple Sugar

“Maple sugar will nevertheless ever continue a favour¬

ite luxury, if not a necessity, with the Canadian peasant,

who has not unaptly been considered as having for it the

same sort of natural predilection that an Englishman has

for his beer, a Scotchman for his scones, and a Mexican for

his pulque.7'

52. Evans, Francis A. 1833

The Emigrant's Guide to Canada. Dublin, 1833, p. 105-7.

On Making Maple Sugar

Descriptive.

53. 1837

Graham /., 1, 87

Maple Sugar

Farmers owning “sugar lots" are urged to give atten¬

tion to the subject of making sugar at home, for there are

many who would purchase maple sugar if it were brought

into market in a suitable state for common use. It would

make the best of loaf sugar, and the molasses made from it

would be of a superior quality. Those who are opposed to

the use of “free labor produce" should be interested in this

phase of the subject.

54. Jameson, Anna B. 1838

Winter Studies and Summer Hambies in Canada, Lon¬

don, 1838, Vol. ill, p. 217.

(Manufacture of sugar by the daughter of Waub Ojeeg)

“A large tract of Sugar Island is her property; and

this year she manufactured herself three thousand five

hundred weight of sugar of excellent quality."

55. Ducatel, _ 1846

Catholic Mag ., 5, 92.

A Fortnight among the Chippewas of Lake Superior

“The birchbark is made into troughs (pisketahnahgun)

in which the maple sugar (sinzibuckwud) is gathered in

March and April. _ _ With the birch bark is also manu¬

factured the sugar basket (mukkuk) - - "

56. Sparks, Jared 1848

The Library of American Biography. Boston, 1848,

L2J Vol. Vll. p. 189.

230

Wisconsin Academy of Sciences, Arts, and Letters.

Life of Sebastien Rale

“His constant food was Indian corn, of which, pounded

in a mortar and boiled, he made hominy. The only condi¬

ment he could have was supplied by maple sugar, prepared

in the spring by the women, who collected the sap of the

trees in vessels of bark, and boiled it down.”

57. Gesner, Abraham 1849

The Industrial Resources of Nova Scotia. Halifax,

1849, p. 213.

Manufactories — Maple Sugar

“This sugar may be made as white and as lively as any

from the tropical climates. The kind usually made is sold

in small brown cakes. The sap also affords a delicious syrup,

and the ‘last run' makes excellent vinegar.”

58. Morgan, Lewis H. 1851

League of the Ho-de-no-sau-nee, or Iroquois. Rochester,

1851, p. 369.

(Sugar from the maple)

“Our Indian population have been long in the habit of

manufacturing sugar from the maple. Whether they

learned the art from us, or we received it from them, is un¬

certain. One evidence, at least, of its antiquity, is to be

found in one of their ancient religious festivals, instituted

to the maple, and called the Maple dance.”

59. Schoolcraft, Henry R. 1852

History Condition and Prospects of the Indian Tribes

of the United States. Philadelphia. 1852. Vol. II, p. 55.

Sugar-Making

“It forms a sort of Indian carnival. The article is pro¬

fusely eaten by all of every age, and a quantity is put up

for sale in a species of boxes made from the white birch

bark, which are called mococks, or mokuks. These sugar-

boxes are in the shape of the lower section of a quadrangu¬

lar pramid . miniature mokuks are ornamented with

dyed porcupine quills, skilfully wrought in the shape of

flowers and boxes.

“The heydey scenes of the Seensibaukwut, or sugar¬

making, crown the labors of the spring. The pelt of ani¬

mals is now out of season, winter has ended with all its

Schuette & Schuette — Maple Sugar Bibliography 231

vigors, and the introduction of warm weather prepares the

Indian mind for a season of hilarity and feasting, for which

the sale of his ‘golden mokuks’ gives him some means/’

60. Jones, Electa F. 1854

Stockbridge, Past and Present. Springfield, 1854, (a)

p. 28, (b) p. 26-7.

(Maple sugar made by Stockbridge Indians)

“The Muh-he-con-ne-ak . . . manufactured large quanti¬

ties of Maple Sugar. And indeed we seem to be chiefly

indebted to them for the knowledge of this luxury, for as

late as 1749, Mr. Hopkins, in writing of Stockbridge and

its Indians, not only describes its taste, and the manner in

which it is made , but tells what it is, as if very little

known.”

6. (Legend of the origin of maple sap)

“They had a rare acquaintance with heavenly bodies;

even the children could tell their names ; and it is an inter¬

esting fact, that not only Muh-hu-con-ne-ok, but other New

England Indians, gave the name of “The Bear” and “Great

Bear ” to the same constellation which is so called by Euro¬

pean nations. Their mythological account was this: —

that these stars were so many men engaged in a bear hunt.

They commenced the hunt in the spring, and by autumn

had wounded the animal, so that his blood was falling upon

the forests, and dyeing them with those beautiful hues of

the season. In the winter they slew him, and the snow was

but his dripping oil. - This melted in the spring, and

furnished the trees with sap.”

61. Kohl, Johann G. 1860

Kitchi-Gami. Wanderings around Lake Superior. Lon¬

don, 1860. p. 318.

(Maple sugar as preservative and condiment)

“The Indians dry it (‘pagessaneg’ des prunes sauvages

— wild plum) at times, but more usually boil it with maple

sugar, and make it into a sort of cake, or dough. They boil

and stir the plums in the kettle, until the mass becomes

thick; they spread it out on a piece of skin or birch bark

for the thickness of an inch, and let it dry in the sun. It

supplies a tough, leathery substance, which they roll up and

232 Wisconsin Academy of Sciences , Arts , and Letters.

pack in their ‘makaks’ (birch-bark boxes). These are then

placed in holes in the ground, like so many other things of

their housekeeping, and covered with earth. It keeps sweet

a long time and in winter they cut off pieces, which they

boil with dried meat. ‘(Test bon-bon, monsieur — tout a

fait\

“Whether the art of preserving fruit with sugar is an

old invention of the Indians I am unable to say, but I be¬

lieve so, for it has been ascertained that the manufacture

of sugar was pre-European among the Indians. Besides,

the use of sugar as the universal and almost only condiment

in Indian cookery is most extended. Sugar serves them,

too, instead of salt, which even those who live among Euro¬

peans use very little or not at all. They are fond of mixing

their meat with sweets, and even sprinkle sugar or maple

syrup over fish boiled in water. They have a perfect aver¬

sion for salt. _ That great cookery symbol, the salt-

box, which is regarded among many salt-consuming nations

with a species of superstitious reverence, is hence hardly

ever found in an Indian lodge. But the larger sugar makak

may be always seen there, and when the children are im¬

patient, the mother gives them some of the contents, and

they will sit at the door and eat sugar by handfuls.”

62. _ 1860

Canadian Settlers’ Guide. 1860, p. 66; Chamberlain,

Am. Anthropol., 4, 39 (1891).

(Flavor due to bark vessels)

“The Indian sugar (maple), which looks dry and yellow

and is not sold in cakes but in birch boxes, mowkowks, as

they call them, I have been told owes its peculiar taste to

the bark vessels that the sap is gathered in, and its grain

to being kept constantly stirred while boiling”.

63. Hind, Henry Youle 1860

Narrative of the Canadian Red River Exploring Expe¬

dition of 1857 and of the Assinniboine and Saskatchewan

Exploring Expedition of 1858. London, 1860, Vol. I, p.

127-8.

(Sap of ash-leaved maple used for sugar making)

“The maple, which at one time grew in considerable

quantities near Sugar Point, is not the true sugar maple

Schuette & Schuette — Maple Sugar Bibliography 233

(Acer saccharinum) so common in western Canada, but an¬

other species, generally known as the ash-leaved maple

(Negundo fraxini folium) , also furnishing an abundance of

juice from which sugar is made as far north as the Sas¬

katchewan.”

64. Poore, Ben Perley 1866

Report of the Commissioner of Agriculture for the Year

1866. Washington, 1867, p. 500.

(An Indian festival dish)

“From the sap of the maple tree they made a coarse¬

grained sugar, which, when mixed with freshly pounded

‘suppaun’ and seasoned with fried whortleberries, was

baked into a dainty dish for high festivals.”

65. Wheeler, Timothy 1869

New England Farmer , Boston, Oct. 9, 1869.

(Indian rule for predicting the character of the

sugar season)

“If the maple leaves ripen and turn yellow, and the buds

perfect themselves so that the leaves fall off naturally, with¬

out a frost, then there will be a good flow of sap the follow¬

ing spring ; but if there is a hard frost that kills the leaves

and they fall off prematurely, before the bud is perfected,

then we may look out for a poor yield of sap. In other

words, the flow of sap will be more or less abundant in pro¬

portion to the ripness of the tree before the frost of the

previous autumn”.

66. Leland, Charles G. 1884

Algonquin Legends of New England. Boston, 1881, p.

121.

(A Penobscot legend)

“Now Wasis was the Baby. And he sat on the floor

sucking a piece of maple sugar, greatly contented, troubling

no one.”

67. Brinton, Daniel G. 1885

The Lenape and Their Legends. Philadelphia, 1885,

p. 255.

(Algonkin Indians prefer maple sugar.)

The general name applied by the Iroquois to the Algon-

kins is given as Ratirontaks, from karonta, tree, and ikeks,

234 Wisconsin Academy of Sciences , Arts, and Letters .

to eat, “Tree-eaters”. They were probably so called from

their love of the product of the sugar maple.

68. Blackbird, Andrew J. 1887

History of the Ottawa and Chippewa Indians of Michi¬

gan. Ypsilanti, 1887, p. 72.

(Indian legend of sugar trees)

“The legends say, that once upon a time the sugar trees

did produce sap at certain seasons of the year which was

almost like a pure syrup; but when this mischievous Ne-

naw-bo-zhoo had tasted it, he said to himself, ‘Ah, that is

too cheap. It will not do. My nephews will obtain this

sugar too easily in the future time and the sugar will be

worthless'. And therefore he diluted the sap until he could

not taste any sweetness therein. Then he said, ‘Now my

nephews will have to labor hard to make the sugar out of

this sap, and the sugar will be much more valuable to them

in the future time.' ”

69. Henshaw, H. W. 1890

Amer. Anthropol., 3, 341-51.

Indian Origin of Maple Sugar

“Considering the great familiarity of the Indians with

the natural edible products of America, and the general ig¬

norance of the European on this subject, it is fairly to be

inferred that the a priori likehood of the discovery of the

properties of the maple sap is all in favor of the Indian.”

70. Chamberlain, A. J. 1891

Ibid., 4, 39-43.

The Maple Amongst the Algonkian Tribes

The legend of the Menomine Indians on the origin of

maple sugar making runs as follows: “One day Nokomis,

the grandmother of Manabush, was in the forest and acci¬

dently cut the bark of a tree. Seeing that a thick syrup

exuded from the cut, she put her finger to the substance,

and upon tasting it found it to be very sweet and agree¬

able. She then gave some of it to her grandson, Mana¬

bush, who liked it very much, but thought that if the syrup

ran from the trees in such a state it would cause idleness

among the women. He then told Nakomis that in order to

give his aunts employment and keep them from idleness

Schuette & Schuette — Maple Sugar Bibliography 235

he would dilute the thick sap whereupon he took up a ves¬

sel of water and poured it over the tops of the trees, and

thus reduced the sap to its present consistency. This is

why the women have to boil down the sap to make syrup.”

71. Chamberlain, A. F. 1891

Ibid., 381-3.

Maple Sugar and the Indians

Evidence is presented in support of the claim that the

American Indian first made maple sugar.

72. Carr, Lucien 1895

Proc. Amer. Antiq. Soc., [ns] 10, 155-90.

The Food of Certain American Indians

and Their Methods of Preparing it

“It was made wherever the tree grew, and it found spe¬

cial favor as an ingredient in their preparation of parched

cornmeal or as we call it, nocake or rockahominy. They

also cooked corn in the syrup 'after the fashion of par-

lines/ which was a favorite dish with them, as a similar

preparation is today with us; and in more recent times

they also made a preserve of plums which is said to have

been good. Among some tribes, and in recent times, this

sugar may be said to have taken the place of salt, though

this latter article was known from the earliest times.”

INDEX

Chemical technology in sugar making

Addition of lime aids in scum formation30

Addition of lime to prevent small grain28

Advantages of refining26

Beef suet aid to sugar formation11

Concentration of sap by freezing 38 42

Excess lime imparts reddish cast to sugar23

Excessive boiling causes small grain28

Fats added to inhibit foaming25 36

Graining, claying and refining operations30

Lime, eggs, milk as clarifying agents36

Milk a superior clarifying agent36

Commerce in maple sugar

Cash crop in Vermont38

Future predicted43

Home market wanting33

Market news26 32 53

Profitable article of exportation29

Surplus sold43

Neglected “branch of trade”12

Concentration of maple syrup

Directions for making sugar25 28 30 52

Freezing38 42

Three methods39

With hot stones38 49

Yield of sugar6 7 11 14 28

Dietary virtues of maple sugar

Antiscorbutic40

Might lessen desire for alcoholic stimu¬

lants39

More wholesome than brown sugar27

Readily digested40

Source of health28

Indian festivals

Maple dance of the Iroquois58

Maple dance of the Ojibwas63

Sugar making (seensibaukwut)59

Indian legends

Algonkian68 70

Chippewa68 70

Menominee70

Mohican70

Ottawa68 70

Stockbridge60

Indian names

Makak (sugar basket)01

Michtan (maple tree)1

Mokuk (sugar basket)59

Mowkowk (sugar basket)62

Mukkuk (Chippewa for sugar basket)55

Nin-au-tik (Ojibwa for sugar maple, “our

own tree”)50

Pisketahnahgun (birch bark sugar tub)53

Ratirontak (nickname for Algonkin)67

She-she-gum-maw-wis (Ojibwa for river ma¬

ple, “sap flows fast”)50

236 Wisconsin Academy of Sciences, Arts, and Letters.

Sho-bo-maw-kun-ka-zho (Menominee for

sugar moon)50

Sinzibuckwud (Chippewa for maple sugar)35

Stone tree27

Medicinal virtues of maple sap

Great friend of the breast12

Pectoral12

Medicinal virtues of maple sugar

Antiscorbutic40

Deemed highly by Nova Scotians24

Exceeds West Indian sugar11

Preferred by apothecaries19

Medicinal virtues of maple syrup

Antidote for “the stone”2

Fortifies the stomach10

Remedy for colds and rheumatisms19

Used in preparation of syrup of maiden

hair7

Miscellaneous

Baptismal rites with “maple water”2

Condimental use of maple sugar by Fr.

Rale56

Thos. Jefferson’s maple orchard36

Miscellaneous products made from maple sap

Beer25

Molasses25

Rossolis9

Vinegar25 57

Wine25

Properties of sugar from the maple

American product28

As good for common use as West Indian

cane11

Comparable to good Muscovado14 28

Comparable to that “from the tropical cli¬

mates”57

Equal to best white Havana53

Equals Jamaican in quality18

Flavor due to bark vessels62

Not inferior to “best clayed sugars”35

Pectoral12

Same as West Indian cane31

Smoky taste15

Substitute for common sugar26

Superior to that “of the islands”12 36

Uncontaminated by insects and worms36

Used in preference to cane sugar36

Properties of maple sap

Comparable to waters of Pougue2

Delicious and wholesome12

Pleasant beverage46

Relative qualities of hard and soft maple

saps21 27

Relished by French and Indians2

Superiority of first runnings15

Sweet and pleasant16 _

Taste superior to “limonade or cherry -wa¬

ter”10

Properties of maple syrup

American “melasses” very little inferior to

West Indian18

Indians extravagantly fond of it48

Sap flow

Color of bark an indication of yield35

Effect of temperature upon44

Presence of snow necessary9

Rate2

Relation to side tapped39

Relative yields from hard and soft ma¬

ples21 27

Time27 39

Yield per tree11

Sap gathering and boiling

Children collect “mapnie-water”10

Duty of squaws13 27 44 48

Yield of syrup from sap11

Source of maple sugar

Ash-leaved maple63

Connecticut37

Isles de Richelieu12

Kentucky35

Lake Erie5

Montreal4

New Hampshire23 45

New York28

New York28 35

Nova Scotia24

Ohio22

Pennsylvania28 35

Sweet maple20

Vermont33 38 40 43

Virginia14

Varieties of maple21

Sugar-making accessories

Metallic kettles11 12 27 61

Quill spouts2

Sap receivers of bark13 15

Spouts of cedar,10 elder,39 sumac39

Sugar baskets of birchbark55 59 62

Sugar tubs of birchbark55

Sugar tubs of elm bark42

Troughs of poplar, linden, white ash, ma¬

ple, etc.39

Sugar making a “home” industry

Advantages of culture of the sugar maple25

Estimated acreage to meet nation’s sugar

needs29

Maple sugar not product of slave la-

bor32 41 53

Plea for greater interest in28 29 35 53

Profit from31

Source of national wealth29

Sugar making an Indian art

Ancient practice6 7 69

Antiquity of (Iroquois)58

Pre-European61

Priority of (Kickapoo)49

Stockbridge Indians60

White man instructed Indians12

Sugar-making season

Character of predicted65

Spring, summer and winter27

Thirteenth month47

Synonyms for maple sugar and syrup

American melasses18 28

American sugar28 32

Canada sugar6

Indian sugar62

Technique of sap gathering

Description2 11

Indian’s method42 48

Tapping the tree11

Yield of syrup11

Use of maple sugar by Indians

Condiment61 72

Festival dish71

Highly prized food21 48 61

Measure of wealth54

Parched corn meal preparation72

Principal (seasonal) food21 26 44

Sale of “golden mokuks” a source of rev¬

enue59

Sweetening agent26

Whortle berry-“sup-paun”-sugar mixture71

Wild plum preserve61

Use of maple sap by Indians

Beverage in time of want1

Ceremonial beverage19

Favored beverage2 12

Use of maple syrup by Indians

Corn cooked in maple syrup72

Plum preserve72

Relish of Indian wheat8

Sagamite19

White man’s introduction to the maple tree

Accidental discovery by English14

Birds puncture bark to drink sap35

Frenchmen learn of it from Illinois In¬

dians19

THE GEOGRAPHY OF THE CENTRAL SAND

PLAIN OF WISCONSIN*

J. Riley Staats

The Landscape

The Central Sand Plain of Wisconsin (Figure 1) is a region

in which the physical and cultural elements of its landscape

sharply delimit it from surrounding more highly developed re¬

gions. (Figures 3 and 4) This sandy area, whose flatfish sur¬

face is interrupted locally by numerous sandstone knobs rising

from 100 to 300 feet above the surrounding plain by terraces

along the Wisconsin River, by ridges extending from the Fran¬

conia escarpment, which forms the southern and western bound¬

aries of the region, and locally by low sand dunes, presents a

stage upon which a succession of cultural landscapes has been

imposed.

The development of the Central Sand Plain landscape may

be divided into: (1) Landscape in the Period of French and

Indian Occupance, which was dominant previous to 1830; (2)

Landscape in the Lumbering and Pioneer Stage, 1830 to 1880;

(3) Landscape in the Transitional Stage, 1880 to 1920; and

(4) the Present Landscape.* 1

The Present Landscape

The present scene is distinguished by features of cultural

decadence in the rural areas, and by specialized manufactural

and commercial urban centers along the chief waterways. Va¬

riations in one or more of its physical forms is locally reflected

in cultural responses. Areas of better soils, in scattered loca¬

tions, contain a higher than the average per cent of cultivated

land and better kept farmsteads. Such sections stand as islands

of higher utilization surrounded by or intermingled with areas

of sand which are decidedly less developed. On the other hand,

in some portions of the plain, the development is below the av-

* This study, which was prosecuted largely on the basis of field work of a reconnaissance

nature, on detailed statistical studies, and on a study of the literature, is an abridgement of a

dissertation. Abbreviation of the paper has necessitated the omission of much data, most of

the pictures and maps, many footnotes, and the bibliography.

1 On account of a limited space a description and interpretation of the present scene only

is attempted.

237

238 Wisconsin Academy of Sciences , Arts, and Letters.

Ph.D. thesis of Loyal Durand, Jr., University of Wisconsin, 1930.

Stoats — Central Sand Plain of Wisconsin

239

I. Eastern farming and dairying area

a. Outwash plain

b. Marshland and associated sandy strips

c. Sand dunes and brush section

d. Flood plains, terraces, and areas of better soil

II. Southeastern farming and dairying area

III. Central marsh area

a. Island of higher utilization

IV. Western brush and dairying area

_ _ _ regional boundary

240 Wisconsin Academy of Sciences , Arts, and Letters ,

crop land total pasture

Fig. 3

Fig. 4.

RYE POTATOES

Fig. S.

Fig. 6.

TRANS. WIS. ACAD., VOL. 29.

PLATE IV

Fig. 2. — A view of the wooded portion of the Central Sand Plain Fig. 4. — A typical view in the Southeastern Farming and Dairying Area

Stoats— Central Sand Plain of Wisconsin 241

erage for the region as a whole. These parts contain little in

the way of cultural imprint other than evidences of past forest

exploitation as shown by many stumps among the brush.

Fluvio-glacial sediments, alluvium, residual materials, and

loess, individually or in combination, have given rise to a vari¬

ety of soils that may be classified under the general term of pine

sands.2 Of the soil types, including sands, sandy loams, and

peat, sand is by far the most widespread, and it is to this soil in

its various phases more than to any other element of the physi¬

cal complex that the economic aspects of the region are closely

related. In addition to its low plant food content, the coarse

soil imposes a severe restriction upon general agriculture be¬

cause of its droughty nature.

A high percentage of the Central Sand Plain is cut-over and

brush lands (Plate 4, Figure 2) in which jack pine (Pinus

Banksiana) and scrub oak (Quercus nan, Quercus marilandica,

and Quercus coccinea) predominate. Only remnants of the for¬

mer cover of white (Pinus strobus) and Norway (Pinus resin-

osa) pines, which extended over a considerable portion of the

region, are found scattered over the area. These remnants stand

as lone sentinels or in small, widely-scattered patches of a few

acres in extent. In some localities, instead of pure stands of

jack pine and scrub oak, one finds variations in the natural

cover, including other species such as aspen (Populus tremu-

loides), willow (Salix Viminalis and Salix Purpurea), and tam¬

arack (Lerx laricina). The aspen, willow, and tamarack are

generally found in areas of poorly drained soils, such as the

Dunning series which surround the marshes, and in the marshes

themselves. Where scrub oak is in pure stands or is mixed with

jack pine, hazel (Corylus Americana) and sweet fern (Comp-

tonia aspenifolia) undergrowth are common; in other parts of

the region where jack pine is found in pure stands the ground

cover is generally a sparse growth of grass which incompletely

mantles the underlying sand.

Not all of the untilled areas are covered with brush and

scrub timber, since marsh occupies large tracts of the Central

Sand Plain. In these sections the dominant soil is peat. The

utility of such land is low, since it is limited to the production of

marsh hay, cranberries, blueberries, and moss, unless it is

2 The term “pine sands” is used to include several varieties of quartz sand which is very-

low in plant food and in which pines and scrub oak are practically the only trees that thrive.

242 Wisconsin Academy of Sciences , Arts , and Letters.

drained. If drained, a general type of farming is carried on

with varying results.

The climatic conditions of the Central Sand Plain are those

peculiar to middle latitudes in continental locations. This region,

according to Koppen’s classification, lies in the Dfb climate. A

growing season of 125 to 146 days, too short for corn to mature

properly, constitutes another phase of the restrictions imposed

upon the use of the area for general agriculture. An annual

precipitation of 30 to 35 inches, with the maximum coming in

the growing season, is usually sufficient for the kind of crops

that are commonly grown in an environment of sandy soils,

short growing season, and mean summer temperatures not ex¬

ceeding 71 degrees. It frequently happens, however, that dur¬

ing the latter part of summer of some years crops suffer from a

lack of moisture, since a small decrease of precipitation during

this period of the year is critical in sandy soils that are charac¬

teristically droughty.

Of the region as a whole, only 43.9 per cent is in farms, and

of this amount 38.7 per cent is in crop land and 40.1 per cent in

pasture.3 The latter percentage represents all types of pasture,

the greater part of which is wooded. These figures reveal the

main uses of the farm land, yet they are somewhat misleading,

since neither the quality of the pasture nor the crop yields are

comparable to those of surrounding regions. Dairying is the

chief occupation, but here again, in comparison with contiguous

areas its development is meager indeed. (Figure 7) In most

parts of the Central Sand Plain the money derived from the

sale of milk and cream is the chief farm income. The silo, which

is characteristic of adjacent dairying areas, is generally absent

from the region except in areas of better soils where it is found

in connection with nearly every barn.

Crops of the Central Sand Plain include small grains (rye,

oats, buckwheat, and some barley) ; corn, tame hay (including

clover and timothy, alfalfa, soya beans, and German millet) as

feed and forage crops for the dairy industry; potatoes, cran¬

berries, and vegetables, all three as cash crops. The sandy plain

does not, however, have a large acreage of any of the above

crops. Furthermore their acreages are unevenly distributed be-

8 All percentages, unless otherwise stated, were compiled from data obtained at the Crop and

Livestock Reporting Service of Wisconsin, and from the United States Census, both giving figures

for civil towns.

Staats— Central Sand Plain of Wisconsin 243

cause of the variations in the fertility of the soils in the differ¬

ent parts of the region. In comparison with surrounding areas

no crop, with the exception of rye, is very important. (Figures

3 and 5)

The cropped fields in portions of the plain are rectangular

following the pattern of the land survey of the Northwest Ter¬

ritory, (Plate 4, Figure 1) while in other sections their shapes

are adjustments to the local physical conditions. In the gently

rolling areas where the land between the the ridges is cleared

and the ridges are generally wooded (Plate 4, Figure 4), the

shapes of the cleared and cropped fields tend to conform to the

trend of the ridges and dunes. Thus it is that the cleared areas

break the monotonous continuity of brush and marsh lands, which

are so characteristic of the Central Sand Plain.

The farms of this region average 160.4 acres in size, an

acreage exceeded in Wisconsin only by that in the Western Ridge

and Valley Region wherein the farms average 161.7 acres. Of

the farm land of the region, 21.3 per cent is waste, used neither

for crop land, farmsteads, nor pasture, a percentage of waste

and unused land that is exceeded only by that of the farms in

the northern part of the state.4

Small farmsteads, situated in small clearings or on the edges

of marshes, are characteristic landscape features. A few farm¬

steads are found widely scattered in the drained portions of the

marshes adjacent to the drained ditches or along the highways

in those parts of the marsh lands which have been brought

wholly under cultivation. A three to four room house, often

unpainted, a small barn, and a few sheds and small outbuildings

comprise the structures of the typical farmstead — the sand

country cannot support a larger capital investment in improve¬

ments.

The Central Sand Plain of Wisconsin is sparsely settled, and

for the region as a whole, rural in character. The density of

population is 24.4 per square mile, and 34 per cent of the people

live on farms, giving a density of farm population of 8.3 per

square mile.5 Country stores, small cross-road hamlets, and vil¬

lages of a few hundred inhabitants are more characteristic than

larger villages and cities. Moreover, the cities that are closely

related to the Central Sand Plain have two general types of lo-

4Ebeling, Walter, H., et. al., Wisconsin Agriculture, Bulletin 140, p. 42, 1932.

5 Ibid, p. 40.

244 Wisconsin Academy of Sciences , Arts , and Letters .

cations, namely: (1) along the Wisconsin River, and (2) near¬

by in other regions.

The cities and many of the villages have in their brief his¬

tories passed through a series of functional changes. Generally

they started in connection with the lumbering industry as sites

for saw-mills, as outfitting centers for the lumber camps or

both. If they were able to adapt themselves to changed condi¬

tions at the close of the lumbering operations, they declined but

temporarily, and then took on new activity in commercial or

industrial functions. If, on the other hand, their sites offered

no special advantage for industry or their locations no oppor¬

tunity for commerce they passed into decadence — -the Central

Sand Plain, without some special advantages other than those

contained in its general upland, cannot support a city of much

magnitude.

MILK COWS

The small hamlets and the villages in the Central Sand Plain

are, for the most part, located either on branch railroad lines or

improved trunk highways. These small and widely scattered

units are tied by lines of transportation to the cities along the

Wisconsin River and to the near-by cities of the surrounding

regions.

Regional Subdivisions

As previously stated, the Central Sand Plain of Wisconsin is

delimited by sharp changes in both the cultural and physical

elements of the landscape. (Figures 3, 4 and 7) Passing from

Staats— Central Sand Plain of Wisconsin 245

the plain into contiguous regions, an observer passes from a

region of sandy soils, low percentage of improved land, small

farmsteads, and a high percentage of wooded and waste lands

to regions of heavier soils, (except in the Glaciated Sand Re¬

gion), a higher percentage of cleared and improved land, and

larger and better kept farmsteads. Thus, the human imprint is

far more conspicuous in the surrounding regions than in the

Central Sand Plain itself. Moreover in the region as a whole

there is a high degree of unity in soils, topography, natural veg¬

etation, and cultural forms, but by a more detailed survey suffi¬

cient differences were discovered to warrant subdividing the

region, for the purpose of description and interpretation into:

(Figure 2)

(1) An Eastern Farming and Dairying Area (1240 sq. mi.)

(2) A Southeastern Farming and Dairying Area (250 sq.

mi.)

(3) A Central Marsh Area (610 sq. mi.), and

(4) A Western Brush and Dairying Area (1110 sq. mi.

The Eastern Farming and Dairying Area

Similar to the region as a whole, a general view of the East¬

ern Farming and Dairying Area presents a wooded terrain

(Plate 4, Fig. 2) whose general flatness is broken locally by

minor surface configurations. From the crest of one of the

sandstone knobs an observer can look over a monotonously flat

surface that fades into obscurity in the distance. A closer ex¬

amination, however, reveals irregularities that result from

sandstone outliers, areas of sand dunes, and terraces along the

rivers and their larger tributaries. In the localities of sand

dunes the terrain becomes gently undulating; and adjacent to

the Wisconsin River, in many places, the rise from one terrace

to another appears, from a distance, as a low range of hills.

Varying degrees in intensity of utilization, and often a lack

of proper methods to maintain soil fertility characterize the

agricultural use of the Eastern Farming and Dairying Area.

It is not uncommon to find a farm that has been properly man¬

aged having its buildings in good repair while a farm near-by,

which has not been properly cared for, has been abandoned

(Plate 4, Figure 3). Where the better soils prevail, due to the

246 Wisconsin Academy of Sciences , Arts , and Letters.

silt content or the proximity of the underlying clays laid down

in Glacial Lake Wisconsin, well developed farms are prevalent.

Between such areas of better soils, however, wood expanses pre¬

vail in which small clearings are found which may or may not

contain a small farmstead. The percentage of land in farms in

this subdivision varies from approximately 20 per cent in the

western part to over 80 per cent in the eastern as compared

with approximately 43 per cent for the region as a whole. From

20 to 40 per cent is in crops and plowable pasture, which is a

higher percentage than that of the other subdivisions save the

Southeastern Farming and Dairying Area. In fact, in some

parts of the western portion of the Central Sand Plain less than

2 per cent of the total area is in crop land and plowable pasture.

The Eastern Farming and Dairying Area is one of mixed

farming and dairying with a tendency towards specialization

in one or more crops. In the northeastern portion of the area

potatoes are the chief cash crop (Figure 6), and in the south¬

eastern part rye becomes the chief cash crop (Figure 5). In

both of the above sections, however, dairying is important, and

landscape features associated with dairying are conspicuous.

The chief crops are rye, oats, and corn, occupying 18.3, 11.2,

and 12.3 per cent of the crop land respectively. These are raised

primarily for feed and forage crops, although some rye is sold.

In its detailed features the Eastern Farming and Dairying

Area may be divided on the basis of these differences into: (1)

an outwash plain, (2) a marsh and associated sandy strip, (3)

a sand dune and brush land section, and (4) flood plains, ter¬

races, and islands of better soils. (Figure 2)

The Outwash Plain-— An outwash plain, varying in width

from two to four miles, lies adjacent to the western base of the

moraine which forms the eastern boundary of the Central Sand

Plain. This seemingly flat land is highly developed in compar¬

ison with other parts of the region. A substratum of finer soil

and the incorporation of minerals leached from the moraine

give to the soil in this strip a higher quality than that gener¬

ally found elsewhere in the sandy area. It is this quality of soil

which accounts for the higher development, a development in

which over 40 per cent of the total area is in crops.

In the northern half of the outwash plain one finds a ten¬

dency towards specialization in the cropping practices in which

Stoats — Central Sand Plain of Wisconsin

247

potatoes are generally the chief source of farm income. In this

section fields of potatoes varying from 5 to 15 acres in size are

found on nearly every farm. The loose, friable soil of the out-

wash plain is a favorable environment for a tuber crop.

Even though potatoes are extensively grown, the importance

of dairying is also apparent. Barns are much larger than those

on the average farm of the Eastern Farming and Dairying

Area, most of them have gambrel-roofs instead of inverted “V”

types, and for the most part the barns have one or two silos in

connection.

Towards the southern part of the outwash plain, a change

in the landscape becomes apparent. The crop associations are

slightly different. While potatoes are still raised on a commer¬

cial scale they are less important in the farm economy. This is

due to the droughtier condition of the soil, a condition that re¬

sults from a substratum of gravel rather than one of fine ma¬

terials such as characteristically underly the soil in the northern

part of the plain. Fields of rye largely replace the potato fields,

and rye becomes the chief cash crop in the southern section.

(Figures 5 and 6)

The Marsh Lands and Associated Sandy Strips— The out¬

wash plain merges, along its western border, into a strip of

flattish marsh and associated sandy islands and ridges. In the

northern portion of this marshy area, artificial drainage ditches

carry away excess water, and as a result, the flat area is used

for pasture, tame hay, and to some extent for general farming.

The southern part of the strip, on the other hand, is undrained,

and largely given over to marsh hay, waste land, and occasional

small patches of farm land.

The fields of the drained portion of the marsh are large and

the buildings of the widely spaced farmsteads are more preten¬

tious and in better repair than those in the undrained part. In

the undrained portion of the marshy strip the farm buildings of

the widely spaced farms are generally small, in poor condition,

and located on some sandy island or ridge within the marsh, or

on its border. Surrounding the barns of the latter farmsteads

are found a number of stacks of marsh hay which add to the

meager supply of forage crops for dairying.

248 Wisconsin Academy of Sciences , Arts, and Letters.

The Sand Dunes and Brush Land Section — The area of un¬

dulating sand dune topography is largely a section of cut-over

brush composed of jack pine and scrub oak in which small scat¬

tered farmsteads are found along the roads at widely spaced

intervals, or in isolated locations. In the latter instances the

farms are accessible only by means of winding trails — merely

two ruts in the sand. Less than 10 per cent of the section is

cleared, and only a portion of this amount is farmed. The

cleared fields are generally fenced to protect the crops from be¬

ing grazed over by cattle that roam at will through the brushy

area in search of the small amount of forage that such vegeta¬

tion affords. In the cleared plots, corn, rye, and potatoes are

the chief crops raised, and, due to the excessive drainage and

low fertility of the soil, these crops often produce meager yields.

The farm income from this section is derived primarily from

two sources, namely: (1) from a small amount of cream, the

end product of the meager dairy industry on most of the farms

of the section, and (2) from pine bolts which are hauled to the

paper mills at Nekoosa, Port Edwards, or Wisconsin Rapids.

Conspicuous features associated with the latter source are the

many piles of bolts that one may observe along the roads, and

in small clearings which may be reached by winding trails

through the brush.

Another conspicuous feature connected with the pulp wood

industry is the acreage of reforested lands which the paper

companies are enlarging at the rate of approximately 3,000

acres per year. These forested plots, with their rows of trees

of uniform heights and of the same species, stand in marked

contrast to the heterogeneous expanses of cut-over lands that

characterize the general wooded landscape. Beginning in 1950

the Nepco Paper Company hopes to have available a sufficient

supply of pine bolts of its own in the Central Sand Plain.

Flood Plains, Terraces, and Islands of Better Soils — The

present flood plain of the Wisconsin River is wooded and its use

is limited to a scanty amount of grazing together with a patchy

development of summer cottages. On either side of the present

flood plain of the river the surface rises through a series of ter¬

races which are more pronounced in the northern than in the

southern portion of the plain. These terraces contain fragmen¬

tary patches of sandy loams interspersed with small areas of

Stoats — Central Sand Plain of Wisconsin

249

sand. Such a distribution of soils is reflected in the cultural

pattern of the section. The better soils show a high percentage

of improved land while sandy areas are either largely wooded,

or having been cleared, they are now mostly abandoned. The

patchy character of the areal scene is further epitomized by the

types of homesteads; the prosperous appearance of those on

the loams stand out in contrast to the smaller homesteads, often

in disrepair or even abandoned, that are found in the sandy

portion of the terraces.

Islands of better soils, varying in size from less than 5 to

over 10 square miles, are found beyond the terraces on the east¬

ern side of the Wisconsin River. Here again, such islands in

comparison with surrounding localities are centers of intensive

farming and dairying.

Urban Landscape

As previously stated, the Central Sand Plain, generally is

rural. However, 42 per cent of the people live in cities of over

1,000 inhabitants. These urban centers have a distinct pattern

in their arrangement. The cities of the region, save Wisconsin

Dells, are in the Eastern Farming and Dairying Area, located

on the Wisconsin River, a stream with which they are inti¬

mately related. The greater number of the villages are, like¬

wise, located in this eastern subdivision, the largest and one of

the best developed areas of the plain. Classified according to

their functions, the urban centers may be divided into: (1)

villages whose functions are primarily commercial, (2) urban

centers, dominantly manufactural, and (3) urban centers, com¬

mercial and manufactural.

Commercial Villages— The small commercial villages, rang¬

ing from' less than fifty to several hundred inhabitants, serve

local trade areas, and each is connected by rail or by improved

highways to cities within the area or with those on the periph¬

ery. Some of these centers contain creameries where butter is

made while others are merely centers for collecting cream to be

sent to larger towns and cities.

Plainfield, a village of 537 inhabitants, is representative of

the small urban centers of the Eastern Farming and Dairying

Area. The whole village complex reflects its functions. It is

dominantly a commercial center, containing feed stores, lumber

250 Wisconsin Academy of Sciences , Arts , and Letters .

and coal yards, potato warehouses, gasoline storage tanks, and

a small stockyard, in addition to the usual array of merchan¬

dising and service establishments that characterize a commer¬

cial village of comparable size.

Urban Centers , Dominantly Manufactural — Nekoosa, Port

Edwards, and Biron, urban centers which are dominantly manu¬

factural, started as saw mill towns making use of the available

water power at rapids in the Wisconsin River where the stream

had cut its channel to the underlying crystalline rocks. With

the passing of the timber, the saw mills were replaced by paper

and pulp mills, the industries upon which the cities are now de¬

pendent for their existence. The trade areas which these neu-

clei serve are practically confined to their corporate limits ; and

the residential sections are occupied almost exclusively by peo¬

ple connected either directly or indirectly with the paper indus¬

try.

As previously stated, the paper mills, located at these centers

were established at sites of former saw mills where both water

power and an adequate water supply were available, factors

which have located the paper mills in the Central Sand Plain.

In relation to raw materials and markets, location in this region

is disadvantageous, since most of the raw materials come from,

and the finished products must be sent to outside regions. The

pulp wood comes largely from northern Wisconsin, Minnesota,

and Canada; the sulphur from Louisiana, and the limestone

from northeastern Wisconsin and Michigan. The markets are

more widespread, since they include central and eastern United

States.

A distant view of Nekoosa, a representative of the manufac¬

tural group, includes lofty smokestacks of the paper mill, located

on the west bank of the Wisconsin River, and the city’s water

tower rising above what appears to be a grove. Approaching

the city, down the terraced descent to the river, the outlines of

the mill and the city’s structure come into view beyond the arti¬

ficial lake that has been formed by a dam across the river at

this point. The paper manufacturing plant and its associated

storage yard and railroad tracks are the most distinctive fea¬

tures of this urban landscape, since the residences and the com¬

mercial establishments are, in the main, like those features of

other cities of comparable size. The rail lines leading north-

Stmts— Central Sand Plain of Wisconsin 251

ward from the mill separate into a series of spur lines which

penetrate the various parts of the storage yard, an area of ap¬

proximately 30 acres, in which are the long ricks of bolts and

the coal sheds.

Urban Centers , Commercial and Manufactural— Stevens

Point (13,000) and Wisconsin Rapids (8,000) are the urban

centers in this classification. They differ from the preceding

class in: (1) their functions, which include both commercial

and manufactural activities, (2) their size, and (3) their rela¬

tion to the Central Sand Plain in which they are located. As in

the preceding class of urban centers, paper mills are important,

but in Stevens Point and Wisconsin Rapids other establishments

such as furniture factories which had their beginning during

the lumbering stage, feed mills which supply dairy feeds to the

Central Sand Plain and contiguous areas, and milk products

plants are included in the manufactural aspects of these cities

as well. Due to the range of functions found in these cities,

they have become considerably larger than the manufactural

centers previously discussed.

The commercial functions are more closely related to the

Central Sand Plain than are the manufactural. The trade areas

of these cities include the northern part of the plain as well as

parts of contiguous regions. Their manufactural functions,

while related in part to the region in which they are located,

are dominantly related to more remote regions, even to the ex¬

tent of the whole United States and a part of Canada.

In the general structure of Stevens Point and Wisconsin

Rapids, the manufactural and commercial sections are adjacent

to the Wisconsin River where power facilities and transporta¬

tion lines are concentrated. The manufactural plants are usually

next to the stream, and these in turn are flanked by storage and

wholesaling establishments, which are succeeded by retailing

centers. The retailing establishments are concentrated along

one or more of the principal streets, both paralleling the storage

and warehousing facilities and at right angles to them. Resi¬

dential aspects of these cities are not unusual for cities of com¬

parable size.

252 Wisconsin Academy of Sciences, Arts, and Letters.

The Southeastern Farming and Dairying Area

The Southeastern Farming and Dairying Area, embracing

the southern portion of Adams and parts of Juneau, Columbia

and Sauk Counties, has greater relief than any of the other sub¬

divisions of the Central Sand Plain. Differences of 200 to 300

feet between the ridge tops and the intervening valleys, within

short distances, are not uncommon. (Plate 2) The sandstone

ridges and knobs become more numerous towards the southern,

western, and northwestern borders of the region, where por¬

tions of the strata that formerly continued into the West Central

Valley Region and the Western Ridge and Valley Region (Fig¬

ure 1) have been cut off from the main body by erosion.

The soil of the Southeastern Farming and Dairying Area,

while varied, is, for the area as a whole, more suitable for farm¬

ing than that of the other subdivisions of the Central Sand

Plain. An exception to this quality of soil, however, is found in

the central portion of the area where there is a predominance

of sandy soil of relatively low utility. In this particular section

“blowouts” are not uncommon; jack pine is dominant in the

tree cover ; and fallow and waste land occupy a high percentage

of the cleared portion of the section. Leaving the central part

of the area and going towards the borders, jack pine is largely

replaced by oak and sandy loams become the dominant soils.

The sandstone knobs and ridges in these parts are generally

capped with Lower Magnesian dolomite rather than Dresbach

sandstone, and as a result of the weathering of the dolomite, the

soils have higher silt content which gives to them a higher qual¬

ity.

Soil and topographic differences within the area give a dis¬

tinctive pattern to the cleared and wooded sections. The crests

and upper slopes of the ridges and mounds are wooded, while

the lower slopes and intervening valleys are either farmed or

devoted to cleared pasture. (Plate 4). A second Characteristic

location of wooded lands is along the streams, where narrow

ribbons of brush wind about, in conforming to the stream

courses.

This subdivision differs from the Eastern Farming and

Dairying Area, moreover, in having a better quality of soils, a

higher degree of development, and a generally better quality of

homes. For the area as a whole there is a higher percentage of

Stoats — Central Sand Plain of Wisconsin

253

improved land.6 In this subdivision the average farm contains

from 8 to 15 dairy cattle ; the figure for the Central Sand Plain

as a whole is 7.8.7

The homesteads are tied to each other and to the surround¬

ing areas by a net of highways, whose rectangular pattern cor¬

responds in general to section lines, a pattern that is interrupted

in parts of the Central Sand Plain by marshes and brushy waste

lands. The surfaced roads are thus spaced sufficiently close to

give accessibility to all the farms within the subdivision, an im¬

portant factor in an area where the inhabitants must make fre¬

quent use of the highways in transporting milk and cream to

the condensaries and creameries.

The urban development of this subdivision is limited, since

most of the urban centers serving the area have a peripheral

location in relation to it. An exception is the city of Wisconsin

Dells, which is well within its borders. This center is predom¬

inantly a resort city. It owes its importance as a resort center

to its proximity to the scenic features of the Dells of the Wis¬

consin River, which is a part of the gorge of the glacially-di¬

verted Wisconsin River. The most conspicuous features of this

city are many hotels and amusement establishments connected

with the resort business. In addition, summer cottages along

the west bank of the river, boat houses, boats anchored at the

piers, parking spaces, and booths with souvenirs and tourists’

needs add to the unique character of Wisconsin Dells and cause

it to be distinctly different from other cities of the Central Sand

Plain.

by fallow and the stubble of small grains. Most of the wood-

Minor manufactural features in addition to its resort aspect

are found within the city, such as a small creamery and a dairy

fixtures plant. The creamery draws practically all of its cream,

amounting to 7,000 pounds per day,8 from the Central Sand

Plain; the dairy fixtures plant might easily be missed by the

casual observer passing through the city.

•Subdivision Per Cent of Total Per Cent

Area in Farms of Total

Area in Crops

Eastern Farming and Dairying Area . . . 60 27.0

Southeastern Farming and Dairying Area . . 71 32.0

Central Marsh Area . 26 5.6

Western Brush and Dairying Area . 34 9.5

Statistics from Crop Reporting Service of Wisconsin for 1929 and United States Census

of 1930 (Figures for 1929).

1 Ebeling, Walter, H., et. al., Wisconsin Agriculture, Bulletin 140 (1932), pp. 36-43.

8 Personal interview with the manager.

254 Wisconsin Academy of Sciences , Arts , and Letters .

The Central Marsh Area

The Central Marsh Area, comprising 610 square miles, is

the least developed of the subdivisions of the Central Sand

Plain (p. 17). It is a level featureless plain except for a few

sandstone crags that rise abruptly from its surface, and numer¬

ous low sand islands. The islands are but a few feet above the

general level, and therefore do not produce marked topographic

features. In the marsh localities jack pine, aspen, willow, and

tamarack, with an undergrowth of sedge and blueberries, com¬

prise the natural cover; on the sand islands scrub oak with an

undergrowth of hazel and sweet fern, and scattered jack pine

make up the vegetation complex; while the tops of the sand¬

stone knobs are either devoid of vegetation or have a sparse

growth of pine. In fact, the change in vegetation is so sharp

that the sandy islands may be located by noting the vegetation

of the area.

The small amount of general farming that is carried on in

the Central Marsh Area is confined primarily to the northern

border, adjacent to the North Central Dairy Region of Wiscon¬

sin, (Figure 1) a region intensively developed, and to an island

of approximately 60 square miles of higher utilization in the

southeastern portion of the area (Figure 2). The well kept

farmsteads, with comfortable homes, large red dairy barns and

their accompanying silos found in this island, are comparable

to like features in the West Central Valley Region to the south.

The remainder of the general farming is found scattered over

the subdivision, generally along the drainage ditches. In this

portion of the Central Sand Plain there is a large drainage pro¬

ject that hindered rather than helped to adjust the use of the

area to its natural conditions.

The chief importance of the Central Marsh Area for farm¬

ing lies in its specialized crops, such as cranberries, sphagnum

moss, and blueberries. Wisconsin ranks third among the states

of the United States in the production of cranberries, having

1,150 acres in producing marshes, of which the Central Marsh

Area contains 1,090 acres,9 or approximately 95 per cent of the

state's acreage. The present production lies mainly in two sep¬

arate areas: (1) southern Wood County, for which the com¬

mercial outlet is Wisconsin Rapids, and (2) southeastern Jack-

0 15th Census of United States.

Stoats — Central Sand Plain of Wisconsin 255

son, northeastern Monroe, and west central Juneau Counties,

for which the outlet is Mather.

The natural environment in these two districts is very favor¬

able for cranberry production. The summers are fairly cool

(p. 4), which is a normal requisite; the necessary acidic peat

soils are at hand; when sand is used in the marshes it can be

obtained near-by at the margin of the marsh ; and the flat ter¬

rain can easily be converted into wide reservoirs to supply the

necessary water, by building low dams at right angles to the

natural drainage.

Sphagnum moss, which requires an abundance of ground

water, is found in the same general areas in which cranberries

are produced. Artificial drainage has almost restricted this

plant, formerly far more widespread, to the northern and wes¬

tern fringes of the Central Marsh Area where the water table

is higher. Over 80,000 bales, each weighing approximately 20

pounds, are shipped yearly from Wisconsin Rapids. This is es¬

timated to be from one-half to two-thirds of Wisconsin's out¬

put, a state which produces from two-thirds to three-fourths of

the commercial crop of the United States.10

The Western Brush and Dairying Area

As the name implies, the Western Brush and Dairying Area

is largely brush covered. In it small clearings are found where

feed and forage crops are raised for a limited dairy industry.

Similar to the Central Marsh Area, large portions of this sub¬

division are waste land (Figures 3 and 4) in which there are

practically no inhabitants. In such areas one may drive for

miles and see no cultural imprint save the two ruts in the sand

that suffice for roads. In some localities, however, as near some

of the sandstone outliers, and in isolated patches, the ubitiquous

sand is replaced by sandy loams, and as a result the expanses

of waste and isolated farmsteads are replaced by limited areas

of intensive agriculture. It often occurs that these well devel¬

oped areas encircle the outliers in bands varying in width from

a half mile to over a mile.

On the average farm the greater part of the land is covered

with brush and cut-over timber, but most of the timbered por¬

tion of the farm is pastured even though such types of pasture

10 Data supplied by the Wisconsin Moss Company of Wisconsin Rapids

256 Wisconsin Academy of Sciences , Arts, and Letters.

furnish but scanty forage. Woodland pasture is supplemented

by fallow and the stubble of small grains. Most of the wood¬

land is unfenced, and whether privately, state, or county owned,

it is grazed over indiscriminately by cattle from near-by farm¬

steads.

The three chief crops, clover and timothy hay, oats, and

corn occupy 76 per cent of the cropped land. To supplement

the forage produced by clover and timothy and that supplied by

corn fodder, soya beans are raised, and wild hay is cut from the

scattered areas of marsh. The importance of marsh hay is em¬

phasized by stacks of it about the barns throughout the area.

The urban development in both the Western Brush and

Dairying Area and the Central Marsh Area is meager. With

respect to both of these subdivisions the cities are peripheral,

and only cross-road villages are found within the areas. These

small centers serve as foci for collection of the small amount of

produce, and for distribution of commercial products to their

limited and relatively barren trade areas.

Urban Development of the Entire Region

The limited agricultural development and the sparse popu¬

lation of the Central Sand Plain are not sufficient for extensive

urbanization. Most of the Central Sand Plain is included in

the trade areas of peripheral cities, whose chief sources of pro¬

duce are the better-developed regions in which they are located.

The low fertility and the droughty condition of the sandy soils,

which are so widespread in the Central Sand Plain, restrict the

agricultural development of the region. Furthermore, these ad¬

verse environmental conditions augment the decadence of the

rural areas so that more and more land is either abandoned or

reforested. It is only where special features exist, such as sites

for water power with their associated paper mills, and scenic

features such as the Dells of the Wisconsin River, that urban

centers within the Central Sand Plain seem to be stable.

INSOLUBLE RESIDUES FROM WISCONSIN

SEDIMENTARY ROCKS*

PART 1. INSOLUBLE RESIDUES AS AN AID IN THE STUDY OF

SEDIMENTARY ROCKS

R. R. SHROCK

University of Wisconsin

Introduction.— it is a well known fact that almost every sed¬

imentary rock leaves some insoluble residue when digested in

hydrochloric acid. The amount of this material may range from

less than 1% (as in very pure limestones and dolomites) to

more than 99% (as in very pure sandstones). While percent¬

ages of insoluble material have been recorded from chemical

analyses for many years, little attention has been paid to that

material until very recently. It is the purpose of this brief dis¬

cussion to call attention to the uses that may be made of insol¬

uble residues in the study of sedimentary rocks.

POSSIBLE USES OF INSOLUBLE RESIDUES

1. All students of sedimentary rocks are aware of the uses

which have been made of diagnostic fossils (both macroscopic

and microscopic), heavy detrital minerals, and insoluble resi¬

dues for purposes of correlation. Work now in progress on

Wisconsin Silurian dolomites has shown that certain residues

appear to mark definite horizons in wells and exposures. Many

problems of a similar nature, involving the determination of the

amount, character and possible stratigraphic significance of in¬

soluble residues in both well cuttings and exposures are await¬

ing investigation in Wisconsin as well as elsewhere.

2. Paleontologists are following with interest the rapid in¬

crease in number and variety of micro-fossils (foraminifera,

conodonts, scolecodonts, etc.), which are being recovered from

calcareous rocks by acid digestion (See Plate V, Figs. 4, 6-8,

* Part 1 is written specifically to acquaint thesis students in Geology with the methods and

potentialities of insoluble residue analysis, and to suggest problems for future investigation. It

represents one of the research projects made possible by a grant from the Wisconsin Alumni

Research Foundation to the writer for the fall semester, 1933-34. Part 2 gives in abstract form

the results of four thesis projects, investigated in the Sedimentation Laboratory of the Department

of Geology at the University of Wisconsin, under the direction of Drs. W. H. Twenhofel and

R. R. Shrock. Laboratory materials for these projects were furnished by the Milwaukee Public

Museum.

257

258 Wisconsin Academy of Sciences , Arts, and Letters .

13). Not only are these of value for correlation but they also

help to picture life conditions and relations at the time they

lived. Investigation, by insoluble residue analysis, of Pre-Cam¬

brian calcareous rocks might produce significant results con¬

cerning the amount and character of the life of that time. Much

may be learned about the very early growth stages of certain

organisms if their youthful shells can be found. Such shells

have been recovered from certain Wisconsin dolomites, and fur¬

ther investigations along these lines may produce interesting

results.

3. Many investigators have found that insoluble residues

aid in reconstructing conditions of sedimentation. The basal

beds of the Black River limestone in Indiana, Illinois and Wis¬

consin frequently contain a high percentage of detrital quartz.

When released from the calcareous matrix, the grains are found

to be well rounded and frosted (See Plate V, Figs. 5, 9). These

conditions obviously suggest affinities with the underlying St

Peter sandstone. The basal beds of the St. Lawrence dolomite,

the Oneota dolomite, the Black River limestone and the Devon¬

ian of Wisconsin need to be studied for their detrital content.

Original chert nodules, siliceous oolites, and other insoluble sub¬

stances often furnish information of value (See Plate V, Figs.

1-3). Dolocastic chert has been reported frequently and chert

masses containing impressions of crystals other than those of

dolomite should be looked for. Small chert nodules in the May-

ville dolomite of Wisconsin contain calcitic crinoid fragments

and brachiopod shells, both of which are suggestive of the origin

of the chert. The detailed study of these nodules, both as to

faunal content and time of origin, promises some interesting

results. Much has been written concerning the significance of

glauconite in sedimentary rocks. Most of the Wisconsin dolo¬

mites contain some glauconite, and investigations of this ma¬

terial will be aided by insoluble residue analysis.

4. It is widely recognized that almost all sedimentary rocks

have undergone certain changes since deposition as sediments.

Most of these are so well known that they need not be men¬

tioned. A few, however, are cleared up considerably by light

thrown on them from studies of insoluble residues. European

geologists, and very recently Americans also, have reported a

number of authigenic minerals from sedimentary rocks. Quartz

Shrock — Insoluble Residues from Sedimentary Rocks. 259

and feldspar are the chief ones, and both have been found in

abundance in Wisconsin dolomites. Microcline from the Silurian

dolomites occurs as prismatic crystals with well developed

brachy- and macropinacoidal faces. The crystals are composed

of a shell of authigenic origin built around, and in crystallogra¬

phic continuity with, a nuclear detrital grain of microcline

(See Plate V, Figs. 11-12). Other kinds of authigenic feldspar

have been found in insoluble residues from Wisconsin rocks

( e.g . the Mendota dolomite), but they have not yet been defi¬

nitely identified. Their thorough study constitutes an impor¬

tant problem. The recovery and accurate identification of all

authigenic minerals in sedimentary rocks are needed to under¬

stand adequately one of the important changes that takes place

in a sedimentary rock after deposition (See Plate 5, Fig. 10).

Recent work by the writer on certain Wisconsin dolomites

shows that, when these rocks are treated for a short time with

a weak solution of hydrochloric acid, the crystals are not only

separated from each other but are also broken up into cleavage

fragments. This fragmentation is due to the fact that solution

proceeds most rapidly along the cleavage planes of the mineral.

Small rhombs of carbonate carried by certain spring waters

may have originated by the solution action just described. In¬

vestigations of the way various minerals behave during solu¬

tion might produce significant results.

5. Students of sedimentary rocks are familiar with the way

English, Scotch and French petrologists have utilized heavy

detrital minerals in determining provenances of sediments. Sim¬

ilar work has been initiated in America recently, and many

problems have been outlined and suggested. The presence of

detrital minerals in certain of the limestones on Anticosti island

suggests that it might be very much worth while to recover the

insoluble residues of all calcareous rocks lying on the periphery,

or upon the main mass, of the Canadian Shield, and study them

along with the heavy detrital minerals from the clastic rocks to

determine the relations between the sediments and the crystal¬

line terrane from which they presumably were derived. By util¬

izing exposures and well cuttings studies of the calcareous for¬

mations, as well as of the clastic ones, which lie around buried

or partly exhumed Wisconsin Pre-Cambrian hills might be pur¬

sued with significant results.

260 Wisconsin Academy of Sciences, Arts, and Letters .

6. Geologists have been able, by using chemical and strati¬

graphic data, to estimate roughly the relative percentages of

sandstone, shale and limestone in the world's sedimentary col¬

umn. Insoluble residue analysis, extended to include all types

of sedimentary rocks, might well alter those estimates some¬

what. In order to carry out such a program, however, it would

be necessary to sample all rocks in some uniform manner.

7. Many field geologists occasionally find themselves at a

loss to describe satisfactorily and accurately some sedimentary

rock. A chemical analysis showing soluble material, sand, silt

and clay would make this much easier and more definite. The

entire Wisconsin sedimentary column needs to be studied with

this in mind.

PART 2. STUDIES OF WISCONSIN SEDIMENTARY ROCKS

1. INSOLUBLE RESIDUES FROM WISCONSIN

SILURIAN DOLOMITES

George B. Burpee*

Purpose. — This investigation was initiated to determine

whether the lithologic units of the Wisconsin Silurian possessed

characteristic insoluble residues which might be of assistance in

correlating exposed formations with those encountered in wells

down the dip from the outcrop.

Procedure. — The samples used in this study were collected

by Dr. R. R. Shrock in eastern Wisconsin during the summers

of 1930 and 1931. They were analyzed in the Sedimentation

Laboratory at the University of Wisconsin with laboratory ma¬

terials furnished by the Milwaukee Public Museum.

A 20-40 gram sample was crushed into fragments about

one-half inch in greatest dimension, and then dissolved in a

50% solution of hydrochloric acid at a temperature slightly be¬

low boiling. It was necessary in some cases to wash the sample

and treat it with fresh acid several times before all of the solu¬

ble matter was dissolved. When all possible reaction had ceased

the acid was decanted and the residue washed clean of acid and

* Submitted as a thesis for the degree of Master of Arts (Geology) at the University of

Wisconsin, 1932. Abstracted by R. R. Shrock.

TRANS. WIS. ACAD., VOL. 29.

PLATE V

Plate V. 1. Thin section showing siliceous oolites in a calcareous matrix (xlO). 2. Same

as 1 , enlarged (x25); showing . several siliceous oolites under crossed nicols. Note the central core

of quartz. 3. Residue of siliceous oolites from same rock as 1 and 2 (xlS). 4. Siliceous

foraminifera (x8). 5. Detrital quartz grains in a clacareous matrix, shown in a thin section

(xlS). 6. Siliceous sponge spicules (x4). 7. Siliceous foraminifera (x8). 8. Several enlarged

(x20) specimens of 4. 9. Residue of detrital quartz grains from same rock as 5 (xlO). Note

rounding and frosting. 10. Authigenic pyrite crystals (xlO). 11. Authigenic microcline crystals

(x25). 12. An authigenic microcline crystal, greatly magnified (x200) and with crossed nicols,

showing the nuclear detrital grain around which the later authigenic shell grew (Photograph by

W. L. Wilgus). 13. Chitinous jaws (xlO).

Shrock— Insoluble Residues from Sedimentary Rocks. 261

clay material. From this procedure only a granular residue re¬

mained. This was then carefully washed onto a tared watch

glass, dried and weighed. It was then filed for future micro¬

scopic study. No attempt was made to determine the amount

of clayey matter in the residues.

Results. — Three general types of insoluble residues were

found. First, almost all of the samples analyzed contained a

small percentage (0.5% — 3%) of very fine to fine authigenic

quartz, and a second mineral then unidentified and thought also

to be quartz with a peculiar crystal habit, but now known to be

authigenic feldspar (microcline). The quartz consists of clear,

anhedral or euhedral prismatic crystals with rhombohedral

terminations. The feldspar usually occurs as euhedral prismatic

crystals with well developed brachy- and macropinacoidal faces.

They sometimes have a nucleus consisting of a detrital grain of

microcline. In these cases the authigenic shell of the crystal is

developed in crystallographic continuity with the nuclear grain.

Second, silicified fossils although far less abundant than the

authigenic minerals just described comprise the diagnostic frac¬

tion of the residues. In order of abundance these fossils are

sponge spicules, silicified foraminifera, fragments of brachiopod

shells, internal casts of ostracods and bryozoans, fragments of

crinoids and corals, and minute silicified gastropods or glauconi¬

tic casts of such shells. The foraminifera, and possibly the os¬

tracods, have stratigraphic significance. It might be added that

a residue composed chiefly of rounded detrital grains of quartz

was found in the basal beds of the Devonian, which immediately

overlie the Silurian at Gedarburg.

The Mayville dolomite is characterized by residues of chert

and hexactinellid sponge spicules. The amount of the residue

ranges from less than 1% to as much as 15%, but these percent¬

ages do not include the chert nodules which are abundant at

certain horizons throughout the formation. The Byron dolomite

rarely contains more than 2% of insoluble material, but has a

distinctive foraminiferal-ostracod fauna which has been found

at Burlington, Waukesha (in what may be the so-called Wauke¬

sha beds), and in well cuttings from a deep well near Racine.

The Coral beds as exposed in the vicinity of Valders contain

from 3% — 26% of insoluble material, consisting largely of small

crystals of quartz and feldspar (Burpee, without optical study,

262 Wisconsin Academy of Sciences , Arts , and Letters .

called all of the crystals quartz). The Racine beds, as developed

in the vicinity of Racine, contain only a very small residue (sel¬

dom over 1%), consisting of a little chert, some very fine anhe-

dral crystals of either quartz or feldspar and a few masses of

marcasite (?). A reef facies of the Racine (“Guelph”) at Ced-

arburg contains no appreciable amount of insoluble material,

but is overlaid unconformably by Devonian strata which carry

a distinctive residue of rounded detrital quartz grains. The

Waubakee dolomite, youngest of the Silurian formations of Wis¬

consin and known only from a few scattered exposures in the

vicinity of Milwaukee, contains no appreciable residue.

In summary it may be stated that the Silurian dolomites of

Wisconsin carry a diversity of insoluble material, usually in

small amount; but that with the exception of certain silicified

fossils, the residues so far obtained and studied do not have

stratigraphic significance. The foraminifera, and possibly the

ostracods, seem to be limited to a narrow stratigraphic range in

the Byron formation, and may with further investigations be¬

come a useful horizon marker.

2. THE INSOLUBLE RESIDUES OF THE ONE OTA

DOLOMITE OF WESTERN WISCONSIN.

Joseph J. Drindak*

Purpose . — This study was undertaken to ascertain the

amount and character of the insoluble material in the Oneota

dolomite, and in the immediately underlying arenaceous strata,

in western Wisconsin; and to determine whether that insoluble

material might be of stratigraphic or economic importance.

Procedure. — Samples were collected in the summer of 1982

with the aid of E. H. Powell and R. R. Shrock. They were taken

in vertical sections about five feet apart unless there was a dis¬

tinct change in lithology, in which case a sample was taken

where the change occurred, and then the same procedure as be¬

fore was followed. In one section samples were taken every six

inches in the lower part and every foot in the upper part. The

analytical work was carried on in the Sedimentation Labora-

* Submitted as a thesis for the degree of Bachelor of Philosophy (Geology) at the University

of Wisconsin, 1933. Abstracted by R. R. Shrock.

Shrock — Insoluble Residues from Sedimentary Rocks . 263

tory at the University of Wisconsin, with laboratory materials

furnished by the Milwaukee Public Museum.

About a 20-gram sample was crushed into small fragments

averaging about one-half inch in greatest dimension, wetted

with distilled water and then dissolved in a 50% solution of hy¬

drochloric acid. In some instances the sample had to be washed

and treated with new acid several times before all of the soluble

rock was dissolved. After solution had ceased completely the

acid was decanted, and the remaining insoluble residue was

washed until all of the finely divided silt and clay had been re¬

moved and only a granular residue remained. This was then

washed onto a tared watch glass, dried, weighed, and studied.

Results.— The insoluble constituents of the Oneota dolomite

show considerable diversity. (1) Silicified corals and sponge

spicules, mainly monaxons, are present throughout the forma¬

tion in the sections studied, but do not seem to have stratigra¬

phic significance. (2) Quartz is abundantly represented in the

lower part of the formation by rounded and frosted detrital

grains, and throughout the formation by minute crystals and

clusters of crystals. (3) Siliceous oolites are usually present in

the oolitic beds of the formation, and may show concentric

structure. (4) Glauconite is abundant in certain beds (“Green

speckled beds”)* and is common throughout the formation in

small amount. Dolocastic chert was encountered in some beds

and mica flakes are not uncommon in the arenaceous strata

immediately underlying the base of the formation.

On the basis of lithology and field relations, and to a very

limited extent on the basis of insoluble residue content, it is

possible to subdivide the Oneota dolomite into several fairly

distinct zones, as shown in Fig. 1. These have interesting,

though rarely distinctive, residues.

Zone 1 belongs below the Oneota dolomite and is probably

to be correlated with the Madison sandstone of the central and

eastern parts of Wisconsin. It bears such a close relation to the

basal part of the Oneota, however, that it was thought advis¬

able to study its insoluble content along with that of the over-

* The basal part of the Oneota dolomite is characterized by thin beds of bluish-gray dolomite

speckled or spotted with roughly spheroidal masses of greenish, or when weathered yellowish or

slightly bluish, clay-like matter composed in large part of glauconite. These beds are specially

designated in the sections of Fig. 1.

264 Wisconsin Academy of Sciences, Arts, and Letters ,

Fig. 1. Chart of stratigraphic sections showing the relative amounts of soluble and insoluble

material at various horizons in the Oneota dolomite of western Wisconsin. The Madison-Oneota

boundary is uncertain in some sections, hence it is not indicated.

Shrock — Insoluble Residues from Sedimentary Rocks . 265

lying formation. The base of zone 1 was taken at the top of the

Jordan sandstone where the first strong dolomite influence ap¬

peared. The top of the zone, which has been reported as an un¬

conformity, could not always be determined, hence no definite

contact between the top of the Madison and the base of the

Oneota is indicated in the sections in Fig. 1. Sand lenses and

layers, dolomitically-cemented arenaceous beds, shale layers and

conglomerates characterize the zone, but neither their presence

nor number is consistent. The residues are composed mainly of

rounded and frosted detrital grains of quartz in the sandstone

layers, and both detrital and authigenic quartz in the dolomitic

beds. Chert, glauconite and mica flakes were also obtained from

some of the residues, but usually in small amount. The insoluble

material tends to decrease upward in the zone.

Zone 2i comprises the basal part of the Oneota dolomite and

is characterized by a diversity of lithology. The “green speckled

beds” are always prominent, but there are also oolitic beds,

shale layers, algal layers and conglomerates, all of which vary

in thickness and number. All are highly dolomitic and contain

small percentages of insoluble material. The detrital quartz of

zone 1 gives way upward to authigenic quartz pretty largely,

which occurs in the form of single crystals or crystal clusters.

Chert is found in the algal layers, while the oolitic beds yield

siliceous oolites. Glauconite is found especially abundant in the

“green speckled beds” and may make up as much as 30% of the

residues. Sponge spicules and casts of cup corals occur sparsely

throughout the zone.

Zone 3 is characterized by a well developed algal biostrome.

The insoluble material is present in small amount, and consists

of authigenic quartz, chert masses, siliceous oolites and rare

sponge spicules. Large chert masses may be seen in this zone in

the field but they are not considered in percentages of insoluble

material, for in the collection of samples every effort was made

to obtain dolomite free of visible chert.

Zone 4 is an interval of thin-bedded dolomites, designated as

“punky beds”* in the field, alternating with thicker strata of the

same general lithology. The insoluble content is quite small, and

is composed mainly of fine authigenic quartz, small chert masses,

* These fine-grained, thin-bedded dolomites have been designated in the field as the “punky

beds”, because when struck with a hammer they give a thud.

266 Wisconsin Academy of Sciences, Arts, and Letters .

siliceous oolites, dolocastic chert, sponge spicules and glauconite

grains.

Zone 5 is characterized by thick, massive beds of dolomite,

somewhat cherty in places and often containing a discontinuous

algal biostrome or a conglomerate horizon. Authigenic quartz

crystals, both singly and in clusters, are common in this zone

along with spongy chert. Chert and siliceous cement increase

upward in some sections. The large masses of chert are not in¬

cluded in the residue percentages.

Zone 6 represents a sequence of arenaceous strata above the

Oneota dolomite and probably correlates with either the Neiv

Richmond or St. Peter sandstone. The insoluble residue which

comprises nearly the entire rock consists of siliceous cementing

material, detrital quartz grains and a few siliceous oolites.

In summary it may be said that this study of the Oneota dolo¬

mite has shown that underlying and overlying formations (Madi¬

son sandstone below and New Richmond or St. Peter sandstone

above) can be sharply differentiated from the Oneota on the

basis of the amount and character of the residues ; but that it is

not possible to identify definite zones or horizons within the dolo¬

mite formation with any degree of success.

3. A SEDIMENT ATIONAL STUDY OF A PART OF THE

TREMPEALEAU FORMATION IN SOUTHERN

WISCONSIN .

Bernhard 0. Hougen .*

Purpose. — This study was made to determine: (1) the per¬

centages of soluble (calcareous) and insoluble (sand, silt and

clay) materials in selected exposures of the Trempealeau for¬

mation in southern Wisconsin; and (2) whether the insoluble

constituents possessed characteristics which might be used either

for interpreting conditions of sedimentation at the time of depo¬

sition or for correlation purposes.

Procedure. — Samples were collected, with the aid of G. 0.

Raasch, from four exposures near Avoca, Muscoda, Gotham and

Kingston, along or near the Wisconsin river valley (See Fig. 1).

* Submitted as a thesis for the degree of Bachelor of Philosophy (Geology) at the University

of Wisconsin, 1933. Abstracted by R. R. Shrock.

Shrock— Insoluble Residues from Sedimentary Rocks. 267

Hand speciments (about 2"x3"x3") were taken every three feet

throughout the section unless individual strata were under that

figure in thickness, in which case several small fragments were

taken from each of the thin beds in the three-foot zone.

After a microscopic examination of the sample was made

(both before and after crushing), about 20 grams of the crushed

rock (fragments about % inch in greatest dimension) were di¬

gested in a 50% solution of hydrochloric acid at a temperature

slightly below boiling for twenty four hours. The sample was

then washed and treated with new acid. This was continued

until all solution action ceased. The acid was then decanted and

the residue washed free from acid with distilled water. The

residue was then dried and weighed, after which it was moist¬

ened and poured into a specially devised water classifier, where

the clay content was separated from the sand-silt fraction.* The

latter fraction, saved in the process of separation, was then

dried and weighed. Percentages of soluble material, sand-silt

material and clay were then computed. It must be emphasized

that while the methods used did not give highly accurate results,

nevertheless, they are quite satisfactory for the purposes in¬

tended because of the variation of the formation both vertically

and horizontally.

Results.— The insoluble residues from the Trempealeau for¬

mation in the four sections studied consist mainly of clay; silt;

rounded and angular quartz grains ; authigenic quartz, and pos¬

sibly feldspar, crystals ; and glauconite. The soluble material is

almost entirely calcite and dolomite (relative percentages were

not determined), with a very small amount of iron oxide.

The results of the analyses are shown on Fig. 1. The St. Law¬

rence dolomite always is impure carrying as much as 36% of in¬

soluble material. The Lodi shale carries a surprising amount of

soluble matter, in some cases well over 56%. It is also apparent

that the clay and sand-silt material may or may not be fairly

evenly balanced. The Jordan sandstone was found to contain a

considerable amount of calcareous material in most of the samp-

es. This situation is not always apparent in outcrops. The main

importance of the insoluble residue data is to give a more accu-

* Considerable experimentation was necessary before the proper procedure and apparatus were

discovered. Finally, however, it was possible to remove the clay (particles less than %gg mm.

in largest dimension) rather, successfully.

268 Wisconsin Academy of Sciences, Arts, and Letters .

rate picture of rock composition than has been available before.

In themselves the residues do not possess characteristics which

are distinctive, and hence are of little value for correlation pur¬

poses.

4. INSOLUBLE RESIDUES OF THE MENDOTA

(ST. LAWRENCE) DOLOMITE.

Ray E. Wilcox A

Purpose. — This investigation was undertaken to find the

amount, character and possible uses of any insoluble materials

that might be present in the Mendota dolomite of the Madison

* Submitted as a thesis for the degree of Bachelor of Philosophy (Geology) at the University

of Wisconsin, 1933. Abstracted by R. R. Shrock.

Shrock— Insoluble Residues from Sedimentary Rocks. 269

vicinity. Correlation of the Mendota with the St. Lawrence dolo¬

mite follows the practice of Wisconsin geologists.

Procedure. — Samples were collected with the aid of Mr. F.

T. Thwaites. Some well cuttings were also obtained from him.

The analyses were made in the Sedimentation at the University

of Wisconsin, with laboratory materials furnished by the Mil¬

waukee Public Museum.

From 25 to 50 grams of the sample were crushed to frag¬

ments averaging about 0.3 cms. in greatest dimension, placed in

a beaker and wetted with distilled water, and then dissolved in

a 50% solution of hydrochloric acid at a temperature slightly

below boiling. It was in some cases necessary to wash the sam¬

ples and then add new acid several times before all of the solu¬

ble matter was dissolved. After all action had ceased the acid

was decanted and the residue washed free of acid. Care was

taken not to pour off any of the residue. It was then washed on¬

to a tared watch glass, dried, and weighed. It was then returned

to a beaker, wetted and the fine clayey material removed by

careful decantation. The result was a clean, granular residue

whose particles were above clay dimensions. This residue was

then washed onto a tared watch glass, dried and weighed. The

method just outlined gave a reasonably satisfactory separation

of the sand-silt and clay fractions of the original residue.

Results. — The results of the investigation are tabulated on

Fig. 1. The insoluble residues consisted of the following: (1)

white, green or brown clay of about the same amount in most

of the samples; (2) rounded and frosted detrital grains of

quartz, scattered throughout the formation, and showing inci¬

pient secondary enlargement; (3) authigenic feldspar crystals

consisting of a shell of feldspar of authigenic origin around a

nuclear detrital grain of microcline; (4) fairly large flakes of

detrital muscovite; (5) fine to coarse, dark to light green,

rounded and polished grains of glauconite; and (6) irregular

masses of soft, argillaceous, cinder-like material containing fer¬

ruginous matter.

Three significant facts are apparent from the data tabulated

on Fig. 1. (1) The basal part of the Mendota dolomite is marked

by a thin conglomerate, which is composed of sand and clay, and

270 Wisconsin Academy of Sciences , Arts , and Letters .

dolomite. The analyses show that the insoluble material is high

(25% to over 50%). It consists of rounded and frosted grains

of quartz, some of which show secondary enlargement ; polished

glauconite grains; mica flakes; and some silt and clay. (2) The

algal layers or biostromes are conspicuous by their very low in¬

soluble content. Obviously the water in which the algae grew

must have been free of muddy matter. (3) The strata in the

Pheasant Branch section, and the upper beds of the Farwells

Point section, carry considerable insoluble material, but it is to

be noted that algal layers are absent. Perhaps the high residues

furnish the explanation for the absence of the algae.

While the amount and character of the insoluble residues of

the Mendota formation are distinctly different from those of the

underlying Franconia or of the overlying Lodi shale, it will be

necessary to obtain much more information on all of the residues

before they can be used successfully for correlation purposes.

The best procedure will be to use the residues in conjunction

with the lithologic data and field relations.

Shrock — Insoluble Residues from Sedimentary Rocks . 271

Fig. 1. Chart of stratigraphic sections showing the amount and character of the insoluble

residues of the Mendota dolomite. The heavy horizontal line marks the top of the conglomerate

which characterizes the base of the dolomite.

■Sdd&i

PRELIMINARY LIST OF THE HYDRACARINA

OF WISCONSIN

PART IV

Ruth Marshall

Parts I, II and III of the Preliminary List of the Hydraca -

rina of Wisconsin recorded fifty-three species belonging to eigh¬

teen genera. The present paper adds fourteen species represent¬

ing four genera of the family Hygrobatidae. None of these

are new but additional data are given for several of them. As in

the preceding Parts, drawings and some of the outstanding

characters of each species are given, together with distribution

data as far as known. For complete descriptions of the species,

the student of the group is referred to the titles in the biblio¬

graphy.

The preparation of this work has been greatly aided by a

grant from the Society of Sigma Xi. For most of the material

the author is indebted to the Wisconsin Natural History Survey.

Following the system of Viets, the Koenikeas are included

with the Unionicolas and the Neumanias in the subfamily Uni-

onicolinae; superficially they resemble the Mideopsinae. The

genus Koenikea was set up by Wolcott (1900) with one species,

K. concava ; in reality, two species were here included, as was

noted by other investigators and pointed out by Koenike and

Lundblad. Dr. Wolcott also erected another genus, Tanaogna-

thus, with one species, T. spinipes, now regarded as a Koenikea.

It was evident that other species were also present and this led

to further confusion in identifications, of which the present au¬

thor was also guilty. Viets (1930), after the examination of a

small amount of North American material, reviewed the litera¬

ture and established three new species and more clearly defined

the original two. These five species are all present in Wisconsin ;

the present paper adds data on the differentiation of the sexes.

The Koenikeas are small mites, usually less than one millimeter

in length ; the body is greatly compressed and concave dorsally ;

the skin is heavily chitinized and pierced with fine pores and the

273

274 Wisconsin Academy of Sciences , Arts , and Letters .

dorsal exoskeleton shows a furrow near the margin which thus

encloses a large circular area. The genus appears to be most

abundant in the New World.

The Pionas form one of the largest genera of the water mites,

both in numbers of species and in individuals. They are soft

bodied, with striated skin, often brightly colored, usually ellip¬

tical in form and of medium size, although a few attain a length

of two millimeters. The shape of the large fourth spimera, each

with a prominent posterior angle, together with the presence

of two or more hair papillae on the fourth segment of the pal¬

pus, will usually serve to identify the genus. The genital open¬

ings lie close to the epimera; sexual dimorphism is marked,

shown especially in the peculiarities of the third and fourth

legs in the male, the latter always showing a large concavity in

the fourth segment. This paper lists only six of the several spe¬

cies of Pionas in the state ; it is hoped to include the remainder

in the next Part.

The Forelias, a small group, closely resemble the Pionas from

which they are readily distinguished by the shape of the fourth

epimera; in this plate the anterior and posterior margins meet

in the median line to form an acute angle. The Huitfeldtias are

rare ; they resemble the female Pionas in the form of the epimera

but lack the other distinctive features of that genus.

Koenike concava Wol.

PL VI, fig. 3-5

The original account of the species (Wolcott, 1900) gave the

male and female; both forms described, however, were obviously

females. Dr. Wolcott deposited co-types in the National Museum

and these have been examined by the author. Little can be de¬

termined from them, since the preservation is poor, but it is

evident that two species are present. Dr. Viets (1930) desig¬

nated Wolcott’s male (in reality a female) as K. concava and

called the other female K. ivolcotti n. nom. The true male is now

known and here described. The body is nearly circular in the

male and measures from 0.60 to 0.69 mm. in length. The females

are more elongated and vary in length from 0.70 to 0.93 mm, de¬

pending upon the age. A lyre-shaped dark marking is seen on

the dorsal side, over which is a broadly U-shaped red mass. (The

mark shown in Wolcott’s fig. 15 was not observed.) The epi-

Marshall — Hydracarina of Wisconsin

275

mera are united in one plate, their places of union marked by

heavy lines, The genital plates are large and close to the fourth

epimera ; their small and scattered acetabula, set into the body

wall, form indistinct wing-shaped areas, narrow in the male.

The legs are weak. The rostrum of the maxillary organ is a little

prolonged and is seen as a rounded body in ventral view. In the

palpus the second segment is nearly as wide as the first leg ; the

fourth segment is distinguished by the presence of a hair-bear¬

ing papilla near the distal end.

Specimens have been found in Michigan, Indiana, Illinois,

Iowa, Florida and Louisiana. In Wisconsin they have been found

in ponds in Adams county and in lakes Wingra and Green, in

the latter to a depth of six meters.

Koenikea haldemani Viets

PI. VI, fig. 1, 2

A species closely related to K . concava, with which it has

been confused, the female was recognized and separated from

it by Viets (1930). The male is now known. The body in both

sexes is nearly circular. Males measure from 0.525 to 0.575 mm.

in length; females, 0.62 to 0.72 mm. The center of the body

shows a large magenta or violet blotch. The epimera are fused,

the divisions between them shown by heavy lines;, the fourth

pair are wider than in K. concava . The genital regions are at

some distance from the epimera ; the genital cleft and plates are

large in the female, much smaller in the male. The scattered

and indistinct acetabula cover a narrow and widely extended

area. The second segment of the palpus is wider than the legs,

which are weak; the fourth segment lacks a papilla. The ros¬

trum is but little projected.

Specimens have been found in ponds in Adams county, in

Green lake and lakes in Iowa.

Koenikea wolcotti Viets

PI. VI, fig. 6, 7

First described by Wolcott (1900) as the female of K. con¬

cava, it was separated as a distinct species by Viets (1930), the

male of which is still unknown. The body of the female is nearly

circular, 0.75 to 0.90 mm. in length. The body covering is very

heavy ; blotches of red show anteriorly and posteriorly. The epi-

276 Wisconsin Academy of Sciences , Arts, and Letters .

meral plates join except for a considerable space between the

third and fourth of either side; the first pair have projecting

rounded anterior ends. A distinctive feature of this species is

the large curved snout or rostral spine, with two long curved

bristles near its base. The palpi are more slender than the legs

and resemble those of K, haldemani . The fourth pair of legs

bear several scattered pectinated spines. Genital acetabula are

few, scattered and indistinct ; they lie in the body wall in a nar¬

row area extending outward from the large genital plates.

Individuals were collected in lake Wingra; they have also

been found in Michigan, Indiana and Illinois.

Koenikea marshallae Viets

PI. VII, fig. 8-11

The male is from 0.55 to 0.65 mm. in length, the width being

only slightly less; the anterior region projects conspicuously

and there is a protuberance over each eye. The female is from

0.675 to 0.775 mm. in length, relatively slimmer than the male,

with anterior projections not so large. Blue green and red blot¬

ches are conspicuous on both surfaces. The epimeral groups

are separated, with a wide space between the third and fourth

of each side in the female; the first pair have projecting rounded

anterior ends. The genital clefts are long and near the last epi-

mera; the plates are very large in the female. The genital ace¬

tabula are irregular and indistinct and form broad wing-shaped

areas. The rostrum of the maxillary organ is greatly drawn out

and prolonged into a very conspicuous slim sickle-shaped spine;

this bears tw~o large hairs at its base. The palpi bear numerous

hairs but no papillae. The legs are weak, scarcely wider than

the palpi.

Material has been found in Ontario, Michigan, Illinois and

Louisiana ; in Wisconsin in ponds in Adams county, in Spooner,

Allequash, Twin and Green lakes, in the latter to a depth of six

meters.

Koenikea spinipes (Wol.)

PI. VII, fig. 12-15

Originally described and set aside in a new genus, Tanaogna -

thus , by Wolcott (1900) from the study of one specimen, a male,

this species was placed with the Koenikeas by Viets (1930).

Marshall — Hydracarina of Wisconsin 277

While this is probably its place, nevertheless the peculiarities of

the palpi and legs set it apart from other described species. Ink

the character of the rostral snout it resembles K. wolcotti . The

female is now known. The male is from 0.625 to 0.70 mm. in

length; the female, 0.90 to 1.10 mm. The body is nearly circular

in form with an anterior projection, or slightly wider than long

in the male. The skin is very thick and blue blotches show on

this in life. The epimera in the male come close together except

for a space in the center of the field ; in the female the third and

fourth of each side are separated from the others by a small

space. The genital areas lie close to the last epimera ; in the fe¬

male the cleft is long and the plates are large, while in the male

the reverse is true. The genital acetabula lie in the body wall

in an area which is broad but not laterally extensive. The ros¬

trum of the maxillary organ is drawn out to form a heavy

greatly curved snout, broad at the base, with a conical tip ; dor¬

sal to this are two long heavy curved hairs. The palpus is very

small; the basal segment is the broadest, while the second is

longer than succeeding ones; the fourth segment has a large

projection on its distal end projecting parallel to the fifth. The

legs are rather stout and bear pectinate hairs ; the fifth segment

in each is enlarged in the center, while the sixth is slightly cur¬

ved. The first leg ends in an expanded tip which bears a pair of

large sickle-shaped claws.

Specimens were collected in lakes Wingra and Allequash.

They have also been found in Michigan and Illinois.

Huitfeldtia rectipes Thor

PI. VIII, fig. 16-18

An uncommon species, the only one in the genus, a few speci¬

mens, all males, have been found in northern lakes. The body is

oval, thin-skinned, with brown blotches. Males measure up to

1.25 mm. in length; females are a little longer. The epimera

resemble those of the female Pionas but are smaller and the

groups are well separated. The genital areas are removed from

the epimera and have lunate plates, united at the ends in the

male, which bear a small number of conspicuous acetabula. Palpi

are slender ; the third segment bears a long hair and the fourth

exceeds the others in length.

278 Wisconsin Academy of Sciences , Arts , cmd Letters .

The species has been reported from northern Europe and

Germany. It has been found in Ontario, Saskatchewan and in

four lakes in Vilas county. In two collections individuals were

found at a depth of about fourteen meters.

Forelia liliacea (Mull.)

PI. VIII, fig. 22-25

This cosmopolitan species measures in the largest males

nearly 0.70 mm., and in the females up to 0.80 mm. The poster¬

ior end of the body is elongated, especially in the male ; the gene¬

ral color is brown. The epimeral groups are close together in

the male, well separated in the female; the genital area lies al¬

most entirely within the bay formed by the large triangular last

pair. In the female the genital plates, well removed from the

long aperature, are broad and bear from fifteen to thirty aceta-

bula each; in the male the plates are narrower, joined at either

end of the short aperature, and lie close to the last epimera,

scarcely extending beyond their sharp posterior angles. The

palpi are stouter than the first pair of legs. The fourth pair of

legs are a little longer than the body ; in the male the sixth seg¬

ment is bent strongly dorsally and bears ten or more blunt spines

on the inner surface.

The species is found throughout Europe and is reported for

Africa. It has been found in Ontario, Wyoming, Washington

and Michigan. In Wisconsin collections have been made in

Spooner and Bass (Waupaca) lakes and in several lakes in Vilas

county, in one case to a depth of eight meters.

Forelia ovalis Mar.

PL VIII, fig. 19-21

Closely related to F. liliacea, this species is a little larger with

stouter legs and shows a red trident-shaped dorsal mark. The

genital plates extend laterally beyond the posterior blunter angles

of the fourth epimera and bear many small acetabula; in the

male the plates meet for a short space below the short genital

opening. The fourth leg in the male is a little shorter than the

body and all of the segments are stout ; the sixth segment, lying

nearly in the same plane as the fifth, has a broad proximal part

Marshall — H ydracarina of Wisconsin

279

which is extended into a finger-shaped process bearing a row of

short blade-like bristles.

Collections have been made in Ontario and Illinois. In Wis¬

consin the species has been found in the Madison and Jordan

lake regions, in Green lake and in eight lakes in Vilas county, in

one case to a depth of thirteen meters.

Fiona crassa (Wol.)

PI. X, fig. 37-42

This is an unusual Piona, being heavily built and chitinized.

The entire dorsal surface of the male is thinly chitinized and

somewhat flattened, while the female has a small anterior dor¬

sal plate, characters not given in the original description of the

species. The remainder of the cuticula is thick, with heavy rid¬

ges. Living specimens show the usual brown blotches on the dor¬

sal side: a faint pink shows in the center of these, while the

edges of the ventral plates are tinged with red and the append¬

ages are deep blue. Males are about 0.65 mm. in length; fe¬

males, from 0.80 to 1.00 mm. The posterior angle of the fourth

epimera is especially sharp; all of the plates are united in the

male. The conspicuous genital tongue-shaped plates, bearing a

large number of acetabula, are strongly curved and extend far

out beyond the lateral boundaries of the epimera. In the male

these plates meet medially below the small opening to form an

oblong area devoid of acetabula to which the anal plate is fused.

The short stout flattened palpi are very unusual in that the

fourth segment has eight small papillae arranged in two rows

on the projecting distal margin. The author has not observed

any marked difference in the palpi in the two sexes; Wolcott

(1902, fig. 60) in a description based upon a single individual

gives a much slimmer palpus for the female. The legs are stout,

with short terminal segments in the first three pairs. The third

leg has a truncated sixth segment, while the long fifth segment

bears distally several heavy spines and one very long hair, the

last character not given in the original description. The fourth

leg has unusually stout proximal segments with the character¬

istic concavity on the fourth.

Specimens have been found in Montana, Michigan, Illinois

and in Wisconsin in lake Wingra and in three regions in Adams

and Vilas counties.

280 Wisconsin Academy of Sciences , Arts , and Letters .

Piowa rotunda (Kram.)

PL IX, fig. 26-81

A cosmopolitan species and widely distributed, its variability

and the lack of very distinctive characters make the identifica¬

tion of P. rotunda difficult, especially in the case of the female.

Confusion in this matter is evident in some of the older litera¬

ture. The body is broadly elliptical and shows brown blotches

under a finely ridged surface. Largest males measure about 0.80

mm., females, 1.10 mm. The genital areas in both sexes extend

laterally hardly beyond the posterior angles of the moderately

concave borders of the fourth epimera. The fused genital plates

of the male show a broad concavity on the posterior border;

there are from, fifteen to thirty acetabula on either side, with a

large shallow free area under the genital opening. The anal

plate lies well back of the genital plates. In the female there

are about the same number of acetabula placed on a sickle¬

shaped plate on either side, arranged in an irregular crowded

row, with a few others embedded in the wall on the concave side.

The palpi are wider than the first pair of legs, with a slim fourth

segment bearing two small papillae. The sixth segment of the

third leg is about half of the length of the fifth, a little expanded

at the end ; one of its claws bears a long straight tip.

This species has been found all over Europe and in parts of

Asia and Africa. It has been collected in Alaska, British Colum¬

bia, Ontario, Montana, Michigan and Nebraska. In Wisconsin

there are records for lakes Wingra, Coma, Mason, ponds near

Cable, Wisconsin Dells, Green Bay and waters of Adams and

Vilas counties.

Fiona reighardi (Wol.)

PI. IX, fig. 32-36

The females of this species closely resemble P. rotunda ,

from which it is separated with difficulty, since both species

show great variability in the genital plates. Co-types of both

species, deposited in the National Museum by Dr. Wolcott, and

examined by the author, are not well preserved ; there appears,

however, to have been some confusion of the two forms. Living

material, believed by the author to be the true P. reighardi , us¬

ually shows a bright red spot ventrally at the meeting of the

Marshall — Hydracarina of Wisconsin

281

epimeral groups ; dorsally there is a yellowish or white T-shaped

mark surrounded by dark brown blotches ; the eyes are dark red

and the appendages deep blue. Ventral plates are rather heavy,

often blue tinged. The body is broadly oval with little or no in¬

dentation anteriorly; the average length is about 0.90 mm. The

fourth epimera show a greater concavity on the posterior border

than in the related species, and the sickle-shaped genital plates

(rarely broken) are more elongated and may extend a little far¬

ther laterally than the posterior angle of the epimera. The geni¬

tal acetabula are highly variable in number; they are usually

larger and more irregularly arranged than in the related species,

and bunched at both ends of the plates, while in the body wall

are from two to four more on either side. The palpi are much

like those of P. rotunda , but the second segment is a little stouter.

The original description of the species (Wolcott, 1902, p. 285)

gave an incomplete and inaccurate description of the male, since

only one specimen, poorly preserved, was known. Males in the

present collection are abundant; they are often much smaller

than the females, averaging about 0.63 mm. in length, and show¬

ing the same coloration. Small spaces separate the epimeral

groups, which cover the greater part of the ventral surface. The

united genital plates are much like those of P. rotunda, but they

extend laterally beyond the angle of the fourth epimera (rather

than 4 ‘about even with the tip of the process in the posterior

margin”), and the genital opening extends through about half

of the length of the plate (instead of “throughout the entire

length of this genital area”). There are from twenty to twenty-

five acetabula on each side. The posterior concavity of the united

plates is deeper than in the related species, with the anal plate

well within it. The third leg has a shorter terminal segment,

with a shorter claw, and the fourth segment of the last leg is

likewise shorter and stouter bristles than in P. rotunda .

Dr. Wolcott in the same paper (p. 240) identified as P. ob~

turbans (Piers.) two females in a collection from Louisiana. In

the author's opinion these were probably P. reighardi, since the

females of the two species are much alike and the specimens were

believed to have shown some red color when alive.

This species appears to be the most abundant Fiona in cen¬

tral North America. It has been found in Ontario, Michigan,

Minnesota, Montana, Indiana, Illinois, Iowa, Louisiana and

282 Wisconsin Academy of Sciences , Arts , and Letters .

Georgia. In Wisconsin it has been collected in nearly all regions

visited, often in large numbers, from the surface to a depth of

12 M. The author’s record for Alaska is probably wrong; in a

re-examination the specimens (females) appear to be P. ro¬

tunda. Likewise the record of the Canadian Arctic Expedition

(1913-18, v. Ill: 13H) is in doubt; some of the specimens, de¬

posited in Ottowa, have been examined by the author.

Piona media (Wol.)

PI. XI, fig. 49-53

Described originally from one preserved female, this species,

including the male, can now be fully characterized. The body is

broadly oval ; the dorsal side shows a yellowish T-shaped figure

surrounded by brown blotches on a pale yellow or bluish back¬

ground, while the plates and appendages are blue. The eyes are

very large and the antenniform bristles are small. Males average

about 0.57 mm. in length, females nearly 1.00 mm. Plates and

legs are heavy. The posterior margin of the fourth epimera

shows only a moderate concavity. The genital areas resemble

those of P. rotunda and P. reighardi but they are of greater ex¬

tent and carry more acetabula. In the female the broadly sickle¬

shaped plates extend laterally beyond the posterior angle of the

fourth epimera and carry each some thirty-five or more aceta¬

bula arranged in two irregular rows with some bunching, to¬

gether with a few more embedded in the wall. In the male the

tongue-shaped plates reach laterally to the margin of the body

and bear each about thirty-five acetabula; the genital opening

is large and the margin of the united genital plates below it

shows a deep concavity in which lies the anal plate. The palpi

are unusually large, especially in the male; the second segment

is nearly twice the width of the first leg and the fourth segment

bears two large slim papillae nearly opposite each other. The

pectination on the spines of the palpi described by Wolcott was

not observed by the author. The third and fourth legs of the

male resemble those of P. reighardi.

Specimens have been found in Ontario, Michigan, Illinois and

Indiana. In Wisconsin collections have been made in lakes Win¬

nebago, Starr, Wingra, Twin, Little John, in ponds near Bur¬

lington and in five bodies of water in Adams county.

Marshall — Hydracarina of Wisconsin 283

Dr. Walter* has described a single female found in a Swiss

lake which bears a close resemblance to P. media ; its identifica¬

tion is in doubt.

Piona inconstans (Wol.)

PL XI, fig. 54-57

The body is elliptical, covered with fine wavey lines, often

showing a faint red T-shaped dorsal mark in the center of the

dark brown blotches. Females are from 0.75 to 1.10 mm. in

length, males a little smaller. The females closely resemble P.

rotunda and P. reighardi, from which they are distinguished by

peculiarities of the genital plate. Here the ten to twenty aceta-

bula of each side lie on a broken sickle-shaped plate, a small

anterior piece with a few acetabula and hairs and a posterior

piece (sometimes again broken) carrying the others in an irreg¬

ular row, while two or more lie free in the body wall or on small

plates. Great variability is seen in this matter; even the two

sides of the same individuals are unlike. The genital plates do

not extend laterally farther than the posterior angles of the last

epimera. The palpi are slim, much like those of the related spe¬

cies.

The male, not included in the original description of the

species, is now known. It is relatively large, measuring about

0.90 mm. The epimeral groups are separated by unusually wide

spaces for males of this group of Pionas, and they occupy only

about two-thirds of the ventral surface. The genital plates are

small, in form like those of P. rotunda, with a small and vari¬

able number of acetabula, about fifteen on each side, and they

do not extend laterally quite as far as the posterior angle of the

last epimera. The sixth segment of the third leg is broadened

near the proximal end and again distally where it ends in two

curved claws, one of them very long. The fourth segment of

the fourth leg has a large concavity and the proximal border

bears a bunch of long slim bristles.

Specimens have been found in Alaska, Ontario, Maine, Mas¬

sachusetts, Michigan, Illinois, Missouri, Nebraska, Louisiana

and Florida. In Wisconsin they have been collected in lakes

Winnebago, Mendota, Mason and Twin and in ponds near Wis¬

consin Dells.

Die Hydracarinen der Schweiz. Rev. Suis de Zool., v. IS, 1907, p. S33-S, fig. 40, 41.

284 Wisconsin Academy of Sciences , Arts, and Letters.

Fiona conglobata (Koch)

PL X, fig. 43-47

This cosmopolitan species has already been reported from

Wisconsin. The females collected conform closely to identified

material; unfortunately the author has no specimens with

which to compare the one male so far found and published ac¬

counts are not in agreement as to certain details of the genital

plates. However, as the palpi and details of other parts of both

sexes conform, it is believed that these individuals represent the

true P. conglobata. The body is oval, the surface coarsely ridged.

Antenniform bristles are long. The dorsal surface shows brown

blotches on a dull yellow background with sometimes faint red

showing in the center. Largest females measure 0.90 mm. in

length, the single male, 0.45 mm. The epimera are well sep¬

arated in the female, closely approximated in the male; the

fourth pair show a moderate posterior medial concavity. The

genital areas in both sexes are extensive and extend laterally

beyond the posterior angle of the last epimera. Acetabula are

few, scattered and very irregularly placed; in the female these

are embedded directly in the body wall except for a few on two

small plates near either end of the long genital opening. In the

male the united genital plates are not very clearly outlined;

there is a deep concavity where they meet and the anal plate

does not join them. The genital opening is small and the mark¬

ings near it are complex and difficult to make out. The palpi

are large, a little stouter in the male than in the female; the

fourth segment bears three large hair papillae in the center of

the flexor surface. In the male the first two pairs of legs have

the terminal segment much thickened distally; the last segment

of the third leg is moderately long and has a reduced claw, while

the fourth leg has a very large concavity on the fourth segment.

The species is found over all of Europe and there are records

for Turkestan and Mongolia. In Wisconsin material has been

found in a mill pond at Oxford and in lakes Spooner and Little

John. Rockford College,

Sept. 1, 1933

* Since the completion of this manuscript the author has received identified specimens from

Germany through the kindness of Dr. K. Viets. It is now clear that the American form is not

identical with the European; consequently it will be designated as P. conglobata wisconsinensis nov.

var. It is distinguished from the parent species chiefly by the presence of a greater number of

genital acetabula and by the form of the male genital plates; the latter show an upturned margin

where they meet the posterior angle of the fourth epimera.

Marshall — Hydracarina of Wisconsin

285

Bibliography

Titles are limited to papers containing authors’ descriptions of the

species cited and to general papers containing descriptions of cosmopolitan

forms.

Marshall, R.

1929. Canadian Hydracarina. Univ. of Toronto Studies: Biol.

Ser. Pub. Ontario Fisheries Research Lab., No. 39, p. 57-93,

PI. I-VII

1931. Preliminary List of the Hydracarina of Wisconsin, Part I.

Trans. Wis. Acad. S.A.L. XXVI: 311-319, pi. VII-VIII

1932. ditto, Part II

Trans. Wis. Acad. S.A.L. XXVII: 339^358, pi. VII-X

1933. ditto, Part III

Trans. Wis. Acad, S.A.L. XXVIII: 37-61, pi. I-IV

Piersig, R.

1897. Deutschlands Hydrachniden. Bibl. Zool. XXII: Stuttgart;

1897-1900

1901. Hydrachnidae [und Halacaridae] . Das Tierreich, XIII,

Berlin

Soar & Williamson

1927. British Hydracarina, Vol. II. The Ray Soc., No. 112,

London

1929. ditto, Vol. Ill, The Ray Soc., No. 115, London

Thor, S.

1898. Huitfeldtia, En ny Hydrachnide-slegt. Arch. Naturv.

Christiana, XX (7) : 4, pi. V, fig. 1-7

Viets, K.

1930. Ueber nordamerikanische Koenikea-Arten (Hydracarina).

Zool. Anz., Bd. 92: 266-272

Wolcott, R. H.

1900. New Genera and Species of North American Hydrachnidae.

Trans. Am. Mic. Soc. XXI: 177-200, pi. IX-XII

1902. The North American Species of Curvipes [Piona]. Trans.

Am. Mic. Soc. XXIII: 201-256; pi. XXIX-XXXIII

1905. A Review of the Genera of the Water Mites. Trans. Am.

Mic. Soc. XXVI: 161-243, pi. XVIII-XXVII

286

Wisconsin Academy of Sciences , Arts , and Letters.

Explanation of the Plates

VI

1. Koenikea haldemani , genital area, male

2. Koenikea haldemani, ventral side, female

3. Koenikea concava, genital area, male

4. Koenikea concava, right palpus, female, inner side

5. Koenikea concava, ventral side, female

6. Koenikea wolcotti, rostral spine and palpi, female

7. Koenikea wolcotti, ventral side, female

TRANS. WIS. ACAD., VOL. 29.

PLATE VI

288

Wisconsin Academy of Sciences, Arts, and Letters.

VII

8. Koenikea marshallae, dorsal side, female

9. Koenikea marshallae, genital region, male

10. Koenikea marshallae , ventral side, female

11. Koenikea marshallae , rostrum and right palpus, female

12. Koenikea spinipes, ventral side, male

13. Koenikea spinipes, rostrum and right palpus, female

14. Koenikea spinipes, end of leg I, male (one claw omitted)

15. Koenikea spinipes , genital region, female

.RANS. WIS. ACAD. VOL. 29.

PLATE VII

290

Wisconsin Academy of Sciences , Arts , and Letters .

VIII

16. Huitfeldtia rectipes , genital plates, female (after Thor)

17. Huitfeldtia rectipes, right palpus

18. Huitfeldtia rectipes, ventral side, male

19. Forelia ovalis, ventral side, male

20. Forelia ovalis , leg IV, male

21. Forelia ovalis, ventral side, female

22. Forelia liliacea, ventral side, male

23. Forelia liliacea, genital area, female

24. Forelia liliacea, 5th and 6th segments, leg IV, male (foreshortened)

25. Forelia liliacea, right palpus, female, outer side

292

Wisconsin Academy of Sciences , Arts , and Letter B.

IX

26. Piona rotunda , genital area, male

27. Piona rotunda , end of leg III, male

28. Fiona rotunda , right palpus, male, outer side

29. Piona rotunda , genital area, female

30. Piona rotunda , 4th segment, leg IV, male

31. Piona rotunda, 6th segment, leg II, male

32. Piona reighardi, 5th and 6th segments, leg III, male

33. Piona reighardi, 4th segment, leg IV, male

34. Piona reighardi, right palpus, male, inner side

35. Piona reighardi, female, genital area

36. Piona reighardi, ventral side, male

Trans, wis. acad. vol. 29.

PLATE IX

294 Wisconsin Academy of Sciences , Arts, and Letters .

X

37. Piona crassa , ventral side, male

38. Piona crassa, genital area, female

39. Piona crassa, right palpus, female, outer side

40. Piona crassa, dorsal side, female

41. Piona crassa, leg IV, male

42. Piona crassa , 5th and 6th segments, leg III, male

43. Piona conglobata, genital area, male

44. Piona conglobata , 6th segment, leg II, male

45. Piona conglobata, genital area, female

46. Piona conglobata , 6th segment, leg III, male

47. Piona conglobata , right palpus, male, outer side

TRANS, WI$. ACAD. VOL. 29.

JPLATE £

296

Wisconsin Academy of Sciences , Arts, and Letters .

XI

49. Piona media , left palpus, male, inner side

50. Piona media , genital area, female

51. Piona media » ventral side, male

52. Piona media , 6th segment, leg III, male

53. Piona media , 4th segment, leg IV, male

54. Piona inconstans, ventral side, male

55. Piona inconstans, left palpus, female, outer side

56. Piona inconstans, 6th segment, leg III, male

57. Piona inconstans, genital area, female

TRANS. WIS. ACAD. VOL. 29.

PLATE XI

PRELIMINARY REPORTS ON THE FLORA OF

WISCONSIN. XXLV. SALICACEAE

David F. Costello

The maps in this report are based on specimens in the fol¬

lowing herbaria: University of Wisconsin, Milwaukee Public

Museum, Mr. S. C. Wadmond, of Delavan, Wis., and Iowa State

College, Ames, Iowa. The writer is grateful for the courtesies

extended by the curators of these herberia. The Wisconsin wil¬

lows in the Gray Herbarium were listed by Dr. Norman C. Fas-

sett. Ranges of the poplars as noted by Mr. L. S. Cheney have

been used in compiling the maps for that genus. Many of the

willows in the author's herbarium have been examined and the

identifications checked by Dr. Carleton R. Ball of the University

of California. His kindly criticism and his many useful notes

and suggestions have been a source of inspiration.

1. POPULUS

Localities based on unpublished studies by Mr. Cheney are

here represented by small dots. Additions to the ranges shown

by Cheney, represented by herbarium specimens, are shown by

large dots.

The following key is based on mature leaves.

Costello : Prelim. Rept. Flora Wis. XXIV Salicaceae

a. Leaves serrate or toothed; glabrous beneath

b. Petioles flattened laterally

c. Leaf blades ovate to orbicular in outline

d. Blades 3-6 cm. long, margins finely serrate

_ _ _ . _ _ _ P. tremuloides

dd. Bladies 5-10 cm. long, margins coarsely

toothed _ _ _ _ _ _ _ P. grondidentata

cc. Leaf blades triangular to rhombic ovate in outline

d. Blades broader than long, abruptly acuminate;

marginal teeth slightly incurved; tree with col-

299

300 Wisconsin Academy of Sciences, Arts, and Letters.

umnar habit _ _ __P. nigra var. italica

dd. Blades longer than broad.

e. Blades distinctly triangular, truncate or broad¬

ly cordate at base; marginal teeth distinctly

incurved; tree with broad rounded crown

_ _ _ $ P. deltoides

ee. Blades rhombic ovate, broadly rounded at base ;

margin crenate serrate; tree with pyramidal

habit _ _ _ X P. Eugenei

bb. Petioles round in cross section

c. Blades ovate lanceolate, broadly rounded at the

base _ _ _ _ _ _ _ „_P. bats ami f era

cc. Blades broadly ovate, cordate at the base ; veins

pubescent below _ _ _ _ _ P. candicans

aa. Leaves lobed or coarsely sinute-crenate ; petioles and

lower sides of leaves densely white tomentose - P. alba

1. P. alba L. White Poplar, Silver Leaf Poplar. (Fig. 1).

Introduced from Europe. Extensively cultivated and freely

spreading by root. Two forms are commonly cultivated in the

state: P. alba var. nivea (Willd.) Wesm., characterized by its

maple like leaves which are densely white tomentose beneath;

and P. alba var. pyramidalis Bunge [P. alba var. Bolleana

(Lauche) Wesm.], characterized by its columnar habit, and

smooth green bark. Herbarium specimens of P. alba and its

varieties are much needed.

2. P. tremuloides Michx. American Aspen, Trembling Pop¬

lar, Quaking Aspen. (Fig. 2). Occurs in every county in the

state. The habitat varies from sandy shores of lakes, tamarack

swamps, and dry barrens to limestone ridges and the summits

of wooded bluffs.

The foliage of this tree is exceedingly variable. P. tremu¬

loides f. reniformis Tidestrom1 is, “distinguished by its kidney¬

shaped leaves, which are ordinarily about 5-6 cm. in length

and 7-8 cm. in width.” P. tremuloides var. intermedia Vic¬

toria has heart shaped leaves which are relatively large by the

1 Rhodora 16:206. 1914

a Contrib. Lab. Bot. Univ. Montreal. No. 16. 1930

Costello— Reports on Flora of Wisconsin 301

time the fruits have matured. I have collected a form near

West Allis and also near Newburg which is characterized by

cordate leaves which retain their dark shining green color for a

period of two or three weeks after the leaves of typical P. trem-

uloides have become pale lemon yellow in the autumn. This

form, however, does not appear to be identical with P. tremu-

loides f. reniformis Tidst., which was described from Maryland,

nor does it resemble P. tremuloides var. intermedia Victorin,

described from Quebec, which has much larger leaves. Further

study will be necessary to determine its exact status.

Leaves from vigorous shoots or sprouts frequently are cord¬

ate at the base and much larger than typical leaves in this spe¬

cies. They are frequently misidentified in herbaria.

3. P. grandidentata Michx. Large Toothed Aspen. (Fig.

3). Like P. tremuloides this tree occurs in every county. Com¬

mon along river banks, in rich woods, swamps and on moist

uplands. Variations from the typical leaf form are not uncom¬

mon. Rotund leaves or leaves with cordate bases and typical

leaves are frequently found on the same branch. Occasionally

leaves with entire margins, or with two or three lobes may be

seen. Leaves of sucker sprouts are large, cordate at base, and

covered with a fine pubescence.

4. P. balsamifera L. (P. tacamahacca Mill.). Balsam Pop¬

lar, Tacamahac. (Fig. 4). Particularly abundant in the north¬

eastern one fourth of the state and in the Lake Superior region.

Common along river banks and borders of swamps.

5. P. deltoides Marsh. (P. monilifera Ait.; P. virginiana

Fourg. ; P. balsamifera L., in part). Cottonwood. (Fig. 5).

This tree reaches its northern limits in Wisconsin, occupying

the southern two thirds of the state. It is especially abundant

along the Mississippi, Wisconsin, Kickapoo, Chippewa, Red

Cedar, Embarass, Fox and Sugar River bottoms.

6. P. candicans Ait. (P. balsamifera var. candicans Gray) .

Balm or Gilead. Introduced. Occasionally found as an escape

from cultivation. Cheney's maps show that it has been planted

over approximately the same range as the following.

7. P. nigra L. var. italica Du Roi. Lombardy Poplar. (Fig.

6). Introduced from Europe. Frequently spreading from cul¬

tivation. Only staminate trees are known.

302 Wisconsin Academy of Sciences, Arts, and Letters.

8. X P. Eugenei Simon-Louis. Catal. ex Schneider, Ill.

Handb. Laubholzk. 1:9. 1904. Norway Poplar, Sudden Saw-

log. Planted in the northwestern part of the state. A rapid

growing tree with rhombic-ovate leaves, rounded at the base,

acuminate at the tip, coriaceous, with rounded teeth. This tree

is not well understood. Some have held it to be a hybrid, P.

deltoides X P. nigra var. italica ; others believe it to be a form

of the P. deltoides complex. Good specimens for comparison are

needed.

2. Salix

Since willows are dioecious, and since they hybridize freely

and show great variation in response to environmental condi¬

tions, they are a difficult group to identify. The only satisfac¬

tory way to study them is to tag individual shrubs or trees with

metal labels and to collect material from these individuals

throughout the seasons. Only in this way can one be certain as

to which flowers go with which leaves. Material of this kind

is much needed in herbaria.

The leaves or individuals vary from season to season as does

the condition of hairiness on leaves and twigs. Twigs exposed

to the sun may be pubescent while shaded twigs on the same

plant are glabrous. Second growth, following grazing, cutting

by mowing machines, or defoliation by insects almost invariably

results in the production of pubescent leaves. Hybrids may

usually be expected to show characteristics intermediate be¬

tween the species of the cross. Monstrosities, particularly of

the flowers, are not uncommon.

In spite of the multitudinous variations of willows, it is

hoped that the three following keys may be of use.

Key to Mature Leaves

a. Leaves glabrous or glabrate beneath

b. Leaves green on both sides

c. Leaves linear to linear lanceolate

d. Leaves linear, short acuminate ; margins remote¬

ly denticulate - - _ - - - S. interior

dd. Leaves lanceolate, long attenuate from near the

base; margins closely serrate _ S. nigra

Costello — Reports on Flora of Wisconsin

303

cc. Leaves oblong lanceolate, to broadly lanceolate

or ovate

d. Leaves short acuminate

e. Leaves leathery, bright green and shining

above (cultivated tree) _ _ S. pentandra

ee. Leaves thin, not shining, usually paler

beneath, cordate at base on vigorous

shoots _ _ _ _ S. cor data

dd. Leaves with long acuminate curved tips, shining

on both sides - - - S. lucida

bb. Leaves glaucous or whitish beneath

c. Margins entire or irregularly crenate-serrate

d. Margins entire, revolute; blades linear-oblong to

elliptic-obovate, slightly coriaceous, 2-6 cm.

long, veiny above

_ _ _ -S. pedicellaris var. hypoglauca

dd. Margins irregularly crenate-serrate; blades

lanceolate to elliptic, 5-12 cm. long discolor

cc. Margins distinctly and uniformly serrate

d. Leaves broadly rounded to cordate at base

e. Leaves dull above, paler beneath cor data

ee. Leaves glossy above, glaucous beneath

f. Petioles short, leaves blackening in

drying; stipules large _ S. glaucophylla

ff. Petioles long and slender, leaves

prominently reticulate veined beneath;

stipules obsolete or minute _ S. pyri folia

dd. Leaves acute at base

e. Petioles glandular at apex.

f. Tall shrub; leaves short acuminate,

coriaceous, elliptical _ S. serissima

ff. Large trees; leaves long acuminate,

lanceolate

304

Wisconsin Academy of Sciences , Arts , and Letters .

g. 7-10 teeth per cm. of margin;

branches slightly brittle at base

_ S. alba

gg. 4-6 teeth per cm. of margin;

branches very brittle at base

_ S. fragilis

ee. Petioles not glandular.

f. Leaves ovate lanceolate, long attenuate;

petioles 1-3 cm. long; large tree

_ _ ,-S. amygdaloides

ff. Leaves narrowly lanceolate; petioles

1 cm. or less; shrub _ S . petiolaris

aa. Leaves pubescent beneath

b. Margin entire or nearly so

c. Blades densely white tomentose below, dark green

above, with sunken veins; branchlets sometimes

densely white tomentose _ _ S. Candida

cc. Blades grayish tomentose to somewhat glaucous

below

d. Leaves 4-10 cm. long, petioles distinct; shrub

1 - 3 m. high _ S. humilis

dd. Leaves 1-5 cm. long, petioles very short;

shrub 0.5 m. high _ S. tristis

bb. Margin finely serrate to crenate serrate.

c. Leaves ovate or obovate to broadly lanceolate

d. Leaves serrate-crenate, thin, mostly acute at

base, usually finely pubescent and rugose

beneath _ _ _ _ S. bebbiana

dd. Leaves closely glandular serrate, thick and

leathery, cordate or rounded at base, frequently

permanently silky lanate above _ S. adenophylla

cc. Leaves linear or narrowly lanceolate.

d. Leaves linear to oblong-lanceolate, margins

remotely denticulate; nearly sessile _ S. interior

Costello — Reports on Flora of Wisconsin

305

dd. Leaves lanceolate, evenly serrate; petioles

distinct.

e. Petioles glandular; trees __ - S. alba

ee. Petioles not glandular; shrubs

f. Leaves slightly silky beneath, sometimes

drying black _ _ _ _ _ S. petiolaris

ff. Leaves densely silvery pubescent beneath,

black in drying _____ - — sericea

Key to Staminate Flowers, Twigs, and Young Leaves

Flower scales yellowish

b. Stamens 3 - 5 or more

c. Aments elongated, slender-cylindrical; scales crisp

villous on the inside

d. Stipules large, semi-cordate and pointed, or small

and ovate, persistent or deciduous _ _ S. nigra

dd. Stipules reniform, minute or none

_ ______ _ _ _ _ S. amygdaloides

cc. Aments thick, short-cylindrical or ellipsoid-ovoid,

densely flowered.

d. Aments appearing with the leaves.

e. Young leaves glabrous from the first

_ _ _ _ _ __ _ S. pentandra

ee. Young leaves pubescent with crisp rufescent

or sordid caducous hairs _ _ _ _ S. lucida

dd. Aments appearing after the leaves (in early

summer) _ _ _ _ _ _ _ _ S. serissima

bb. Stamens 2

c. Aments appearing after the leaves, frequently in

clusters of 2 or 3 _ _ _ _ _ _ ,S. interior

cc. Aments appearing with the leaves

d. Aments 1.5 - 2.5 cm. long; leaves sparsely

pubescent when young ; twigs long and slender,

drooping - - - S. babylonica

306 Wisconsin Academy of Sciences , Arts, and Letters.

dd. Aments 3-5 cm. long; twigs not drooping

e. Young leaves glabrous from the first

_ _ _ S. fragilis

ee. Young leaves silky pubescent on both sides

f. Twigs green to olive-brown _ _ S. abla

ff. Twigs yellow or reddish

- - - S. alba var. vitellina

aa. Scales brown to black or colored at tip (stamens 2)

b. Flowers appearing before the leaves,

c. Anthers reddish

d. Filaments hairy; anthers often united so as to

appear as one ; buds sub-opposite on the twig

_ _ _ _ ...S, purpurea

dd. Filaments smooth; anthers not united; buds

alternate ; low shrub of cold bogs — -S. Candida

cc. Anthers yellowish

d. Aments 2.5-5 cm. long

e. Flower scales brown or black, with long silky

hairs

f. Branchlets dark brown to black or purple

_ discolor

ff. Branchlets stout, mostly yellow to

chestnut brown _ glaucophylla

ee. Flower scales rose tipped, thinly villous;

branching divaricate; bud scars numerous

_ _ S. bebbiana

dd. Aments 0.5 - 2.5 cm. long

e. Aments narrowly cylindrical, 2 cm. long,

sometimes leafy bracted at base; young

leaves very silky when they appear -S. sericea

ee. Aments globular, ovoid, or ellipsoid; young

leaves downy but not silky when they appear

f. Aments ovoid or ellipsoid, 1-2 cm. long,

often recurved - - S . humilis

Costello— Reports on Flora of Wisconsin

307

ff. Aments globular or ovoid, 0.5 - 1 cm. long

— — - - tristis

bb. Flowers appearing with the leaves

c. Young leaves silvery silky

d. Young leaves densely lanate, ovate or broadly

lanceolate; aments 1.5 - 3 cm. long; twigs stout,

frequently yellowish _ _ _ _ ,S. adenophylla

dd. Young leaves thinly silky pubescent, narrowly

lanceolate ; aments 1-2 cm. long ; twigs slender,

usually purplish _ _ _ petiolaris

cc. Young leaves not silvery silky

d. Young leaves glabrous from the first

e. Scales glabrous or glabrate, greenish yellow;

twigs slender; aments 1.5 - 2 cm. long

_ _ S. pedicellaris var. hypoglauca

ee. Scales pubescent or silky villous ; twigs

medium to stout; aments 2-3 cm. long

- — _ _ _ _ _ S. pyrifolia

dd. Young leaves puberulent or pubescent

e. Aments 1-2 cm. long, narrowed at base ;

twigs slender, divaricate, usually full of bud

scars _ _ _ _ _ _ _ _ bebbiana

ee. Aments 2-4 cm. long, slender; twigs

medium, not divaricate _ _ ~S. cor data

Key to Pistillate Flowers, Twigs, and Young Leaves

. Mature capsules glabrous (except sometimes in S. interior)

b. Scales yellow, falling before the catkins are ripe

c. Flowers appearing after the leaves ; tall shrubs

d. Leaves linear lanceolate, remotely denticulate;

capsules blunt ; fruiting in late spring or early

summer _ _ _ _ _ __ _ _ _ interior

dd. Leaves ovate lanceolate, shining above; capsules

awl shaped ; fruiting in July and August

_ , _ S. serissirm

308 Wisconsin Academy of Sciences , Arts , and Letters .

cc. Flowers appearing with the leaves

d. Twigs slender, sometimes drooping; catkins

slender and loose (except in S. babylonica) ;

trees

e. Fruiting aments 3-11 cm. long; capsules

distinctly pedicelled, 3-6 cm. long.

f. Stipules minute, very early deciduous;

aments becoming very loose in fruit;

capsules globose-conical, long pedicelled

amygdaloides

ff. Stipules large, semi-cordate, pointed and

persistent, or small, ovate and deciduous;

aments more or less dense; capsules

ovoid-conical, short pedicelled ___£. nigra

ee. Fruiting aments 1.5 - 2 cm. long; capsules

sessile, 1-1.5 mm. long _ S. babylonica

dd. Twigs medium or stout, not drooping

e. Catkins thicks, short and dense; shrubs or

small trees

f. Capsule conic-subulate; pedicel twice

exceeding the gland ; young leaves

glabrous from the first (cult, shrub or

small tree) _ _ _ S. pentandra

ff. Capsules conic-ovoid (long beaked) ;

pedicels 3 or 4 times exceeding the gland;

young leaves sparingly rusty pubescent

_ _ - _ lucida

ee. Catkins slender, loosely flowered, 4-8 cm.

long in fruit; large trees

f. Capsules sessile or nearly so, ovoid

conical (short ovoid) ; young leaves silky

pubescent alba

ff. Capsules short pedicelled, subulate-conical

(long conic lanceolate) ; leaves glabrous

from the first fragilis

bb. Scales brownish to black, persistent (except S.

hebbiana)

Costello— Reports on Flora of Wisconsin

309

c. Aments usually appearing before the leaves; young

leaves glabrous from the first ; scales gray

pubescent with long, matted or twisted hairs

_ _ _ _ -S. glaucophylla

cc. Aments appearing with the leaves

d. Capsules reddish or reddish green, 5-8 mm.

long

e. Young leaves and twigs glabrous throughout;

leaf margins entire; pedicels 2-4 mm. long

_ _ — _ S. pedicellaris var. hypoglauca

ee. Young leaves densely lanate pubescent;

margins serrate; twigs pubescent; pedicels

0.5 - 1 mm. long _ _ _ _ S. adenophylla

dd. Capsules greenish

e. Capsules 8-10 mm. long; pedicels 2.5 - 3.5

mm. long; twigs shining, reddish-castaneous

or olive; stipules obsolete or minute

_ _ _ _ _ S. pyri folia

ee. Capsules 4.5 - 6 mm. long; pedicels 1 - 1.5

mm. long; twigs yellowish to dark brown,

puberulent to pubescent; stipules persistent,

usually conspicuous; young leaves lanceolate

_ _ _ _ _ _ _ _ _ S . cor data

Mature capsules pubescent

b. Aments appearing before the leaves

c. Buds alternate on the twig; capsules distinctly

pedicelled

d. Flower scales brown or black, at least at the tip

e. Capsules densely white woolly; twigs

sometimes white woolly; styles dark red

_ _ S. Candida

ee. Capsules not white woolly; twigs brown or

black

f. Twigs black or purple; mature capsules

7-12 mm. long; aments 2.5 - 7 cm. long;

scales copiously clothed with long glossy

hairs _ S. discolor

310 Wisconsin Academy of Sciences , Arts, and Letters.

ff. Twigs brown; mature capsules 6-9 mm.

long; aments 1-4 cm. long in fruit,

g. Catkins 1.5 - 4 cm. long in fruit; tall

shrub (1 - 2 m. high) _ S. humilis

gg. Catkins 1 - 1.5 cm. long in fruit; low

shrub (0.5 m. high) _ _ S. tristis

dd. Flower scales yellow, rose tipped, elongate

(aments sometimes appearing with the leaves)

_ _ _ _ S. bebbiana

cc. Buds sub-opposite on the twig; capsules sessile

(cultivated shrub or small tree) _ _ S. purpurea

bb. Aments appearing with the leaves

c. Capsules 3-5 mm. long, ovoid-oblong (blunt) ;

fruiting aments dense, sessile or subsessile

_ _ _ S. sericea

cc. Capsules 6-8 mm. long, lanceolate-conic; fruiting

aments lax, leafy bracteate; twigs frequently drying

purplish black - - - - — _ _ _ — S. petiolaris

1. S. nigra Marsh. Black Willow. (Fig. 7). Banks of

streams and moist places in rich soil. Although widely distri¬

buted in the state it is nowhere abundant. Most abundant in

southern two thirds of the state.

2. S. amygdaloides Anders. Peach Leaved Willow. (Fig.

8). Common in moist rich soil throughout southern two thirds

of the state. Cheney states that the range is, “a little south of

the range of S. nigra ” Herbarium specimens of this tree are

few and of poor quality.

3. S. pentandra L. Bay Leaved Willow. A cultivated spe¬

cies introduced from Europe. Seldom escapes.

4. S. lucida Muhl. Shining Willow. (Fig. 9). Widely dis¬

tributed excepting in the southwest corner of the state. Com¬

mon on low ground, in ponds and along streams. I have col¬

lected, in a pond near Cedarburg, a form of elongated capsules

as in S. lucida ; leaves somewhat resembling S. fragilis, but

silky beneath and with 9-11 serrations per cm. of margin as

in S. alba. Dr. Ball has examined my specimens and leaves

Costello— -Reports on Flora of Wisconsin 311

them as S. lucida X S. alba ( ?). I have not seen material from

Wisconsin that appears to be S. lucida var. intonsa Fernald.

5. S. serissima (Bailey) Fernald. Autumn Willow. (Fig.

10) . Apparently confined to the northwestern and southeastern

portions of the state. Found in mossy swamps with Betula pu-

mila var. glandulifera , Larix laricina and Rhus Vernix. Espe¬

cially abundant about bogs in the southeastern counties.

6. S. fragilis L. Crack Willow. Naturalized from Europe.

One of our commonest willows along roadsides and streams in

the southern part of the state. I have collected branched stam-

inate catkins from a number of willows of this species near

Thiensville for three successive years3. It seems probable that

this is an inherited characteristic.

7. S. alba L. White Willow. Introduced from Europe.

Widely planted and freely escaping. Leaves often permanently

silky beneath ; twigs greenish. Var. vitellina L. Koch has more

Koch has more glabrate leaves; twigs yellow or orange.

8. S. babylonica L. Weeping Willow. Naturalized from

Europe. Occasionally found along streams. Cultivated vari¬

eties are numerous.

9. S. interior Rowlee. S. longifolia Muhl. S. fluviatilis of

many authors. Sandbar Willow. (Fig. 11). Common throughout

the state. This willow frequently flowers a second time. I have

collected flowers as early as April 25 and as late as July 20.

When the leaves of this species are eaten by larvae, the new

leaves which are then produced are always more or less silvery

pubescent.

What appears to be S. interior var. pedicellata (Andersson)

Ball has been collected in Grant County (Fig. 11, cross). The

very narrow and finely denticulate leaves agree with typical

specimens collected along the Missouri River in southeastern

Nebraska where it is common.

10. S. cordata Muhl. Heart Leaved Willow. (Fig. 12).

Frequent throughout the state. In wet places, around ponds

and on the banks of streams. The leaves of this willow are

variable. Narrow leaved forms are common, while the leaves

•Costello, Jour. Heredity 23 ; 238. 1932,

312 Wisconsin Academy of Sciences, Arts, and Letters.

of vigorous shoots and water sprouts are broadly oblong lanceo¬

late with cordate bases.

11. S. glaucophylla Bebb. Glaucous Leaved Willow, Blue-

leaf Willow, Broadleaf Willow. (Fig. 13). Follows the western

shore of Lake Michigan closely. Characteristic of sandy shores

of lakes. A very small leaved form grows on the dry clay banks

of the ravines near Lake Michigan in Milwaukee and Racine

Counties. It is doubtfully var. angustifolia Bebb. Wadmond4,

in his Flora of Racine and Kenosha Counties, states that he

finds S. glaucophylla, “inland on low prairies more abundantly

than near the Lake, and even when found near the Lake, it

seems to affect wet clayey soils, rather than sand.”

12. S. pyrifolia Anders. S. balsamifera Barratt, Gray's

Man., ed. 7. Balsam Willow. (Fig. 14). Mostly in the north¬

ern half of the state. Low ground and in spruce swamps. In¬

frequent.

13. S. adenophylla Hooker. S. syrticola Fernald, of Gray's

Man., ed. 7. Gland Leaved Willow. (Fig. 15, dot). The rarest

willow in the state. Has been collected only at Two Rivers,

Manitowoc Co., on the sand dunes. In Indiana it is fairly com¬

mon in the dune region bordering the lake. Usually found

growing with S. glaucophylla in the dunes at the south end of

Lake Michigan.

14. S. pedicellaris Pursh.5 Bog Willow. (Fig. 16, cross).

A single specimen, collected by A. M. Fuller on Washington Is¬

land, Door County, seems to belong here. The sub-coriaceous

leaves are green on both surfaces as contrasted with the pale

glaucous under surfaces of the leaves of the following variety.

S. pedicellaris var. hypoglauca Fernald.6 (Fig. 16). Most

of our material seems to belong here. Common in bogs and

springy meadows.

15. S. discolor Muhl. Glaucous Willow, Pussy Willow. (Fig.

17) . Probably found in every county in the state. Very common

in swamps and wet places.

S. discolor var. eriocephala (Michx.) Anders. (S. erioceph -

ala Michx.). This variety is characterized by pubescent twigs

4 Wadmond, Trans. Wis. Acad. Sci., Arts and Let. 16:826. 1909

* Fernald, Rhodora 11:157-162. 1909

8 Fernald, l.c.

Costello— Reports on Flora of Wisconsin 813

and by leaves more or less pubescent below. Frequently the

aments are more densely pubescent than in the species. Range

the same as the species.

16. S. petiolaris J. E. Smith. Slender Willow. (Fig. 18).

Common in meadows, along banks of streams and in ponds.

Forms with pubescent leaves are frequently collected. The sec¬

ond crop of leaves, following defoliation, is always more or less

pubescent.

17. S. humilis Marsh. Prairie Willow, Upland Willow.

(Fig. 19). Throughout the state, but nowhere abundant. Dry

prairies and sandy areas.

18. S. tristis Ait. Dwarf Gray Willow, Dwarf Pussy Wil¬

low. (Fig. 20). Dry sandy places, borders of woods and clear¬

ings. Similar to above but smaller in every respect. Russel7

states that this species was reported from Milwaukee County

by J. A. Brandon in 1900. I have seen no herbarium specimens

from that locality.

19. S. sericea Marsh. Silky Willow. (Fig. 15, squares). In

moist places. Should be sought for in the Driftless Area. The

station noted as Beloit, Rock Co. is doubtful. This sheet, con¬

taining two species, no date, collected by Dr. Lathrop, possibly

has New York as its locality.

20. S. bebbiana Sargent8 ( S . rostrata Richards., Grays

Man., ed. 7). Bebb Willow, Beak Willow. (Fig, 21). Common

in moist or swampy areas throughout the state. A large shrub

with leaves finely pubescent above and densely pubescent be¬

neath. The glabrate leaved variety, S. bebbiana var. perrostrata

(Hydb.) Schn., grows with the species but is not common.

21. S. Candida Flugge. Sage Willow, Hoary Willow. (Fig.

22). Found mostly in cold bogs at the margin of tamarack

spruce swamps.

S. Candida var. denudata Anders. (Fig. 22, cross). First

collected by A. M. Fuller, No. 2371, in the Cedarburg Bog,

Ozaukee Co., 1928. As in the Indiana specimens collected by

Chas. C. Deam9, the capsules are entirely glabrous. The ma-

7 Russel, Bull. Wis. Nat. Hist. Soc., 5:186. 1907

8 Rhodora, 26:122-123. 1924

9 Deam. Shrubs of Indiana p. 58. 1924,

314 Wisconsin Academy of Sciences, Arts, and Letters.

tured capsules are short pedicelled, and similar in shape to typi¬

cal Candida. The lower fourth of the narrow leaves is inrolled

to such an extent as to give to the leaf the appearance of being

long petioled. The leaves are green on both sides, rugose above

and strongly veined below. I have recently collected specimens

of this variety in the same place.

S. Candida X S. petiolaris. Cedarburg Bog, Ozaukee Co. For

two successive years I have found this cross growing on the

quaking bog in company with S. Candida and S. petiolaris.

Leaves are intermediate between the two species, like £. petio¬

laris in shape and mostly like S. Candida in pubescence. Inter¬

mediates resembling one species or the other more closely than

the cross have also been collected. Carleton R. Ball has seen my

specimen No. 928, July 8, 1932, and states in a letter that al¬

though the thinly tomentose to glabrate undersurfaces of the

leaves might indicate variety denudata Anders., “the leaves are

broader than denudata as seen by me.” He also mentions the,

“sparsely denticulate and scarcely re volute margins, which

would be expected in hybrids with petiolaris .”

22. S. viminalis L. Osier. Introduced from Europe and

cultivated in parks as an ornamental. No evidence that is has

escaped anywhere in the state.

23. S. purpurea L. Purple Willow, Basket Willow. Na¬

turalized from Europe. A common escape along roadsides and

in waste places. The subopposite leaves are a distinguishing

characteristic.

Costello — Reports on Flora of Wisconsin

315

Populus grandidentata Populus balsamifera

316 Wisconsin Academy of Sciences , Arts, and Letters .

Salix interior

Salix cordata

Costello — Reports on Flora of Wisconsin

Salix glaucophylla

Salix pyrifolia

Salix discolor

Salix petiolari^

318 Wisconsin Academy of Sciences , Arts, and Letters .

Salix 'be'bbiana

+ S. Candida var. denudata

ELABORATION OF SETTING IN OTHELLO AND

THE EMPHASIS OF THE TRAGEDY

Julia Grace Wales

I

The Setting

The Significance of the Study of Place for this Play

The sense of place is one of the major factors in Shake¬

speare's imaginative appeal, though not place, necessarily, in

any scientific, geographical signification. That he is no realist

interested in facts for themselves, must not be allowed to ob¬

scure the complementary truth that he is no classicist either,

but a romantic. There is nothing abstract about his scene. It

is individualized through homely detail. His airy nothing is

always given name and habitation; and largely from these ob¬

tains its interpretation and emphasis.

In a study of Othello from this point of view it would be

instructive, did space permit, to bring together all the concrete

detail of the play, whether English, foreign, or neutral, and to

examine and classify it for its significance in creating atmo¬

sphere and building up a sense of locality. The cumulative ef¬

fect of this detail would be greatly enhanced could we gather

and place beside it all the notes and illustrations, collected by

editors1 of the play or scattered through learned articles,2 which

1 We can only refer the reader to the various annotated editions of the play and cite a few

significant passages that have not yet found their way into these editions.

See, for example, Variorum, p. 5 for C. A. Brown’s comment, and p. 8 for Staunton’s, on the

mercantile associations of Florence; p. 46 for Knight’s note on Luccicos; pp. 30-32, a long note

on “as double as the duke’s” (I. ii. 11-17), with quotations from Contareno and Thomas on the

power of the Duke in relation to the rest of the Council. Again, see Variorum, pp. 28, 29, for

a long and interesting note, including Malone’s famous illustration from Lewkenor on Venetian

officers especially charged with protecting the city by night.

See also Hart’s note on nettles (I. iii. 32S-7), Arden Edition (1903), p. 53; that on

chrysolite (V. ii. 143), p. 240; that on coloquintida (I. iii. 355), p. 57.

See a good note in the Yale Edition (1918), p. 134, on the historical background of the war.

2 One passage must be quoted from the researches of Sir Edward Sullivan, “Shakespeare in

Italy,” The Nineteenth Century (1908), pp. 329-330. “The word Veronessa shows an intimacy

with Italy in two distinct directions; first, it is a correctly formed feminine adjective, meaning

‘of, or belonging to Verona’; and secondly, it implies an acquaintance with the fact that Verona

(which was in the Venetian State) furnished war galleys to Venice, or that the Venetians kept

a portion of their own fleet at or near Verona. . . . The importance from a naval point of

view of the Adige, which flows through Verona, may be appreciated by reading in pre-Shake-

spearian Italian histories the account given of the fleet of ships sent by the Venetians up the

river to the Lago di Garda to assist their army in that district against Filippo Visconte, Duke of

Milan. The distance was about two hundred miles, and the flotilla consisted of twenty- five barks

and six galleys under the command of Zeno.”

He refers in a foot note to Alberti, F. Leandro, Descriptions di Tutta Italia, Venice, 1586,

p. 397, “who cites Biondo and other writers. See also Hazlitt’s History of the Venetian Republic

(1860), where a full account is given of this remarkable expedition.”

319

320 Wisconsin Academy of Sciences , Arts, and Letters ,

have extended its temporal and local significance and deepened

its imaginative connotation. To do this, however, would entail

too much repetition of accessible material.

The valuable researches of Mr. J. W. Draper,3 too recently

before us and too extended to be reviewed here, are evidence

enough that the subject of Italian local color in Othello had been

by no means exhausted by earlier writers and that useful ma¬

terial is still coming in.4

The more general works on the sixteenth and early seven¬

teenth centuries are constantly throwing new light on the sub¬

ject;5 and continued search into the materials of history, such

3 J. W. Draper: “Captain General Othello,” Anglia, XLIII (1931), 296; “‘This Poor Trash

of Venice’,” Jour, of Eng. and Germ. Philology, XXX (1931), 508; “ ‘Honest Iago’,” Pub. Mod.

Lang. Assoc. XLVI (1931), 724; “Shakespeare’s Othello and Elizabethan Army Life,” Rev.

Ang.-Am., IX (1932), 319; “Some Details of Italian Local Color in Othello,” Sh. Jb., 1932,

pp. 125-7; “Desdemona, a Compound of Two Cultures,” Rev. Litt. Comp., XIII (1933), pp.

337-351.

Especially interesting is the wealth of new illustration, given in these articles, of army life

in England and on the Continent and of conditions in England and Italy having to do with the

freedom of women.

4 See “Shakespeare’s Venice” by Violet M. Jeffrey, Mod. Lang. Rev., vol. 27 (1932), pp. 24-

35, in which the problem of the identity of “the Sagittary” is reopened.

5 Perhaps it is sheer coincidence that a passage in Hazlitt’s Venetian Republic seems to

illuminate Iago’s figure of speech (I. i. 75-77):

I’ll call aloud.

Do; with like timorous accent and dire yell

As when, by night and negligence, the fire

Is spied in populous cities.

The dread of fire like that of plague was common to all the countries of Europe. Hazlitt gives

an account of a great fire in Venice in 1577: “The damage was incalculable, and speculative

reports were soon spread over Europe of the amount of the losses.” ( The Venetian Republic,

1860. Ed. 1915. Vol. II, p. 133.) Disastrous fires occurred in 1479, 1483, 1574, 1577. ( Ibid .,

pp. 496-497.)

In Shakespeare’s England (Oxford, 1916, I. Ch. VI. pp. 170-171) we find, on the general

background, a passage of the utmost interest:

“For the characters and events of old-time plays, Shakespeare’s Europe is concentrated upon

Athens and Rome, but is extended to the easternmost recesses of the Mediterranean. In the Tudor

period the Turks had pushed this frontier of Europe westward: Rhodes (1522) and Cyprus (1571)

had fallen; Greece and its islands had already become Asiatic; and there was a redistribution

of forces. Accordingly, in the modern-Mediterranean plays of Shakespeare — All’s Well, Much Ado,

Two Gentlemen, The Winter’s Tale, Twelfth Night, Othello, Romeo and Juliet, and The Merchant

of Venice — Athens is not named, and Rome is named only twice {Tam. Sh. IV. ii. 75, Merck,

of V. IV. i. 153); and the scenes are laid in Verona, which is misdescribed as a tidal port

( Two Gent, of Verona, II. iii. 40), Venice, Padua, Milan, Mantua, Florence, Marseilles, Illyria,

Sicily, or Messina; and of these only the last two figure in the old-Mediterranean plays. The

eastern Mediterranean is only once to the fore. In Othello, Rhodes and Cyprus are physical and

political storm centers, where the Turks and Venetians would have fought had not all the Turks

been drowned; at Cyprus the Furies which watch over family life overwhelm all the leading

characters in the play. With this exception — if it is an exception — the modem plays shun the

eastern Mediterranean, Greece, Middle Italy, and all the principal places in the old-Mediterranean

plays. The sea is the same sea as of old, and swarms with pirates {Merck, of V. I. iii. 24),

like those of which Pompey wanted to rid it (Ant. & Cleop. II. vi. 36); and Italy is still the

place where Spaniards, Neopolitans, Frenchmen, Englishmen, Scotchmen, Germans, and Polish

Counts Palatine meet (Merck, of V . I. ii) ; and an occasional ‘Moor’ (Mohammedan) lends an

Asiatic or an African tinge. Italy is still cosmopolitan and dominates the Mediterranean, but

the center of political gravity has shifted, and for Shakespeare, whose instincts draw him to places

where life is rich and full, Italy and the Mediterranean mean different things in ancient and

modern times.”

See also an interesting note on “guards of the ever-fixed pole,” I, p. 453; also a note on

“carrack.” I. p. 153, which should be compared with the note on “lawful prize” in Variorum,

p. 37 — these comments affording an excellent illustration of Shakespeare’s method of piecing out

foreign information with English detail for the general purpose of realism in the sense of sub¬

stantiality. The present writer has endeavored to examine this composite process in two studies:

“Shakespeare’s Use of English and Foreign elements in the Setting of The Two Gentlemen of

Verona,” Wisconsin Academy of Sciences, Arts, and Letters, vol. 27 (1932), pp. 85-126, and

“Shakespeare’s Use of English and Foreign Elements in Much Ado about Nothing,” Ibid., vol. 28

Wales — The setting of Othello

321

as the travel books of the period and of slightly earlier and later

decades, is still rewarded with new and pertinent matter.6

Point by point these researches may seem merely curious.

But they have a cumulative and critical value. They prove be¬

yond a doubt that in Othello Shakespeare is building up a sense

of place, and that deliberately, for an artistic purpose. What

that purpose is becomes evident when we realize that the local

color of the play is not homogeneous but consists of two ele¬

ments in sharp contrast with each other, and that this contrast

is in vital relation to the tension of the tragedy.

The Italian background and the world of Othello's memories

may to our imagination have come to be merged romantically in

the general impression of the play. Yet in trying to get the

tragedy before us as a whole, it is of first importance to dis¬

tinguish the two and to keep the contrast between them vividly

in mind.

Venetian Color

The detail of Venetian color used in this play is less opulent

than that used in the Merchant of Venice, more economical, yet

adequate. We are left in no doubt how to visualize these back¬

grounds. To see how richly they are filled in, we have only to

(1933), pp. 363-398. Mr. J. W. Draper stresses the process as a conscious method in his

“Desdemona, a Compound of two Cultures.” (See note 3 above.)

Shakespeare’s England also affords information on the sword of Spain (I. p. 132):

“In Othello (V. ii. 252) we have ‘It is a sword of Spain, the ice brook’s temper.’ So the

modern version, but in the earliest printed edition. 1622, it was ‘The Ise Brokkes temper.’

Isebrook was the English name for Innsbruck in the Tyrol, whence some of the best steel was

imported into England from the early part of the sixteenth century until the Civil War. This

steel was used for the manufacture of armour in England, and ‘Isebrook’ and other variants of

spelling can be found in documents quoted in the Calendars of State Papers from April 1517

to April 1595. Moreover warm water of various degrees of heat was used here, as in Japan,

by the famous swordsmith Musamane, for tempering the blades. Othello’s expression merely

means a Spanish blade of the best Innsbruck temper.” If, by this interpretation, the passage

loses one element of poetry, it has, on the other hand, gained slightly in exotic suggestion.

e James Howell in his Survey of the Signorie of Venice, London, 1651, gives an elaborate

account of the government of the Republic. Though not available to Shakespeare, the following

passage is of interest:

“The Generali in warr upon the Continent is commonly som forren Prince; He is not chosen

either of the Senatorian or patrician order: he hath an ample salarie, viz. ten captaines pay,

and 4000 crownes a yer: ther goes along with him two Legats or Proveditores, who are Gentlemen

of Venice, and of the Senatorian order, and without the concurrence of their advice he neither

acts nor decrees anything himself without their intervention. These Proveditores are perpetually

assistants to the Generali and they pay the Soldiers Salaries, and their main care is that nothing

be done rashly to the detriment or dishonor of the Republic.”

I regret that I have not this page reference; nor have I been able to check the passage.

Howell also mentions (p. 17) the “Provosts of the night.” Cf. Variorum, p. 47, for more immedi¬

ately pertinent illustrations: Malone’s statement supported from Contareno (trans. by Lewkenor,

1599) that it was against the policy of the Venetian State to entrust the command of the army

to a native; also a passage cited by Reed from Thomas’s History of Italy to the same effect.

For an account of the travel books of the time, see Clare Howard, English Travellers of the

Renaissance (1914), with bibliography.

Fynes Moryson, writing in 1611, says after speaking of Italian love of revenge and skill in

making poisons ( Shakespeare’s Europe, Glasgow, 1907, pp. 405-406): “For which treasons the

Italians are so warye, espetially hauing a quarrell, as they will not goe abroade nor yet open

their doores to any knocking by night, or somuch as putt their head out of a windowe to speake

with him that knockes.”

322 Wisconsin Academy of Sciences, Arts, and Letters.

look at the meager setting of Cinthio’s story, of which the fol¬

lowing passages are the most significant as bringing out the

contrast :

The Venetians resolving to change the garrison which they maintain

in Cyprus, elected the Moor to the command of the troops which they

destined for that island ... he was extremely pleased with the honour

proposed to him (as it is a dignity conferred only on those who are noble,

brave, trusty, and of approved courage).

He had in his company an ensign of a very aimiable outward appear¬

ance, but whose character was extremely treacherous and base. „ . .

Had he not feared the strict and impartial justice of the Venetians he

would have put him openly to death. . . .

The Venetian magistrates, hearing that one of their fellow-citizens

had been treated with so much cruelty by a barbarian, had the Moor

arrested in Cyprus and brought to Venice, where, by means of the torture,

they endeavored to find out the truth. But the Moor possessed force and

constancy of mind sufficient to undergo the torture without confessing

anything.7

In the play, on the other hand, we have in the first scene the

chiaroscuro of a street by night, the taper at the window, flaring

torches and gleaming swords; in the second, the stateliness of

the Council Chamber and the gravity of the discussions going on

there, the stir of arriving messengers, and the background of

war and critical action. And it is worth noting that the theatri¬

cal interest of the two scenes, the suspense which gives to each a

focus and unity of its own, its own rise and fall of intensity, is

largely dependent upon the skillful use of setting. Similarly in

Act II the suspense arises out of the realization of place— the

seaport and citadel, the storm — “than which a fuller blast ne'er

shook our battlements," — the relief and serenity of a safe land¬

ing; the merry-making passing into carousal “in night and on

the court and guard of safety", “ in a town of war, yet wild, the

people's hearts brimful of fear," “the clink and fall of swords,"

voices “high in oath," the clanging of a dreadful bell.

When the main action in finally under way in the third act,

the sense of place is less immediate than indirect. It is the back¬

ground of Venice of which we are again made conscious— and

not now of its justice and dignity, of power worthily held, but

rather of palaces where “foul things" “intrude", of hypocrisies

’ Shakespeare’s Library, II, pp. 286, 306, 307. Cf. the comment of Violet Jeffrey (see note 3

above), “Giraldi Cinthio, in his version of the tale, supplies no details whatsoever of the town:

yet the scenes of the play set in Venice are packed with local color.”

Wales — The setting of Othello

323

too subtle to be divined save by one versed in the ways of the

world, who “knows all qualities with a learned spirit of human

dealings.”8 In the end we return to the earlier impression and

feel that the honor of Venice has remained inviolate.

8 H. N. Maugham ( The Book of Italian Travel, London, 1903) pronounces Shakespeare’s Italy

uniformly that of the Italian novelists as far as local color is concerned, but adds in a footnote

“except in the character of Iago, who is a typical Renaissance Italian”.

The fact is that Shakespeare, as we should expect, reflects in his plays both contemporary

attitudes, the romantic lure and educational advantages of Italian travel and the patriotic and to

some extent even the protestant and puritan condemnation of its evil influences.

But although he reflects the two aspects of Italy, Shakespeare does not of course subject

them to any sort of scientific historical study. In fact he cheerfully combines and mixes them.

In this respect it is worth while to contrast Othello with the Duchess of Malfi. In Webster’s

play there is a more fully developed Italian atmosphere, as Italy was thought of in England —

for instance, in the corruption of the church, of government, the preponderance of evil, the misuse

of the law, wholesale bribery, the tendency to use crime freely as a means to an end and to make

little of it. Of course, for theatrical purposes, Webster uses also many trappings, conventionally

associated with Italy, but no more realistic than romantic — such as the control of marriage by

the family, the emphasis on banishment, on poison and the dagger, the easy isolation of the

palace, superstitions about drugs and charms, the ancient abbey and the echo, etc. Also he brings

in natural references to place and time, the new fortifications at Naples, “I knew him in Padua”,

“the Cardinal hath made more bad faces by his oppression than ever Michael Angelo made good

ones”; the baths at Lucca, the shrine at Loretto, the citadel of St. Benet, etc. But chiefly

Webster seeks to get an Italian effect by emphasizing the intellectual attitude of the Italian

renaissance, the will, force, intellect of his villains, the intellectual doubt of Antonio and the

Duchess.

Dost thou think we shall know one another

In th’other world? ....

O that ’twere possible we might

But hold some two days’ conference with the dead!

From them I should learn somewhat, I am sure,

I never shall know here. I’ll tell thee a miracle:

I am not mad yet, to my cause of sorrow.

This is the miracle. She does not go mad — nor does Antonio — nor Bosola. They have strong

heads — these skeptics. Julia the “great woman of pleasure” is likewise typical. She too dies

like an intellectualist:

’Tis weakness

Too much to think what should have been done: I go

I know not whither.

In his intellectual attitude Bosola is unlike Iago, less convincing, x though in some ways more

interesting. He knows doubt of his own philosophy. The growth of his doubt is gradual; however,

because of over-condensation, far too swift for us to follow satisfactorily. Throughout the play,

evil is the norm of action and atmosphere. The good is rare and stands alone. The characters

struggle with evil as an intellectual problem, not a single issue. At the end it is not so much

the active triumph of good that we feel as the proved futility of evil. Nothing is gained by

wrong; hence you may dare to be good if you prefer it.

Shakespeare gives us no such sense of general human depravity. Even in Othello, good

appears as the norm; the evil, though real and unconquered, is presented as monstrous.

Fynes Moryson in Shakespeare's Europe (1617, p. 408), expresses the popular view of the

Italian character: “Thus the Italyans being by nature false dissemblers in their owne actions,

are also most distrustfull of others with whome they deale or converse, thincking that no man is

so foolish as to deale playnly, and to meane as he speakes.” For earlier comments cf. Maugham,

p. 14: “Young Englishmen did not always come back entirely improved by this southern

experience. Ascham, the gentle master of Lady Jane Grey, was only nine days in Italy, but he

tells us that he saw ‘in that time, in one city, more liberty to sin, than ever I heard tell of

in our noble city of London in nine years.’ Robert Greene, the dramatist, admits that he ‘saw

and practised on his Italian travels such villainy as it is abomination to describe.’ Sir Philip

Sidney has admitted the dangers of Italy, but remarks that he is acquainted with ‘divers noble

personages . . . whom all the sirens of Italy could never untwine from the mast of God’s word’.”

See Einstein’s account of The Subtlety of the Italian, by F. G. B. A., 1591, a book which

argues that the Italians ought “to be shut up from all access or entrance into other countries. If

such means were adopted, ‘wre no more shall be exposed to the lamentable miseries into which they

were wont to bring us headlong at their own lust and pleasure’.” — Einstein, The Italian Renais¬

sance in England, pp. 170-172. See also Ibid., p. 160 — a quotation from Gascoigne: “George

Gascoigne, in his lines to a friend about travel in Italy, advised him to beware of poison when

invited to dinners, never to drink before another had tasted the beverage, to be on the lookout

for poisoned soap, and take care lest the tailor stuff his doublet with what might bring on a

a deadly sweat. The Italian art of poisoning impressed itself on the Elizabethan imagination and

furnished endless material to the dramatists.” He also quotes Nash, Piers Penniless (p. 38) :

“0 Italy academy of manslaughter, the sporting place of murder, the apothecary shop of all

nations! How many weapons hast thou invented for malice.”

On the subject of infidelity and private vengeance, Fynes Moryson says:

“Adulteries (as all furyes of Jelousy, or signes of making loue, to wiues, daughters, and sisters)

324 Wisconsin Academy of Sciences, Arts, and Letters .

The elements of Venetian setting which receive the most

emphasis thus group themselves about four aspects of the story :

the romantic elopement,9 Othello's official relation to Venice, the

are commonly prosecuted by priuate reuenge, and by murther, and the Princes and Judges, meas¬

uring their just reuenge by their owne passions proper to that nation, make no great inquiry after

such murthers besides that the reuenging party is wise inough to doe them secretly, or at least

in disguised habitts.” — Shakespeare’s Europe, p. 160.

Einstein says ( Tudor Ideals, 1921, p. 123):

“The frequency of vengeance on the stage suggests that this motive as an incentive to crime

was readily understood, but it was associated more with Italy where the absence of central authority

and the inadequacy of the law, favored the wronged individual taking the remedy into his own

hands. . . . Perhaps one reason why the Elizabethan drama save in the greater Shakespearian

masterpieces remains so dead to us, is the lack of contact between modem life and private ven¬

geance. The Englishman of the Sixteenih Century had still enough associations with former

recollections of violence to make the crimes of Italy appear not altogether remote.”

Similarly, Boulting says ( Tasso and his Times, 1907, pp. 182-3):

“The Italian gentleman of the sixteenth century felt certain stains as keenly as wounds; and

the growing respect for female relatives and family pride had this consequence, that any unfaith¬

fulness on the part of a wife or any unchastity on the part of a sister were visiied by the speedy

removal of the suspected lover, and in time it became de rigeur that she also should pay for her

fault with death. The restriction of the power of the nobles to their own domestic circle and

the growth of honor provided the world with a terrible series of family tragedies which struck the

imagination of our English dramatists and gave us ‘Othello’ and the ‘The White Devil,’ and ‘The

Duchess of Malfi’.”

Mr. J. W. Draper (cf. note 3 above) finds a similar attitude among English army officers of

the period.

9 On the subject of Othello’s marriage, some passages from Boulting’s Woman in Italy (1910,

pp. 72, 74) are worth quoting:

“In the middle of the sixteenth century, the Council of Trent published the decision De Sac-

ramenti Matrimonii, which insisted on ecclesiastical marriage and the prior publication of banns.

. . . . Not merely was marriage subjected to family interests, but the State also had a word to

say. ... By reason of the peculiar patrician government of Venice a noble marrying a plebian

woman was excluded from the Venetian Council, until the contract was submitted to the Govern¬

ment and allowed.”

That Desdemona’s difficulties were not unparalleled is seen in the story (given by Boulting,

Woman in Italy, pp. 77-79) of Giulietta Spinoia (Genoa, 1545), who somewhat independently

married the man of her choice. An official inquiry was instituted as to whether she had been

forced into wedlock. She declared that she wished to return to her husband. “The Vicar vainly

endeavored to get her uncles and trustees to accept the marriage, and the question of its validity

was referred to the Archbishop, who decided that it was valid, and, therefore, a sacrament. The

trustees, not to be balked, then appealed to Pope Paul III himself. . . . Meantime Giulietta

was removed, first to another convent, where it was deemed impossible for her husband to hold

any communication with her, and then to the house of a lady of the Spinoia family, where she

was again interrogated by the Vicar. . . . But the spirited girl, determined not to be thwarted,

contrived to make her escape, gained her spouse’s castle and resumed the interrupted honeymoon.”

Mr. J. W. Draper (cf. note 3 above) says of Desdemona:

“Her hybrid origin is surely a secret we are not intended to explore; and so may future

critics continue as heretofore to find her only ‘angelic’ and ‘innocent’ and ‘shy’ and forget, as

Shakespeare wished us to forget, that she wooed a husband for herself, deceived a father and

made him die of bitterness, and then stepped back into child-like innocence at the dramatic behest

of four acts of mighty tragedy.” We are inclined to wonder, however, whether it is not the shy

and innocent girl who might do with simplicity what Desdemona does. Is it not because she is

simple and acts “all of a piece” that she can do it and so sincerely? A more complex woman

would have thought twice. Even Juliet is more complex by nature — though younger.

A distant parallel to Desdemona’s story is found in the story of Bianca Cappello, given by

W. C. Hazlitt, The Venetian Republic (1915), Vol. II. pp. 139-140. “Bianca, sole child and

heiress of Bartolommeo Cappello, a noble Venetian” yielded “to the advances of Pietro Bona-

ventura, a young Florentine of good but poor family, employed as a book-keeper at the Salviati

bank, who resided in a house near the Casa Cappello at S. Apollinare, adjoining the Ponte

Storto. Love letters were exchanged; and Bonaventura, allured by the beauty of the girl and

her probable fortune, . . . persuaded her to elope with him on the night of the 28th November

1563. The fugitives had engaged the services of a gondolier named Girolamo, and had taken into

their confidence the uncle of Bonaventura and three or four others, whose silence or aid they

deemed imperative. . . . They crossed the frontier and reached Florence in safety. Bianca

carried with her all her jewellery.

“The amazing news was spread over the city the next morning. The Council of Ten and

the Avogadors took immediate proceedings; prices were set on, the heads of the principals. . . .

The afflicted parent added a reward of 6,000 lire to that of the government for the recovery of

his misguided child, who was only sixteen years of age at this time.” The rest of Bianca’s story

differs widely from Desdemona’s. The warning “she has deceived her father and may thee”,

unjust in Desdemona’s case, would have been just in Bianca’s.

Wales — The setting of Othello

825

corrupt side of Venetian life, and on the other hand, the dignity

and integrity of the Venetian State.10

Othello' s Memories

As already indicated, Venetian color is, however, not the

only significant element of the setting; another kind of detail is

used in profusion in building up a remoter background to be

seen in the mind's eye only— a world of memory and imagina¬

tion.

We would fain hear Othello “run through'’ the continuous

story of his pilgrimage, the “distressful strokes" that his youth

suffered, the “disastrous chances" of his later years. But (like

Desdemona as she went about her house-affairs) we have only

snatches— besieged cities, capture, slavery, and redemption; his

mother receiving the enchanted handkerchief from the ancient

sybil who devined her thoughts j* 11 the pomp of war, the Pontic

sea and its icy current;12 the encounter with the turbaned Turk

at Aleppo;13 curious peoples in their own lands, caves, deserts,

10 For an account of (lie official machinery of Venice in the sixteenth century, cf. W. C.

Hazlitt, Tke Venetian Republic, Vol. II, Chapter XLVIII: “The provision for the public service

was at once exhaustively comprehensive and jealously minute. No labor, ingenuity or cost was

spared in rendering all the departments of the state, spending and administrative, efficient and

adequate to current wants. A brief survey of the offices and magistratures engaged in the man¬

agement of affairs suffices to impress on us the magnitude of the responsibility and charge, which

gradual conquest and aggrandizement had laid on Venice, as well as the corresponding genius,

which manifested itself for the control and protection of a dominion so extensive and so scattered,

no less than of a territory at home beyond everything precious.” (pp. 448-9.)

Cf. Variorum , p. 43. Lloyd: “Central in the First Act is the scene in the Council Chamber;

and the consideration, by the Duke and Senators, of the news from Cyprus is no mere surplusage;

it strikes a tone of dispassionate appreciation of evidence and opinion that dominates all the

succeeding scenes of agitation and disorders. From inconsistent intelligence, the main point of

agreement is carefully adopted for further examination, notwithstanding predisposition to under¬

rate it; intelligence, otherwise of good authority, is condemned as fallacious from collateral

indications; and lastly, thus prepared for, the last courier has full credence, and the critical

circumstances once understood, action follows at once. Othello is dispatched that very night. The

same solid perspicacity distinguishes the reception of the complaint of Brabantio.”

11 In Shakespeare’s England (II, 485-6) we have an account of the gypsies in England in

Shakespeare’s day:

“It was in the beginning of the sixteenth century that they made their first appearance, and

the mystery of their coming and going was still unsolved. Though they were called Egyptians,

or in derision, Moon-men, there were few who believed in their eastern origin. ‘Ptolomy. I war¬

rant.’ says Dekker, ‘never called them his subjects, no, nor Pharao before him.’ And the same

writer, declaring that their complexion is filthier than the tawney face of a red-ochre man. is

sure, in defiance of the truth, that it is not their own. ‘Yet are they not borne so,’ says he,

‘neither has the Sunne burnt them so, but they are painted so’ ... . They lightly deceived the

common people, ‘wholly addicted and given to novelties, toyes and r.ew fangles;’ whom they

delighted with the strangeness of their headgear, and of whose credulity they took an easy ad¬

vantage. Wherever they went they practiced legerdemain, or fast and loose, they professed a

knowledge of physiognomy, palmistry, and other abuc°d sciences, and by foretelling in the hand

destinies, deaths, and fortunes, they robbed poor counfrv girls of money and linen.”

Shakespeare may have had in mind the gypsies of his own country; but in Othello’s words

he characteristically utilized their Oriental suggestions +o contribute to an Oriental atmosphere.

12 See Variorum, pp. 210. 211, for the passage in Holland’s Pliny on which this is based;

also Swinburne’s eloquent comment.

13 These regions made a strong and familiar appeal to the English mind.

The malignant and turbaned Turk was a real person to the Englishman as well as to the

Venetian. We have for example an account of the escape of John Fox from the captivity of the

Turks in Alexandria. See C. R. Beazley, Voyages and Travels mainly during tke sixteenth and

seventeenth centuries (Arber’s English Garner ), 1903. I. 139-149. Even Englishmen knew what

it meant to be taken captive and “sold to slavery.” See also Ascham’s account (1552) of ex-

326 Wisconsin Academy of Sciences , Arts, and Letters .

and mountain passes, with rocks sharp against the sky.

Though the lines beginning

Wherein of antres vast and deserts idle

are among the most familiar in literature, they never lose their

power to take us by surprise. Shakespeare never saw these

things or anything like them. Whence is the peculiar quality

and coloration of this landscape ? The most satisfactory answer

is that given by Professor H. B. Lathrop,14 who quotes a strik¬

ing passage from a sixteenth-century translation into English

of de Changy’s summary of Pliny’s Natural History:

Towards the west there is a people called Arimaspi, that hath but one

eye in their foreheads, they are in the desert and wilde Countrey. The

people called Agriphagi, liue with the flesh of Panthers and Lyons: and

the people called Anthropomphagi which we call Canibals, liue with hu-

maine fleshe. The Cinamolgi, their heades are almost lyke to the heades

the pillers of Hercules , (among the floudes) there is Onylus that doth in¬

gender Cocodrils. There are goodlye Forrests with vnknowen trees, some

of the which trees beare threades, of the which is made clothing of cotton.

Gyrenes and Syrtes, make their houses of salt stones cut out of the moun-

taines, there is the mountaine of Ciry , the which doth ingender and bring

forth many precious stones. In Libie, which is at the end of the Ethiopes ,

there are people, differing from the common order of others, they haue

among them no names, and they curse the Sunne for his great heate, by

the which they are all black sauing their teeth, and a little the palme of

their handes, and they neuer dreame. The others called Troglodites haue

Gramantes , they make no mariages, but all women are common. Gampha-

santes they go all naked. Blemmyis is a people so called, they haue no

heades, but haue their mouth and their eyes in their breastes.

changes of Turkish and Christian atrocities: The English Works of Roger Ascham, Cambridge,

1904, pp. 130, ff.

For interesting passages on these regions, including Aleppo, see the narration of John Eldred,

“the first Englishman who reached India, overland, 1583-1589.” (See Beazley I, 295-303). See

also Shakespeare’s England , I, Chapter VI, passim ; also the bibliography of this chapter. See

also H. C. Hart’s introduction to the Arden Edition of the play (1903, 1928) for the parallels in

Holland’s Pliny.

Among the many books listed in the bibliography of Care' Howard. English Travellers of the

Renaissance (1914) is that by George Sandys’ published in London in 1615, entitled A relation of

a journey begun An. Dom. 1610. Four Bookes. Containing a description of_ the Turkish Empire,

of AEgypt, of the Holy Land, of the Remote Parts of Italy and Islands adjoyning. This book is

too late of course to be a source of any of the allusions in Othello. But it testifies to the popular

interest in the regions which were the obscure background of Othello’s adventures. Sandys de¬

scribes the Euxine Sea (pp. 39-40) and tells how the Bosphorus “setteth with a strong current

into Propontis.” He describes the habits and dress of the “turban’d Turks.” He tells of the

slave markets. Their slaves “are Christians taken in the warres. or purchased with their money.

Of these there are weekly markets in the Citie, where they are to be sold as horses in Faires;

the men being rated according their faculties, or personal abilities, as the women for their

youths and beauties.” He says (p. 69) that “with their aspects of pity and affection” they

"endevour to allure the Christians to buy them, as expecting from them a more easie servitude

and continuance of their religion. . . . But gally-slaves are seldome released, in regard of their

small number, and much employment which they have for them.”

14 Henry Burrowes Lathrop. Translations from the Classics into English, from Caxton to Chap¬

man. University of Wisconsin Studies in Language and Literature, no. 35 (1933), pp. 219-20.

327

Wales — The setting of Othello

“Here,” says Professor Lathrop, “within the spaces of two

pages, is everything whereof it was Othello's ‘hint to speak'.”15

Everything, we answer as we read, except the perspective of the

picture. Could it have been from the flat surface of these pages

that Shakespeare lifted up his eyes to its depths and distances?

Mr. Lathrop promptly answers our question : “Nothing is omit¬

ted but the loftiness of the hills, ‘whose heads touch heaven.’

And even this omission is but natural. The region named —

though the geography extends further — is Africa, which ‘doth

contain the Moors'-— Othello's own country, and the hills whose

heads touch heaven, as Shakespeare's Ovid would suggest, are

the summits of Mount Atlas itself, bearer of the skies, the loft¬

iest mountain in the land of the Moor, Othello.”

Let us not forget, however, that the scope of the pilgrimage

includes not only strange and shadowy lands, but a world more

tangible, if no less romantic: one familiar to Venetian traders,

travellers, and warriors. And so in the final allusion to Aleppo16

the two main elements of the setting come suddenly and sharply

into relation, and the sweep of memory is indissolubly linked

with Othello's loyalty to Venice.

II

The Emphasis

Two Noble-Barbarian Theories

It is usually best to appeal first to the structure of a play for

light on its emphasis.

Professor Bradley says

Of all Shakespeare’s tragedies, . . . Othello is the most painfully ex¬

citing and the most terrible. . . . Othello is not only the most masterly of

the tragedies in point of construction, but its method of construction is

unusual. And this method, by which the conflict begins late, and advances

without appreciable pauses and with accelerating speed to the catastrophe

is a main cause of the painful tension just described.17

15 See also his earlier article: “Shakespeare’s Anthropophagi. The Source of the Travel’s

History of Othello.” The Nation. 100. Ja. 21, 1915. 76-77.

The scattered parallels with Holland’s Pliny noted by. Hart, Arden Edition, 1903, pp. 26, 39,

etc., though in themselves significant, are much less convincing than this massing of the material

within two pages. For the well-known passage from Sir Walter Raleigh {The Discoverie of

Guiana, 1596; p. 85, Ed. Hakluyt Soc.) usually associated with the lines, see Variorum, p. 56.

16 And here we must recall to the reader’s mind a note provided by Professor Parrott in the

Tudor Edition of the play (1928, p. 168):

“The Venetians had special trading privileges in this town. If the Turkish law that the

Christian who struck a Turk must either turn Turk or lose his right arm prevailed there, Othello

risked his life to uphold the honor of Venice.”

17 Shakespearian Tragedy, pp. 176, 177. Cf. Ibid., pp. 64-7.

328 Wisconsin Academy of Sciences , Arts , and Letters .

Professor Bradley lays a very considerable stress on Othello’s

jealousy; at the same time he does not overlook the importance

of disillusionment as a primary factor in the tragedy; and he

makes much of the point18 that Othello is

not easily jealous, but, being wrought,

Perplexed in the extreme.

Professor Bradley vigorously dissents from the theory that

the play is “primarily a study of a noble barbarian, who has be¬

come a Christian and has imbibed some of the civilization of his

employers,” and that the last acts “depict the outburst” of his

Moorish passions “through the thin crust of Venetian culture”.

Moreover, while admitting that Othello’s race has its importance

in the play, he says,

But in regard to the essentials of his character, it is not important;

and if anyone had told Shakespeare that no Englishman would have acted

like the Moor, and had congratulated him on the accuracy of his racial

psychology, I am sure he would have laughed.19

In the next paragraph, however, he goes on to say,

He [Othello] does not belong to our world, and he seems to enter it

we know not whence — almost as if from wonderland. There is something

mysterious in his descent from men of noble siege; in his wanderings in

vast deserts and among marvellous peoples. . . . And he is not merely

a romantic figure; his own nature is romantic. ... He has watched with

a poet’s eye the Arabian trees dropping their med’cinable gum, and the

Indian throwing away his chance-found pearl; ... So he comes before

us dark and grand, with a light upon him from the sun where he was

born . . . grave, self-controlled, ... at once simple and stately in bearing

and in speech, a great man naturally modest but fully conscious of his

worth, proud of his services to the state, unawed by dignitaries and un¬

elated by honours, secure, it would seem, against all dangers from without

and all rebellion from within.20

But do not these glowing phrases of Professor Bradley put

before us again the idea of the noble barbarian which we were

bidden to discard?21 We cannot of course accept any view which

1S Op. cit., p. 186, p. 194. Cf. Sir Walter Raleigh, Shakespeare ( English Men of Letters

Series ) 1907, p. 204: “Jealousy and suspicion, as Desdemona knows, are foreign to his nature;

he credits others freely with his own noblest qualities.”

19 pp. 186, 187.

20 pp. 187-189.

21 The words

Like the base Indian cast a pearl aside

Richer than all his tribe

may be of considerable significance. For a discussion on which reading Indian or Judean is right,

see Variorum, pp. 327-331. See also a note in the Yale Edition, p. 143, and a note in the Tudor

Edition, p. 167. Obviously the view presented in the present paper accords best with the reading

Indian.

Wales — The setting of Othello

329

would make Venetian civilization responsible, broadly speaking,

for the good in Othello, and his Moorish blood responsible for

the evil. May it not be possible, however, to make the idea of a

noble barbarian the basis of an almost opposite theory, — namely,

that the good in Othello is a native good , and that his temporary

overthrow comes from the failure of a mistakenly idealized civi¬

lization?

The conception of the noble barbarian as in some ways su¬

perior to the more civilized and less natural man was current to

some extent in the Renaissance, and is found in explicit form

in Montaigne's essay on Cannibals — an essay with which we

have other reason to believe Shakespeare was familiar.

"There is nothing in that nation/' writes Montaigne, "that is

either barbarous or savage, unlesse men call that barbarisme

which is not common to them. . . . Those nations seeme there¬

fore so barbarous unto me, because they have received very

little fashion from humane wit, and yet are neere their originale

naturalitie. . . . The very words that import lying, falsehood,

treason, dissimulation, covetousness, envie, detraction, and par¬

don, were never heard amongst them," etc.22

Thos. Palmer has an interesting passage bringing in the idea

of "the noble barbarian" :

So also is it to be understood, that no nation in the world, how Court¬

like soever, but hath the dregs and lees of barbarous incivility; and that

many heathen people, by the light of nature meerly inscribed in their

hearts, rest for ensamples and reproofes to many civill nations governed

by a diviner knowledge, in points of civil actions & conversation.23

Montaigne’s essay expresses the appeal of New World

discovery to the Renaissance imagination. The most striking

product of this appeal in Shakespeare is The Tempest . Yet is

it not possible that we find it in another and in some ways a

more vital aspect, in Othello ? As we have elsewhere24 empha¬

sized, the Englishman of Shakespeare’s day was reacting to two

diverse influences, the stimulus that came from Italy, of a riper

22 Essay of Cannibals, Florio’s Translation.

Mr. George Coffin Taylor ( Shakespeare’s Debt to Montaigne, 1925, p. 32) makes this signifi¬

cant observation: “In Othello, written in 1604, when one would naturally expect to find Mon¬

taigne’s influence at its height, it is scarcely discernible. . . . The scant influence on Othello is

more easily accounted for by the nature of the particular play, in which Shakespeare seldom

introduces matter not germane to the plot or situation.”

23 Thos. Palmer, An Essay of the Meanes how to make our travailes into forraine Countries,

the more profitable and honourable. London 1606, p. 62.

24 “Shakespeare’s Use of English and Foreign Elements in the Setting of The Two Gentlemen

of Verona,” in Transactions of the Wisconsin Academy of Sciences, Arts and Letters, Vol. 27

(1932), pp. 93-4.

330 Wisconsin Academy of Sciences , Arts , and Letters.

culture and a more self-conscious society, and that which came

from the unknown world beyond the Atlantic, and perhaps the

most curious and thought-provoking aspect of these two influ¬

ences is that of the impact of one on the other.25 Do we per¬

haps discern something of this aspect in the tragedy of Othello ?

It is not necessary of course to assume any direct connection

between Montaigne's essay and Shakespeare's play, nor by any

means to insinuate that Othello was after all neither a Moor nor

a Blackamoor, but a North American Indian. So far as any

special barbarian race is concerned, we must agree with Pro¬

fessor Bradley that Shakespeare had no idea of attempting a

study of racial psychology. Yet is it not possible that Mon¬

taigne's essay, together with the plot of Cinthio's novel, may

have suggested to Shakespeare the character of Othello, the

general conception of a noble barbarian, the type of a strong and

in some ways mature man, who was, nevertheless, in the presence

of the complex and more or less corrupt civilization of sixteenth

century Europe, a child and a stranger ?

While he says that “we must not call the play a tragedy of

intrigue as distinguished from a tragedy of character," Profes¬

sor Bradley is struck by the fact that “Iago’s intrigue occupies a

position in the drama for which no parallel can be found in other

tragedies . . . ."26 “The part played by accident in this catas¬

trophe," he says again, “accentuates the feeling of fate." And

again, “It confounds us with a feeling .... that .... there

is no escape from fate, and even with a feeling .... that fate

has taken sides with villainy."27

For further light on these impressions let us turn back to

Professor Bradley's lecture on the substance of tragedy.

“How is it," he asks, “that Othello comes to be the companion

of the one man in the world who is at once able enough, brave

enough, and vile enough to enslave him? By what strange fatal¬

ity does it happen that Lear has such daughters, and Cordelia

such sisters? Even character itself contributes to these feelings

of fatality. How could men escape, we cry, such vehement pro-

pensites as drive Romeo, Anthony, Coriolanus, to their doom?

25 Cf. for interest in the New World, Rachel M. Kelsey, “Indian Dances in the Tempest”,

Journal of English and Germanic Philology, XIII (1914), pp. 98-10C5.

See also, Robert Ralston Cawley, “Shakespeare’s Use of the Voyagers in The Tempest*9

PMLA, XLI (1926), pp. 688-726.

28 Shakespearian Tragedy, p. 179.

27 pp. 181, 182.

Wales - — The setting of Othello

331

And why is it that a man's virtues help to destroy him, and that

his weakness or defect is so intertwined with everything that is

admirable in him that we can hardly separate them even in im¬

agination ?”28 If these questions indeed have a rational answer

at all it would seem to be: Because the world is so constructed

that all men must learn; it is not enough to say — as Professor

Bradley himself does say29 — -that the vital principle of growth is

destructive of all that is evil; it is destructive of all that is in¬

complete, or rather, of all incompleteness. Therefore it is artis¬

tically true to place beside Othello that being who— for the pur¬

pose of dramatic condensation — is best fitted to destroy Othello's

ideal. Professor Bradley touches this concept, though with a

difference.

These defects or imperfections are certainly, in the wide sense of

the word, evil, and they contribute decisively to the conflict and catastro¬

phe. And the inference is again obvious. The ultimate power which

shows itself disturbed by this evil and reacts against it, must have a

nature alien to it. Indeed its reaction is so vehement and “relentless” that

it would seem to be bent on nothing short of good in perfection, and to be

ruthless in its demand for it.30

Whether the imperfections are to be called good or evil would

seem to depend, however, on the direction from which they are

approached— whether from an inferior or a superior plane. An

identical act or attitude may represent either ethical progress or

ethical retrogression.

The moral nature of a man grows by the process of the fail¬

ure of inadequate desires or ideals and the construction of larger

ones. The collapse of an ideal is sometimes attended by moral

prostration. Each of Shakespeare's tragedies presents the fail¬

ure of an ideal or attitude to life and the attendant moral pros¬

tration — these being expressed through the dramatic medium of

crime or error and outward calamity. To many minds some at

least of Shakespeare's tragedies imply also a sense of moral

triumph or the foreshadowing of the reconstruction of the indi¬

vidual ideal.

“Nor .... are the facts ever so presented," says Professor

Bradley, in speaking of the sense of fate in Shakespeare's plays,

28 p. 29.

20 “Yet it appears to engender this evil within itself, and in its effort to overcome and expel

it it is agonized with pain, and driven to mutilate its own substance and to lose not only evil

but priceless good.” — p. 38.

30 p. 35.

332 Wisconsin Academy of Sciences, Arts, and Letters.

“that it seems to us as if the supreme power .... had a special

spite against a family or an individual.”31

No ; on the contrary it would seem that the supreme power

finds it worth while to complete the individual— never to let him

off without putting him through the painful process of the de¬

struction of his illusions. In the various tragedies of Shakes¬

peare we feel varying degrees of sympathy for the central figure

and of blame for his mistakes. The essential point is this : that

Shakespeare — like natural law — does not seem to distinguish in

his catastrophes between sins of ignorance and more deliberate

crimes. Othello merely stands as Shakespeare’s extreme instance

of disaster which must sometimes come upon even those who

have acted in accordance with the dictates of a perfect — though

limited — soul.32

Granted this general view of the play, it is obviously no acci¬

dent that the construction is peculiar — that is, that the tragedy

begins late — or that the action and catastrophe depend upon in¬

trigue. Since this is a tragedy of character, since it not what

Othello does that is his ruin, but what he is, it is all-important

that we be made to grasp his normal character. Hence the long

exposition. As for the intrigue, it is a dramatic concentration

of forces that are bound to act sooner or later for Othello’s en-

lightment. The enlightment may come suddenly or by degrees, —

through the untruth of one man, or of many. In any case, and

this is the point to be borne in mind, it must be, as far as Oth¬

ello is concerned, in a sense accidental, — not due to his deliberate

fault, but in the nature of the universe, inevitable. Hence the

tragedy of Othello has quite as much universal truth as the other

tragedies, since it is an example of well-intentioned human na¬

ture adjusting itself to the knowledge of good and evil. It is

preeminently the tragedy of disillusionment— the disillusion¬

ment of a noble barbarian with a somewhat decadent civiliza¬

tion which he had simple-mindedly venerated. If Montaigne’s

noble barbarian were transferred to civilization, what would

become of him? At what terrible price would his adjustment be

made ? He would believe in the world too much at first ; he would

be bitterly disappointed, losing all faith ; then he would find that

after all what he had loved best was true. Confidence in himself

31 p. 29.

32 Cf. Raleigh, Op. cit. p. 198: “Othello, like Hamlet, suffers for his very virtues, and the

noblest qualities of his mind are made the instruments of his crucifixion.”

Wales— The setting of Othello 333

and the world; disillusionment; reconstruction: that is the tra¬

gedy of Othello.

The Second Theory and the Action

Let us briefly review the action from, this standpoint. With

his usual theatrical wisdom Shakespeare opens the play with a

scene tending by every method of suggestion to prejudice us

against Othello at the outset. We are expecting him to be a bar¬

barian ; we are prepared not to apply to him the standards of

civilization. When the real Othello comes upon the scene, we at

once become his advocates and tend to be over-lenient with his

faults. So far from having to make allowances for him we feel

that he is superior to his European surroundings. Othello's

first words “ Tis better as it is”— -referring to lago’s boasted

wish to punish Roderigo— present him to us as a person of au¬

thority, just but generous. In answer to lago’s further insinua¬

tions, irritating as they are meant to be, he speaks calmly :

Let Mm do his spite:

My services which I have done the signiory

Shall out-tongue his complaints. Tis yet to

know,

Which when I know that boasting is an honour

I shall promulgate, I fetch my life and being

From men of royal -siege, and my demerits

May speak unbonneted to as proud a fortune

As this that I have reached.

On the lines

Good signior, you shall more command with words

Than with your weapons

Sir Walter Raleigh makes this comment :

Fearlessness and the habit of command, pride that would be disgraced

by a street brawl, respect for law and humanity, reverence for age, la¬

conic speech and a touch of contempt for the folly that would pit itself,

with a rabble of menials, against the General of the Republic and his

body-guard— -all this is expressed in two lines.33

To Brabantio's insulting charge he replies reasonably as to a

fractious child,

What if I do obey?

How may the duke be therewith satisfied,

Whose messengers are here about my side,

Upon some present business of the state,

To bring me to him?

33 Shakespeare ( English Men of Letters Series ), 1907, p. 141.

384 Wisconsin Academy of Sciences , Arts , and Letters .

His ceremonious words to the Senate — far from being a mere

form — express genuine confidence and veneration. No forms

are mere forms to Othello. He takes the civilized world ser¬

iously, regards its institutions with reverence, and expects of it

a sincerity equal to his own. His respect for himself, his re¬

spect for others, and his modesty are all essentially related; it

is of the essence of his pride to admit readily his little knowl¬

edge of the world.

Rude am I in my speech

And little blessed with the soft phrase of

peace. . . .

And little of this great world can I speak,

More than pertains to feats of broil and battle.

Yet the consciousness of his ignorance causes him no doubt of

his own perfect soul. Moreover while he has the restraining

sense of fitness which we noticed before, he loves the sound of

his own words. His vivid imagination — a supposedly child-like

and barbarian quality — takes fire the moment he begins to speak

of “antres vast and deserts idle.” He is satisfied with the part

he has played in these adventures. To reflect upon them gives

him pleasure — a pleasure of which he does not think of being

ashamed. The speech makes a favorable, even a delightful im¬

pression upon the Duke. Othello succeeds in justifying his mar¬

riage in the eyes of Venice. In the first act he comes off vic¬

torious, having behaved with tact, wisdom, decision, courage,

unfailing courtesy, unruffled generosity. He has satisfied others,

and in no respect disappointed himself.

By the beginning of Act II, we have advanced far in our ac¬

quaintance with Othello, having seen him face to face with cir¬

cumstances which — though far from tragically serious — were

fairly critical, testing his resource and self-control. His pre¬

monition of evil, early in the second act, is no misgiving of fail¬

ure in himself or his world— only a superstitious dread of the

irony of fate, a passing thought, natural enough to one who is

happy and who has imagination enough to think of himself as

bereft of his happiness. He looks forward with frank pleasure to

renewing old acquaintanceships in Cyprus, but remembering his

standard of good manners, checks his too enthusiastic speech

and turns to give orders to his attendants, punctiliously consid¬

erate of every one, and generous and glad in his recognition of

Wales — The setting of Othello

335

every good quality. Professor Bradley points out that in the

third scene of Act II Othello’s self-control is emphasized and

that here “occur ominous words which make us feel how nec¬

essary was the self-control and make us admire it the more.”34

Now, by heaven,

My blood begins my safer guides to rule,

And passion, having my best judgment collied,

Assays to lead the way.

They indicate not only Othello’s self-control, however, but also

his clearly defined theory of self-control, the fact that it is a

part of his deliberate ideal.

Good Michael, look you to the guard tonight:

Let’s teach ourselves that honourable stop,

Not to outsport discretion.

And later in the scene he shows again his veneration for Chris¬

tian institutions, and his single-minded horror of whatever is

barbaric and below the ideal of civilized life.

Are we turn’d Turks, and to ourselves do that

Which heaven hath forbid the Ottomites?

For Christian shame put by this barbarous brawl. . . .

How comes it, Michael, you are thus forgot?

Cassio is abashed by this reproof and can find no words. Othel¬

lo’s reproof of Montano is also characteristic.

Worthy Montano, you were wont be civil;

The gravity and stillness of your youth

The world hath noted, and your name is great

In mouths of wisest censure: what’s the matter,

That you unlace your reputation thus

And spend your rich opinion for the name

Of a night-brawler ? Give me answer to it.

Othello’s personal sense of the enormity of the offense — the

want of consideration of the public peace — shows that he is still

unaccustomed to evil, especially in civilized men, and discerns it

with surprise and pain. He feels it not only his military but

his moral duty to be very severe.

Give me to know

How this foul rout began, who set it on.

190.

336 Wisconsin Academy of Sciences, Arts, and Letters .

In the early part of Act III we have little new light on the

character of Othello. In the first scene he does not appear. In

the brief second scene we have a glimpse of him in his post of

authority, occupied with his official duties. Not until the mid¬

dle of the act does the tragedy itself begin.

lago’s first task is to cause Othello to distrust human nature.

He sows first a distrust of Cassio, a general suspicion that he

is not truthful, that he is given to drink and brawling, then that

his whole relation to his chief has been one of darkest duplicity.

Next lago skillfully opens Othello’s eyes to the true nature of

life in Venice and the capacities for evil hidden in the bosoms

of super-subtle Venetians. And after this it is easy to sow a

greater doubt in the Moor’s trusting soul. He has been deceived

in much, why not in more? The general fact of his ignorance

of human nature, especially feminine and Venetian human na¬

ture, having once been thoroughly brought home to him, he

abandons himself to Iago’s superior knowledge.

This fellow’s of exceeding honesty,

And knows all qualities, with a learned spirit,

Of human dealings.

The mingling in Othello of credulity with susceptibility to doubt

is psychologically true to life.35 His intense imagination once

at work, Othello’s suspicion becomes a part of him. The devas¬

tation is complete, not only of his recent self but of his past self

as well.

0! now, for ever

Farewell the tranquil mind; farewell content!

Farewell the plumed troop and the big wars. . . .

. . . .Othello’s occupation’s gone!

Yet nothing is actually proved. The struggle begins again, to

end quickly in despair.

In the fourth act Othello reaches the lowest point in his hu¬

miliation. He loses all sense of personal dignity and of respect

for others. He betrays his jealousy and strikes his wife in the

presence of incredulous spectators. Professor Bradley cannot

reconcile himself to the blow. Yet if the theory here put for¬

ward be correct, not only the blow, but the fact (noted by Pro-

35 Cf. Raleigh: Op. cit. p. 204. “If he were less credulous, more cautious and alert and ob¬

servant, he would be a lesser man than he is and less worthy of our love.” P. 20S: “There is a

horrible kind of reason on Othello’s side when he permits lago to speak. He knew lago, or so he

believed; Desdemona was a fascinating stranger. Her unlikeness to himself was a part of her

attraction: his only tie to her was the tie of instinct and faith.”

Wales— The setting of Othello

337

fessor Bradley) that it occurs in the presence of the Venetian

representative is absolutely central in the interpretation of the

tragedy. Its dramatic necessity can be appreciated only by ref¬

erence to the situation in the first act: Othello’s confidence in

himself in the presence of the Senators, his faith in his own

worth and dignity as their loyal servant, his chivalry for his

wife, his simple-minded emphasis on good manners. Again and

again in the play we are made to realize that Othello has a sim¬

ple and noble ideal of good manners as a genuine indication of

high-mindedness. This is part of his worship of civilization.

When he loses his faith in civilization, he loses his manners. To

him manners were not merely outward accessories or matters

of empty, mechanical habit. They were deliberate actions de¬

pending on a conscious state of mind. That is why he could

fail in them. The civilized man receiving his conventions ready¬

made from tradition controls himself automatically, and a de¬

gree of outward self-control does not necessarily mean a pro¬

portionate security from inward collapse. Othello was a genu¬

ine gentleman, but not a mechanical one. Hence the blow and

hence its tragedy. Jealousy and violence are mere indications

of the crash of his universe. He is overthrown to the extent of

failing in his consciously strongest points. In these strong

points, he has placed a confidence which, even in the best of

men, argues incomplete experience. Yet in one sense we are

less in despair of Othello than if he had been capable of saving

his personal dignity and keeping his reproaches for a private

moment. Reputation and honor, hitherto his ruling passion, are

forgotten in a frenzy of anguish.

In the third scene of Act IV Othello has partly recovered

himself and displays the calm of definite resolution. He treats

Lodovico with courtesy and speaks less harshly, perhaps even

kindly, to Desdemona. Emilia observes that “he looks gentler

than he did.” Although in the scene of the blow he has tem¬

porarily ceased to care about his personal dignity, it is clear

that one thing remains to him-— a sense of abstract justice, the

need to avenge that personal honor in which he still believes, to

punish the wrong-doers and to vindicate the right. Though bit¬

terly disappointed in others he still has faith in his duty and his

power to perform it. Othello’s belief as to what his duty was

must simply be taken for granted,36 although it is one that we

338 Wisconsin Academy of Sciences , Arts , and Letters.

*.':v iX'j'vv.

cannot understand at the present day. When he knows Desde-

mona’s guilt, no question enters his mind as to what is to be

done next. And before we blame him for his want of reflection,

we must remember that he is not, like Hamlet, amply supplied

with the materials of thought.37 From his point of view there

is nothing more to consider. Nowhere has Shakespeare entered

more completely into an experience and at the same time wholly

shut off from his consciousness those elements which could not

enter into a mind infinitely simpler than his own. Othello's

quickness of action is the inevitable result of his limited knowl¬

edge and his perfect simplicity, sincerity, and certainty of him¬

self. Relentlessly conquering his grief and pity, he murders his

wife from a conviction of right, without a qualm as to the jus¬

tice and necessity of the deed.

Then comes his appalling enlightenment.

At this crisis he is not even able to avenge himself upon his

destroyer. He runs at lago, who evades the stroke. Montano,

fearing that the Moor may do himself harm, wrenches his wea¬

pon from him.

I am not valiant neither,

But every puny whipster gets my sword.

But why should honour outlive honesty?

Let it go all.

As far as Othello’s estimate of himself is concerned, these words

mark the lowest point of his moral prostration. They are also

the beginning of his triumph. He knows the world now and

himself. In lago he has had an overwhelming revelation of evil.

He himself is no wise and moderate man, but one who “like the

base Indian threw a pearl away, richer than all his tribe.” Yet

in Desdemona he has recovered all, so that he can even be just

to himself. He has done naught in haste, but all in honor, and

one way to vindicate that honor still remains.

Set you down this;

And say besides, that in Aleppo once,

Where a malignant and a turban’d Turk

Beat a Venetian and traduced the State,

I took by the throat the circumcised dog,

And smote him, thus.

36 Cf note 8 above — material from Moryson, Einstein, Boulting, and Draper.

37 cf Raleigh- Op cit. p. 17. “A measure of the subtle speculative power of Hamlet

might have saved Othello from being a murderer; it could not have increased Shakespeare s love

for him.” Compare the delineation of Hotspur and Coriolanus— also simple-minded warriors and

men of action.

Wales — The setting of Othello

339

He will prove himself still loyal to Venice and the trust she

has reposed in him, to his conception of the State and civiliza¬

tion,-— that which is noblest in him ready to do swift execution

upon whatever has betrayed the ideal. So dies Othello, the Moor,

triumphant in Desdemona’s truth and in the sincerity of his own

perfect soul.

The Setting and the Total Effect

The elaboration of the setting has thus served a definite end,

since against its delicate and colorful network, as against the

stained fragments of a mosaic background, has been projected

in boldest relief the large and simple figure of the Moor. The

use of detail of place has done much to present the characters

with solidity to the spectator and to create the dramatic situa-

in which they move. It has contributed to their inner life also.

Some modern critics see Shakespeare's treatment of Othello

(and of many other characters) as for the most part plastic and

external, and contrast this method with what they consider to

be the modern method of conscious psychological analysis. And

yet we wonder whether contemporary and older dramatists alike

do not at their best employ a method which is neither of these.

Coleridge says, “One of Shakespeare's modes of creating char¬

acters is to conceive any one intellectual or moral faculty in mor¬

bid excess, and then to place himself, Shakespeare, thus muti¬

lated or diseased, under given circumstances." The process is

not deliberate or self-conscious, however, but instinctive and

imaginative. Most persons who have tried to write dramatic

dialogue know a little of it by experience. If all personalities

are latent in any personality, can be, as it were, realized from

something in oneself, and if, in order to produce a single one of

these, one can check off all in oneself that is foreign to its con¬

sciousness, let down shutters on the rest of one's mind, then

what remains for use and development is not a constructed, in¬

organic thing, but something almost as organic as the drama¬

tist's own mind, though on a different scale. And hence in great

drama, out of the issues of the heart the mouth speaketh.

The essentials of character may, as Professor Bradley im¬

plies, be independent of specific time and place. Yet, however

transferable, they are most easily realizable to us in a vivid set¬

ting. Shakespeare has in Othello realized an alien personality

840 Wisconsin Academy of Sciences, Arts, and Letters.

in an alien place and has somehow assimilated to Othello's inner

experience the outer details of his environment and made these

symbolic to us of his inner conflict.

Thus Italy in this play serves not merely to represent ro¬

mance, nor primarily a realistic contemporary background. It

is the sophisticated world, rather, in its complexity, at first

idealized, then found to have abysmal possibilities of evil, finally

restored to its human proportions. Othello’s memories, too, stuff

of poetry as they are, serve dramatic ends, being images that

express to us the Moor’s simplicity and the gamut of his exper¬

iences, his sense of being equal with his world, his perplexed

sense of being unequal to it, and finally, even in his humiliation

and tragic remorse, his sense of repudiating its evil and being

at one with its good.

Othello is a barbarian in a general rather than a specific

sense. Romantically he is the Moor; but if we try to locate

him more realistically, we shall find him in many lands and

under many disguises. The suggestion for the type came, no

doubt, from innumerable sources: indirectly, through the inci¬

dents, from Cinthio; perhaps (and if so, more directly) from

Montaigne ; and at once more generally and more vitally, from

any honest soul trying to adjust a simple ideal to a complex en¬

vironment, — Valentine at the Emperor’s court, the Englishman

in Italy, Shakespeare himself going up to London.

THE LITERARY GERMAN LANGUAGE AND ITS

RELATION TO THE GERMAN DIALECTS

Alfred Senn

By the term “Literary German Language" we understand

that standard form of literary German which is used as a com¬

mon means of expression and communication in written or

printed form by all German speaking people whether they live

in Germany proper, in Austria, Czechoslovakia, Switzerland,

Alsace-Lorraine, or wherever else. This is the form of the Ger¬

man language that is taught in German schools. It is the only

connecting link that unites all the members of the German

speaking world.

This standard form of the German language is also that

German language which is being learned by foreign students.

Foreign students who have never been in a German country

are sometimes inclined to regard this form of German as the

only one. This error is especially likely to appear in countries

having a more or less unified colloquial language, e.g., in the

United States. Here English is spoken everywhere in almost

entirely the same way, and those who speak of the existence of

various American English dialects seem to overemphasize the

differences. This is at least inevitably the impression gained

by a person familiar with the German dialects. By this I cer¬

tainly do not want to minimize the dialectal differentiations in

American English. I only want to show that the differences

existing between the German dialects are by far more impor¬

tant. These differences are so enormous that speakers of the

extreme parts of the German territory, if they come in contact,

are not able to understand each other if they have not been

trained in the use of the standard form of German. That could

be easily illustrated here in Madison by confronting two Ger¬

mans both born in America and not educated in German schools,

but one of Low German and the other of Swiss, i.e. High Ger¬

man, especially Upper German, origin. Even if both of them

speak their mother tongue to perfection they will not be able to

understand each other and will be forced to resort to the use of

English.

341

342 Wisconsin Academy of Sciences, Arts, and Letters.

This little excursion shows clearly chat standard German or

literary German has to be acquired even by German people. This

acquisition is a permanent struggle against the local form of

speech, the so-called dialect. Some succeed in overcoming the

immense difficulties, but others do not. The question has been

raised again and again which part of the linguistic German

territory uses the best form of German, in other words, which

local form of speech could be regarded as a model and therefore

imitated. The answer to this question has varied according to

the time and the political or geographical unit in which the liti¬

gant parties lived. These linguistic debates were especially ve¬

hemently waged at the time of the German Sprachgesellschaften

in the 17th century. In more recent times, however, this ques¬

tion seemed to lose importance. It seems as if the German

people have become more tolerant toward local expressions. So

we find not only single dialectal expressions but also entire sen¬

tences and even long drawn-out dialogue in dialectal form in

the works of such outstanding writers as Auerbach, Anzengru-

ber, Sudermann, Maria von Ebner-Eschenbach, and Gerhard

Hauptmann. This tolerance shown with regard to idiomatical

expressions of limited spread is to be explained as a reaction

against the classical standard language. At the same time it

proves that the unification of the literary German language has

been accomplished. The German writers feel so sure of this

accomplishment that they see no danger in cultivating local

forms. This reaction needs an explanation. The explanation

is easy to find if we realize that this linguistic step backwards

accompanied a new literary wave, the so-called realism. The

realistic writers felt the wide gulf that existed between the

written or printed form of standard literary German and that

which was used in colloquial everyday speech. They realized

that the modern literary German language is scarcely more

than a mere fiction. It is merely a written language not actu¬

ally being spoken anywhere. Therefore realistic writers wish¬

ing to depict real life began to use such forms and expressions

that were truer to the standards of living of their characters.

I have just stated that the modern literary German language

is not actually spoken anywhere. What is actually spoken by

most of the cultivated German people is a form of speech that

stands between the literary standard form and the dialect. In

Senn — Literary Language and German Dialect . 343

some parts of Germany this elevated form of colloquial speech

is closer to the literary standard, in other parts, especially in

the South, it comprises more local dialectal elements. In Swabia

and Austria, e.g., the colloquial speech of the higher social stra¬

tum is rather more a unified Swabian or Austrian dialect than

an adapted form of the standard language. The extreme in this

direction is represented by the Swiss. Even today there exists

nothing that could be called “common colloquial Swiss”. And

when in 1921 a Swiss scholar, namely Karl Stucki, undertook

to write for practical use a grammar of the German Swiss lan¬

guage,1 he was compelled to confess that the title of his book

was misleading. In his introductory remarks he admits the

impossibility of writing a practical grammar embracing the

whole German Swiss territory, because even the most cultivated

inhabitants of such cities as Zurich, Basel, Bern, and St. Gallen,

represent four entirely different linguistic types. Stucki’s Swiss

grammar may be regarded as a first attempt toward leveling

out the dialectal differences on the basis of the dialect of Zurich.

Now to come back to the question of correctness we see that

there are two sides to it: (1) the correctness of the written

language and (2) the correctness of pronunciation. As to the

first one, it is clear and has already been stated above that we

possess now a strict common norm for written expression,

which only in rare cases permits a choice between two equally

recognized forms.2 In regard to the second side, there exists

still great variety.

Standard literary German is however not the only form of

language used for literary purposes. In all periods of literary

production we find writers who refused to use the generally

adopted standard language. A large dialect literature was de¬

veloped especially during the last century and particularly dur¬

ing the last few decades. It must however be duly emphasized

that not all of the so-called dialect literature represents pure

dialectal forms. We distinguish three types:3

(1) The literature which is mainly based on the standard

literary form, but abundantly interspersed with dialectal forms

and idiomatical expressions of dialectal character. The best

known author of this type is the Swiss Jeremias Gotthelf. In

1 Schweizerdeutsch. Abriss einer Grammatik mit Laut- und Formenlehre. Zurich 1921.

2 Cf. Paul, Deutsche Grammatik I 129 f.

3 Cf. Hans Reis, Die deutsche Mundartdichtung. 1915. Sammlung Gdsche'n. p. 10 f.

344 Wisconsin Academy of Sciences, Arts, and Letters.

the same category I would also place some of the most promi¬

nent Austrian writers: Ludwig Anzengruber, Peter Rosegger,

Paula Groger, etc.

(2) The great bulk of dialect literature belongs to the second

group which pretends to represent real dialects but actually is

an ennobled form of dialect and even a new kind of literary

language based on local dialects but frequently abandoning the

most striking characteristics of the own dialect and using in¬

stead a synonymous form or expression from a neighboring dia¬

lect. For an illustration of this statement may I refer to just

one author, namely Alfred Huggenberger, one of the prominent

Swiss writers of today. He lives in the eastern part of Switzer¬

land, in the canton Thurgau, and the grammatical forms in his

popular comedies and farcical plays are essentially those of his

dialect. But there are also elements, which for some reason of

predilection were taken from other dialects. This is true es¬

pecially for the expression mira meaning “meinetwegen” ; e.g.

V erheimleched! s mira vor de Lute, ihr zwoo, wenn I sab recht

tunkt “Verheimlicht es meinetwegen vor den Leuten, ihr zwei,

wenn euch das recht diinkt”. This sentence is taken from Hug-

genberger’s most important poular comedy Dem Bollme si bos

Wuche 4 (a Swiss Malade imaginaire) p. 61. The word mira is

characteristic of the Bernese dialect. This type of dialect

literature is developing a great multitude of new literary dia¬

lects which differ from the spoken dialects in the same way as

standard High German differs from colloquial High German.

As I have just shown, this is to a certain extent true even of the

Swiss dialect literature of which we possess an excellent survey

written by Otto von Greyerz, the competent representative of

Swiss literature at the University of Berne.4 5

(3) A third group comprises that literary production in dia¬

lect form which is a faithful copy of the spoken dialect. The

student of dialects is particularly interested in this type of liter¬

ature, and because he does not trust the professional writers of

dialect too much, he is often compelled to write literature him¬

self by collecting texts. He accomplishes that work by writing

down fairy tales or stories directly from the lips of some imag-

4 Verlag Huber und Co., Frauenfeld, 1914.

5 Otto von Greyerz, Die Mundartdichtung der deutschen Schweiz. Geschichtlich dargestellt.

H. Haessel Verlag, Leipzig, 1924; cf. also the article “Schweizerische Dichtung” in Merker and

Stammler’s Reallexikon der deutschen Literaturgeschichte III 213-233 written by the same author.

Senn — Literary Language and German Dialect . 845

inative country-man or country-woman who is supposed to

speak the dialect correctly. Of course, this type of dialect liter¬

ature is only used for the purposes of scientific investigation.

The question may arise : why did the dialect literature grow

up so rapidly just at the time when the unification of the stand¬

ard literary language had been accomplished ? We have already

partly answered this question above when we saw that the real¬

istic authors refused to use the sublime forms of standard High

German in the speech of personages not belonging to the culti¬

vated class. Thus dialect expressions were used as a means of

characterization. In other cases a certain local patriotism was

the motive force for the use of the dialect. It is at the same

time interesting and important to know that not even one of the

German dialect authors wrote dialect because he could not write

otherwise. All of them are people of high cultural standard,

thoroughly educated, and most of them wrote also in standard

High German form at the same time. As examples it may suffice

to mention the names of three, namely Johann Peter Hebei, Al¬

fred Huggenberger, and Otto von Greyerz.

In my opinion the best justification for the existence of a

dialect literature has been given by Behaghel in the introduction

to his edition of Johann Peter Hebei's works.6 p. XV - XVI.

Behaghel states justly that only those things or happenings for

which the language already possesses adequate means of ex¬

pression can be easily described. But only those words which

are used again and again are active parts of the language.

Therefore a language is capable to express with ease only those

things which have already often been expressed in that langu¬

age. This statement is corroborated by the fact that all those

languages which as a result of the world war obtained the role

of official languages (e.g. Lithuanian and Lettish) where ob¬

liged to create first the necessary terminology before they could

be successfully used in the state administration, in the courts, in

the schools, etc. The range of possible expressions in a langu¬

age varies from period to period and from nation to nation.

Generally this range of expression must be as large as the field

of interest. In a standard literary language, i.e. a language of

the higher social stratum, the range of interest diminishes in

6 O. Behaghel, Hebels Werke. Erster Teil. Allemannische Gedichte. Deutsche National-

Litteratur. 142. Band, Erste Abteilung.

346 Wisconsin Academy of Sciences , Arts, and Letters.

the same proportion as a thing approaches the simple life of the

lower classes. This was especially the case in the superidealis-

tic German literature of the 17th and 18th centuries. Therefore,

still today, there are a great number of objects, especially num¬

erous plants and animals, meals, household-effects, etc., for

which the standard literary language lacks names generally

recognized and understood.7 I have some experience of my own

in this field from my work on the Lithuanian-German Diction¬

ary, where it became necessary to add to the German transla¬

tion the corresponding Latin names in order to avoid misunder¬

standings. For the same reason the literature proper is unable

to grasp and to describe the petty everyday manifestations of

emotion that appear as expressions of anger and irritation or in

the form of abusive terms and imprecations or as interjections

of pain and joy or as innumerable fond and tender words. The

standard literary language stands aloof from the speech of the

common people. And as a result of this aloofness we find a

certain poverty in the speech of the educated class. Therefore,

Behaghel regards the creation of the dialect literature as an

enlargement of poetry in general.

7 Cf. Paul Kretschmer, Wortgeographie der hochdeutschen Unmgangsprache.

Gottingen 1918.

THE WINNEBAGO VISIT TO WASHINGTON IN 1828

Louise Phelps Kellogg

Governor Lewis Cass of Michigan Territory was a man of

ideas and resource. Moreover he was profoundly interested in

the Indian race and sympathetic with the troubles caused by

encroaching frontiersmen. In 1827 had occurred the so-called

Winnebago war, which might have proved disastrous for the

Northwest frontier had not Cass by rapid movements and wise

councils prevented serious bloodshed. The occasion of the out¬

break was a false rumor that two Winnebago prisoners had

been done to death by the garrison at Fort Snelling on the upper

Mississippi. The real cause was the invasion of Winnebago

lands between the Mississippi and Rock rivers by lead miners,

who overran their territory, drove off the game, and made free

with all the Indians considered their own. An even deeper cause

was the hatred of the tribe for Americans, to whose occupation

of the region they had never become reconciled.

The Winnebago were an eastern offshoot of the Siouan race,

which in the dawn of history occupied most of the southern por¬

tion of what we now call Wisconsin. During the wars of the

French regime, they became weakened in numbers, but not less

proud, intractable, and aloof in spirit. They were among the

most active associates of Tecumseh and the Shawnee Prophet

in their rebellion against the might of the United States. They

fiercely resented the coming of American troops in 1816 and

were subdued only by fear of the cannon that were placed upon

the forts, and the bayonets of the soldiers that mounted guard.

When Fort Howard was built at one end of the Fox-Wisconsin

waterway and Fort Crawford at the other, they withdrew to

their villages on the upper Rock and upper Fox and in sullen

silence watched suspiciously the movements of the white men.

After Red Bird, their guilty chief, had surrendered in 1827

at the Portage, and Governor Cass and General Atkinson had

put down the incipient rebellion, Cass, in the summer of 1828,

called the Winnebago chiefs to a council at Green Bay. Before

the council met he wrote to the commissioner of Indian Affairs

recommending that a party of their chiefs “be permitted to

347

348 Wisconsin Academy of Sciences, Arts, and Letters.

visit our cities to impress them with our power; it may tend to

quiet their restlessness and tame their ferocity.” The Commis¬

sioner favored the plan as a manner of conquering more merci¬

ful than with cannon and bayonets. The War Department ap¬

proved, and gave orders that fifteen chiefs be invited to visit

Washington and that John H. Kinzie accompany them.1

Meanwhile their beloved chief Red B£rd had died in prison

at Prairie du Chien ; the government had ordered a fort built at

the Portage— in the heart of Winnebago land- — and Cass and

Pierre Menard of Illinois had met the chiefs in council at Green

Bay. At that conference Cass invited the chiefs to pay a visit

to their Great Father, the President of the United States.2

There was much discussion at the council concerning the per¬

sonnel of the delegation. The two principal families among the

tribe were the Decorah and the Caramaunee, rivals for the

headship. The leaders of the Washington delegation were the

Caramaunee. Their village was first on Green Lake of Fox

River, but after this visit to Washington and the treaty of 1829,

the Caramaunee removed to the Baraboo River. The elder Cara¬

maunee, known as Naw-kaw or Wood was at this time the prin¬

cipal chief of the nation. He had been Tecumseh’s adviser and

was present at his death at the Brattle of the Thames. Mrs.

Kinzie describes him in 1830 as “a stalwart Indian, with a

broad, pleasant countenance, the great peculiarity of which was

an immense under lip, hanging nearly to his chin.3

Caramaunee’s kinsman, Hoo-wau-nee-kah or Little Elk was

the orator of the group. When Henry Clay visited the Winne¬

bago in Washington “after carefully looking at the countenances

and bearing of all the members of the deputation, [he] had in¬

dicated him [Little Elk] as the one possessing the greatest tal¬

ent; and he was greatly pleased when informed that he was

the principal orator of the nation, and decidedly superior in

abilities to any other individual of the tribe.”4

The Decorah family had a tincture of white blood, being de¬

scendants of the famous chieftess, Glory-of-the-Morning and

her French consort, Sabrevoir de Caris. The head chief of this

family was the White War Eagle or Konokah Decorah, grand-

1 Indian Office Files, Jan. 24, 1828.

3 Ibid. Proceedings of Council, Aug. 17-20, 1828.

3 Juliette A. Kinzie, Wau-Bun, (edition of 1930), 63.

* Ibid., 65.

Kellogg — Winnebago Visit to Washington 349

son of the chieftess. Mrs. Kinzie says of him, he was “the most

noble, dignified, and venerable of his own or any other tribe.

His fine Roman countenance, rendered still more striking by his

bald head, with one solitary tuft of long silvery hair neatly tied

and falling back on his shoulders ; his perfectly neat, appropri¬

ate dress, almost without ornament, and his courteous de¬

meanor, never laid aside under any circumstances, all combined

to give him the highest place in the consideration of all who

knew him.”5 This fine old chief did not go with the party on

this journey as his age forbade. His son, Spoon Decorah, was

a somewhat inconspicuous member of the deputation.

The more prominent Decorah that took the journey was a

nephew of the old chief, whose village was on the upper Missis¬

sippi near La Crosse. His Indian name was W au-kon-haw-ka w,

or Snake Skin, and he usually wore the skin of a rattlesnake

woven about his head as a turban. After this journey he was

universally known as Washington Decorah. He was a large,

handsome man six feet three inches tall and quite an orator,

spokesman for his tribe at the Treaty of 1829 at Prairie du

Chien. At that treaty he said: “Fifteen of us went last year

to see our Great Father; v/hen we shook hands with our Great

Father we did not think him a man like ourselves ; we thought

him the Great Spirit — his house was so grand and everything

around him so splendid ; but when we heard him speak we found

him a man.”6

The chief who guarded the entrance to Lake Winnebago,

with a village on what was later Doty’s Island, was Hootschope

or Four Legs (really Four who Stand). He was a wise old

chief whose wife was a Fox woman. She had been in her youth

as far as New York, and knowing the power of the whites she

had during the Winnebago War pled successfully with her band

not to attack the whites.7 This family was represented by the

nephew of the old chief, whom the whites called Dandy, because

of his fondness for dress.8 He was also called the Little Soldier

on the trip, because he bore the hardships well.

The Rock River band was represented by Kaw-ray-kaw-saw,

or White Crow, whose village was on Lake Koshkonong. White

5 Ibid., 63.

6 Indian Office Files, Proceedings of the Treaty, Aug. 1829.

7 I. 0. F. Jan. 24, 1828.

8 Mrs. Kinzie, Wau-Bun, 6S-66.

350 Wisconsin Academy of Sciences , Arts, and Letters .

Crow was accompanied by his young daughter, eighteen years

old and said to be beautiful. She married, soon after her return,

another member of the party, Yellow Thunder. She was always

called the Washington Woman by her people and Mrs. Kinzie

describes the airs she assumed. “She had a pleasant, old-ac¬

quaintance sort of air in greeting me, as much as to say, 'You

and I have seen the world.' ”9

Other members of the delegation were “Talk English, a re¬

markably handsome, powerful young Indian," who received his

name on the trip, because of his reiteration of that expression ;10

Tshi-zun-haw-kaw, “who united the characters of a conjurer or

medicine man, with that of a brave,"* 11 had a most pleasing

countenance and was accounted the most intelligent and pro¬

gressive observer of the party. We have not the names of the

other six members of the deputation, but all the chief bands

were represented. Their conductors were Robert Forsyth, sec¬

retary of Governor Cass, and John H. Kinzie, recently appoint¬

ed Indian agent at Fort Winnebago. The interpreter was Pierre

Pauquette, the half breed Winnebago who had charge of the

transfers at the Portage and was much beloved by all the tribe.

Moreover he was the only really competent interpreter of the

Winnebago language and spoke also both French and English,

although he could neither read nor write.

The deputation of fifteen Indian chiefs, one woman, and

three conductors left Green Bay with Governor Cass, after he

had finished the council negotiations. The steamboat Henry Clay

conveyed them to Detroit where they arrived about the first of

September.12 They remained a month at Detroit, waiting Gov¬

ernor Cass's convenience and left there Oct. 6 on the Niagara

for Buffalo. At Buffalo they were conveyed by stage across

New York state while Governor Cass traveled on to Washing¬

ton by a shorter route. The Indians passed through Utica,

Schenectady, and Albany, and thence took a river steamer to

New York City where they arrived on October 19. 13

If only some of the chiefs could have kept diaries how amus¬

ing and interesting their impressions would have been. They

9 Ibid., 75-77.

™Ibid., 64-65.

11 McKenney and Hall, Indian Tribes (Phila. 1854), 1, 215.

12 Detroit Gazette, Sept. 4, 1828.

13 Expense account in Congressional Documents, Serial 186, No. 129.

Kellogg — Winnebago Visit to Washington 851

hitherto had believed themselves braver and more virtuous than

the whites and equal in number to them.14 What must have

been their amazement at the numbers of “children of the Great

Father,” whom they met on their journey. Agent Kinzie tells of

their shrewdness when passing by stagecoach through the coun¬

try. The driver connived with the tavern keepers to sound the

horn just as the Indians were seated at table for a meal.

“Do you pay for all these provisions that are set before us

at the hotels ?” one asked their conductor.

“Yes. Why do you ask.”

“Nothing. I thought you paid for just what we ate.”

At the next stopping place a fine breakfast was set upon the

table of which they partook plentifully. As the horn sounded

for the stagecoach, each chief sprang to his feet. One seized

the plate of biscuits and poured them into the corner of his

blanket; another the remains of a pair of chickens; a third emp¬

tied the sugar bowls ; each laid hold of what was nearest him,

and in a trice nothing was left upon the table but empty plates

and dishes. The landlord and waiters, meanwhile, stood laugh¬

ing and enjoying the trick.15

In New York the Winnebago created a sensation. None of

their people had ever been seen there, and their barbarous cos¬

tumes, their singular manners attracted much attention. Pro¬

prietors of theatres sent them complimentary tickets and then

advertised that the Indians would be present, and their houses

were thronged. At the Bowery theatre one of the chiefs was so

pleased with the singing of the cantatrice, that he stripped an

eagle's feather from his costume and sent it by the boxkeeper

to the “singing squaw.” October 27 they visited Peale's Mu¬

seum and enjoyed the experiments, such as the air pump. Naw-

kaw gravely thanked Mr. Peale with the remark that he could

not understand what was done, it must come from the Great

Spirit.16 The next day they went to a balloon ascension at the

Battery, and even Kinzie expected they would be impressed with

the daring of the aeronauts ; but the leading chief thought them

unwise to trifle with their lives in that way. “What good does

it do?” he asked.17 Another chief said when asked what he

14 Niles Register, xxxv, 101.

15 Mrs. Kinzie, Wau-Bun, 78.

10 National Intelligencer, Oct. 28, 1828.

17 McKenney & Hall, Indian Tribes , i, 317.

352 Wisconsin Academy of Sciences , Arts, and Letters .

thought of it, ‘Think nothing of it. Americans foolish.”18 Nor

could the chiefs be made to express their wonder at so many

people. Naw-kaw boasted when his attention was called to a

great crowd, “We have more in our smallest villages.”

New York’s enthusiastic adulation was rebuked by Washing¬

ton journalists to whom Indian visitors were less of a novelty.

Indians are intelligent men and should be so treated, wrote a

correspondent.19 Philadelphia however showed an interested

curiosity in the chiefs’ visit. Arriving there the last of October

they made an unexpected appearance at the French opera and

excited a lively sensation as they filed into the stage box, in all

their savage finery. They stayed at Mr. Wade’s hotel on North

Third Street,20 but soon passed on to Washington where they

arrived the last day of October and were domiciled at Tennison’s

Hotel.

Their stay in Washington lasted for six weeks for which the

innkeeper charged $1,700.00 including $250.00 for “damages

done the house.” For the entertainment of the citizens the Win¬

nebago staged a war dance on the common between the White

House and the river, for which a dollar’s admission was charg¬

ed. Postponed because of bad weather it finally took place the

last of November. The Marine Band played and the chiefs gave

the Discovery Dance, a mock battle, and the Rejoice Dance.

“The movements of the Indians would not offend the most rigid

and fastidious delicacy,” wrote an attendant journalist.21

One of the amusements then claiming the patronage of

Washingtonians was an exhibition at Carusi’s Assembly Rooms

of trained animals — birds and dogs. When the Winnebago chiefs

visited this entertainment they were much delighted. A canary

named “Fairy” was supposed to play dominoes and one of the

dogs was similarly accomplished. Two of the chiefs challenged

these learned animals to a contest. This created a considerable

excitement in Washington, all the newspapers carrying an ac¬

count of the “Challenge Extraordinary.” The result of the play

was not reported. One day the local militia company, known as

the Washington Guards marched in a body to Tennison’s Hotel

18 Washington Telegraph, Oct. 29, 1828.

™lbid., Oct. 26, 1828.

20 Detroit Gazette, Nov. 13, 1828.

21Washington National Journal, Dec. 1, 1828.

Kellogg — Winnebago Visit to Washington 353

and their band serenaded the chiefs, after which their orator

gave thanks in a grandiloquent speech.22

The principal purpose of the trip, however, was for the

chiefs to have an interview with the President, their Great

Father. The President at this time was John Quincy Adams,

who was more familiar with deputations from Europe than from

the Indian tribes. He received the Winnebago with considerable

pomp, surrounded by members of his cabinet and certain in¬

vited diplomats. Naw-kaw presented him with a peace pipe and

Little Elk declaimed a sonorous oration, translated by Pau-

quette, in which he assured the Great Father that the tribe

would maintain perpetual peace.23 On December 2 the Presi¬

dent received the delegation once more, and gave them a docu¬

ment as follows:

This certifies that Tshi-zhunk-kau-kaw, a Winnebago Chief,

in company with fourteen other chiefs and Warriors of that

nation, has visited the seat of Government of the United States,

and held friendly councils and smoked the pipe of peace with

me in my house. Having full confidence in the declarations

freely and solemnly made by him and his associates, of their

determination to maintain perpetual peace and friendship with

the United States, I recommend him to the favorable notice and

consideration of all my white and red children. [Signed] John

Quincy Adams, President of the United States.24

The visit, however delightful, must come to an end. Kinzie

prepared for their return journey and had the tribesmen fitted

out with clothes by a fashionable tailor, who made them fifteen

blue frock coats, as many pair of leggings to match, sixteen

blue cloth caps (perhaps one for Pauquette), cloth and ribbon

for the squaw, three dozen pair of stockings and twelve pairs of

shoes. They traveled across country by stage to Detroit, where

they arrived the last of the year. Horses, saddles, and bridles

were there bought for them, and they journeyed homeward to

their separate Wisconsin villages, with many gifts to testify to

the welcome they had received.

22 Washington Chronicle, Nov. 22, 1828; National Intelligencer, Nov. 20, 1828.

23 Naw-kaw’s portrait was painted in the act of presenting the pipe. A reproduction is in

McKenney & Hall, Indian Tribes, 1, 72. For Little Elk see Ibid., ii, 289. McKenney requested

an appropriation for the portraits of all the chiefs at $20 apiece. The originals are not now in

Washington.

24 The original of this document was presented in 1930 to the Museum of Luther College,

Decorah, Iowa. It had been preserved by a missionary at Wittenberg, Wis. to whom it had been

given by a Winnebago many years ago.

3*54 Wisconsin Academy of Sciences, Arts, and Letters.

The visit of the chiefs was justified by its results. The ex¬

pense of $10,272.05 to the government was less than a war

would have been. The Winnebago never again attempted a re¬

volt against government authority, and four years later, when

the Sauk under Black Hawk went on the warpath, the Winne¬

bago were restrained from joining them by the knowledge that

their chiefs had of the might and power of the United States.

Especially friendly to the whites were the chiefs of the delega¬

tion. White Crow rescued the captive girls from Black Hawk,

acted as guide to General Dodge and served with the whites in

the battle of Wisconsin Heights. Little Elk kept Kinzie informed

of the movements of the hostiles, and a brother of Washington

Decorah brought the defeated Black Hawk as prisoner to

Prairie du Chien. Certainly the trip to Washington of the Win¬

nebago chiefs proved a good investment for the future welfare

of Wisconsin. 1 ! !'MI

THE DETERMINATION OF ORGANIC NITROGEN:

PAST AND PRESENT*

H. A. SCHUETTE AND FREDERICK C. OPPEN

Chemistry Department University of Wisconsin

It is the purpose of this review to present a picture, suffi¬

ciently comprehensive yet not too detailed, of the development

of one of the most fruitful and important fields of organic anal¬

ysis, the determination of nitrogen, as it has evolved from its

crude inceptions down to the numerous specialized techniques

of the present day. The plethora of available material and the

fact that oxidized nitrogen occurs but rarely in organic mater¬

ials as compared to the abundance of this element in its reduced

condition, necessitates a sharp curtailment of the field. The

bulk of the emphasis, therefore, has been placed upon the latter

problem.

By way of background for what shall be a century of prog¬

ress in this field, brief reference to the contributions of the old

masters of analytical chemistry to the beginnings of ultimate

organic analysis may not be out of order. Taking as a point of

departure the pioneering investigations of Lavoisier of about

1784 ( 1 ) on the determination of the elementary composition

of organic substances containing only carbon, hydrogen and

oxygen — not accurate, it is true, by present-day standards yet

none the less most significant because he exploited a method of

analysis which was soon to be improved by others— we come to

the contributions of Berthollet (2) with respect to nitrogen. In

a sense, he applied to this determination the principle of dry dis¬

tillation. The virtue of his method seemed to lie chiefly in the

fact that a large sample was required, a fortuitous situation

since the experimental error was large, doubtless no greater

than was considered permissible in those days.

Following a brief desultory period of experimentation with

gaseous oxygen under atmospheric pressure, attention turned to

the use of solid oxidizing agents. Sometime near the beginning

of the nineteenth century, Abildgaard had suggested (3) and

* Presented as part of the symposium on a Century of Progress in Agricultural and Food

Chemistry before the Division of Agricultural and Food Chemistry of the American Chemical

Society at its 86th meeting, Chicago, Sept. 11 to IS, 1934.

855

356 Wisconsin Academy of Sciences, Arts, and Letters .

later Berzelius (4) used this type of reagent; the one manga¬

nese dioxide, the other lead peroxide. The first satisfactory

values were obtained several years later by Gay-Lussac and

Thenard ( 5 ) who introduced the sample and oxidizing agent,

potassium chlorate, in pellet form into the combustion tube

fixed in a vertical position. The gaseous reaction products were

then analyzed. The presence of nitrogen in the sample made

necessary close control of the quality of chlorate added in order

to prevent its oxidation to the nitrate form. Although the re¬

sults which these investigaors obtained were better than others

before them had been able to get, the accuracy of their procedure

was still far from satisfactory.

It was at this time that Berzelius introduced the horizontal

combustion tube with movable burners, thus obviating the diffi¬

culty of having particles of substance spatter into a cold por¬

tion of the tube and so escape decomposition. Others subse¬

quently clung to the vertical tube ; nevertheless the idea proved

superior and has remained in use to the present.

A suitable oxidizing agent was the next goal for it was ap¬

parent that herein lay one of the limiting factors in the success¬

ful determination of the ultimate composition of a compound.

Various metallic oxides and oxy-salts had been used up to this

time, though rather unsuccessfully except in the case of potas¬

sium chlorate. By 1815 this search appears to have ended for

then Gay-Lussac ( 6 ) introduced the now familiar use of cupric

oxide as oxidizing agent, together with copper turnings for the

decomposition of oxides of nitrogen. Dobereiner (7) announced

the same discovery almost simultaneously and apparently inde¬

pendently.

The Dumas Method

We now pass over a series of methods and modifications to

the work of a great coordinating genius, Justus von Liebig, and

his brilliant contemporary, Jean Baptiste Andre Dumas. It be¬

came increasingly and painfully evident that no simple, accurate

method for organic elementary analysis was available, and that

this lack greatly crippled investigators in the field, for they had

no ultimate foundation upon which to rest their work. To Lie¬

big, in particular, already on the threshold of fundamental dis¬

coveries in organic chemistry, this exigency became so great

Schuette & Oppen — Determination of Organic Nitrogen 357

that he devoted a number of years to work on the problem, and

was rewarded by a method which, though burdened with cer¬

tain inescapable errors, nevertheless was reliable and accurate.

His apparatus was characterized by extreme simplicity and was

little more than a combination of parts well known at the time.

In fact, Liebig said of it (&), “This apparatus involves nothing

new, save its simplicity and the complete dependability which

it affords.” The first thing which he did was definitely to sepa¬

rate the nitrogen determination from those of carbon, hydrogen

and oxygen. Almost like the search for the fabled philosopher's

stone had been the quest after a universal method of embracing

all organic compounds as well as all elements therein, and Liebig

was the first to realize that, since each element has its own pecu¬

liarities, much more is gained by attacking them individually

rather than collectively. It is true that methods of simultaneous

analysis for several elements have been proposed from time to

time, but even though very good results were claimed they have

enjoyed only passing comment (9, 10). At this time, also, Liebig

introduced his famous “Kaliapparat,” for the purpose of ab¬

sorbing the carbon dioxide evolved in the combustion of the

sample. It was small enough to be weighed conveniently upon

the analytical balance and is apparently the precursor of all

later models.

Once the way was shown to a reliable method capable of re¬

producible results, and the limitations of this method clearly

recognized, modifications and improvements were not slow of

appearance. Of the mushroom crop which sprang forth, one in

particular was accepted less reluctantly than the original; in

fact, even the name by which the procedure was eventually

known was that of modifier rather than originator. We refer

to the contribution of Dumas (11).

Dumas' improvements were twofold: the substitution of an

inert but readily absorbable gas for air in the combustion tube

and the use of potassium hydroxide solution, rather than mer¬

cury, as the confining liquid. The first modification eliminated

an error inherent in Liebig's method but introduced not only

the necessity of working with air-free carbon dioxide but also

of expelling air from all parts of the apparatus before begin¬

ning a combustion. The second modification seems to have with¬

stood best the test of time for it has suffered less by way of

change than perhaps any other feature of the procedure.

358 Wisconsin Academy of Sciences , Arts, and Letters.

Dumas produced carbon dioxide by heating lead carbonate in

an extended portion of the combustion tube, a practice which

was superseded later by the introduction of the separate gas

generator (12). The Kipp apparatus has been found especially

suited for this task if precautions are taken to produce air-free

carbon dioxide, such as the procedure of Bernthsen (IS) where¬

by one evacuates the water-covered marble chips, or that of

Kreusler (14) who recommended the use of fused sodium car¬

bonate and sulfuric acid. Numerous other procedures also are

available (15, 16). More recently Pregl (17) recommended a

preliminary etching of the marble chips with hydrochloric acid,

followed by a careful rinsing with water.

As much effort has been spent upon the preparation of suit¬

able reduced copper for the purpose of decomposing the oxides

or nitrogen as securing pure carbon dioxide. Dumas reduced

his copper with hydrogen, an unsatisfactory practice as subse¬

quently became evident when it was observed (18) that copper

thus prepared emitted, upon strong heating, a gas not absorbed

by the potassium hydroxide solution. An explanation for this

circumstance was eventually found in the fact that a copper hy¬

dride is formed during reduction (19) under these conditions.

Further investigation (20, 21) showed that this difficulty could

be corrected by igniting the copper strongly and then cooling it,

all in a current of pure carbon dioxide. An entirely different

approach to a solution of this problem is seen in the suggestion

that a silver spiral be substituted for the copper because of the

claim that it need be neither reduced nor dried, is more efficient

than the latter in decomposing oxides of nitrogen, and serves

to hold back halogens, if present (22, 23) .

Finally, the technique of measuring the nitrogen evolved in

the combustion, like all the other features of this determination,

has undergone many changes, although the attainment of that

development has not, apparently, been so difficult. The history

of the convenient azotometer of Schiff (24) traces its course

from the inverted bell jars of Lavoisier and of Gay-Lussac and

Thenard, through the graduated cylinders of Liebig and Dumas,

respectively. The whole picture of the development of ultimate

organic analysis reflects a fine example of the scientific evolution

of a group of procedures brought about by careful, tireless

workers, over a period of many decades, by means of which the

Schuette & Oppen — Determination of Organic Nitrogen 359

crude seed, sown with prophetic vision, brings forth an hun¬

dredfold of perfect fruit.

The Kjeldahl Method

In a discussion of the wet oxidation method for the determi¬

nation of nitrogen one again finds material for ample develop¬

ment of the central theme of this review, progress and evolution,

for the rise of this procedure embraces fully three-quarters of

a century before, as well as the half-century after, the publica¬

tion of Kjeldahl’ s original communication. Originally developed

to meet the needs of the chemist in the brewing industry, yet it

was not to be exclusively his tool for very soon it found entree

into other fields of technical, as well as scientific, analysis.

Essentially, the Kjeldahl method consists in decomposing

the material under examination with strong sulfuric acid in the

presence of oxidizing agents, accelerators or catalysts, used

either singly or in combination. Following complete decomposi¬

tion, the digestion mixture is made strongly alkaline, where¬

upon the ammonia is distilled off and subsequently determined

by some suitable means.

The technique of effecting decomposition of an organic sub¬

stance with a sulfuric acid-catalyst mixture as outlined above,

appears to have been first employed by Berzelius (4) when he in¬

vestigated the decomposition products from various animal and

vegetable substances after digestion with a sulfuric acid-lead

peroxide mixture. He, however, concerned himself only with

the products of distillation from acid solution.

Sulfuric acid-digestion, followed by precipitation of the re¬

sulting ammonia as the chloroplatinate, first enters the picture

in 1845 as a method for the determination of urea (25, 26) . In¬

asmuch as it was applied to the examination of urine and em¬

ployed no catalyst, this work of Heintz and Ragsky presages,

in a sense, the important modern researches of Folin and co¬

workers. And in this connection it may be timely, too, to recall

that Grete in 1878 (27) offered the suggestion that the decom¬

position of hair, wool, leather and similar substances by the now

obsolete Will-Varrentrup soda-lime method (28) could be expe¬

dited by giving the material under examination a preliminary

treatment with sulfuric acid.

360 Wisconsin Academy of Sciences , Arts , and Letters .

It may also be of interest to make mention of the fact that

Kjeidahl had apparently received the inspiration to try an acid

medium from the researches of Wanklyn and associates (29)

on the use of strongly alkaline permanganate solutions for the

liberation of so-called “albuminoid ammonia”. Reasoning that

the tendency to split off ammonia would be much greater in acid

than in alkaline medium, he tried first dilute and finally concen¬

trated sulfuric acid and was rewarded in the second instance by

a yield of ammonia which compared favorably with that ob¬

tained by the soda-lime method (28). For a series of pure com¬

pounds the results were found to agree well with the calculated

values, a fact which gave the new procedure further weight. Be¬

cause of its extreme simplicity and rapidity, together with the

high degree of accuracy obtainable in the analysis of protein and

related materials, Kjeldahl’s method was not long in receiving

recognition. Indeed, the germ of an idea suggested by the orig¬

inal communication exactly fifty years ago has brought forth

reports of an unprecedented number of investigations, exten¬

sions, and modifications. Obviously, the only rational method

of reviewing them is by classification, selection of typical ex¬

amples, and drawing of summarized conclusions. To this end,

the subject will be considered with as much brevity as is con¬

sistent with clarity of presentation under the topic headings of

Digestion, Distillation of Ammonia, Determination of Ammonia,

and, finally, Scope of Application of the Method.

Digestion. — A Kjeidahl digestion mixture consists, in gen¬

eral, of representatives of two classes of substances: (1) strong

mineral acids, and (2) catalysts or accelerators. There may also

be added a third kind of substance, inert in itself, which dis¬

solves in the acid and serves to raise the boiling point of the

mixture, thus increasing the reaction velocity. Potassium sul¬

fate, first employed by Gunning (SI), or the less expensive sodi¬

um salt (S2), is the most familiar example of this group. Its

indiscriminate use, however, may lead, it is said (33, 34), to

losses of ammonia during digestion when the composition of the

residue approximates that of the acid salt. Kjeidahl himself sug¬

gested phosphorus pentoxide, an ingredient which has since

been used by others either as such (35-37) or in the form of the

acid as a substitute, wholly or in part (38-41), for the sulfuric

Schuette & Oppen — Determination of Organic Nitrogen 361

acid. The desirability of using phosphoric acid has not, how¬

ever, gone unchallenged (42) .

Rapid digestion methods involving phosphoric-sulfuric acid

mixtures containing hydrogen peroxide as oxidizing agent have

been investigated by Wieninger and Lindemann (43) who found

that these methods, although quite satisfactory, nevertheless

yield results appreciably lower than those obtained by the older,

slower Kjeldahl method. They offer the further objections that

these new methods are very hard on glassware and require com¬

paratively expensive reagents, so that the great saving in time

of digestion (10 to 20 minutes as against 55 to 60 minutes) is

still of problematic value at present. These discrepancies may

well be due to the extremely drastic treatment, rather than spe-

fically to the presence of phosphoric acid. Finally, a proposal

which suggests a page from the days when Carius fusions were

popular merits mention. It is to the effect that the Kjeldahl

digestion of substances not readily attacked be carried out at

330° C. with fuming sulfuric acid in a sealed tube (44). Al¬

though this interesting suggestion is probably worthy of de¬

velopment, yet the experimental technique and hazards involved

would preclude its use except under unusual circumstances.

Passing now to a discussion of catalysts, accelerators, and

oxidizing agents used to increase the reaction velocity we find

than an unusually large number of substances has been tried for

this purpose. The accompanying table lists there, together with

significant references and remarks on their efficacy.

Table I

Selected List of Catalysts and

Accelerators Available for Use in a Kjeldahl Digestion

Compound

KMnO*

Author

Kjeldahl (SO)

Remarks

Added as a final step to insure

oxidation.

Salkowski (45)

Ordinarily unnecessary, and

objectionable in the presence

of substances of high halogen

content.

Phelps (46)

May cause loss of nitrogen.

Dowell and Friede- Use discontinued,

mann (47)

362 Wisconsin Academy of Sciences, Arts, and Letters.

Schuette & Oppen — Determination of Organic Nitrogen 363

364 Wisconsin Academy of Sciences , Arts , and Letters .

Schuette & Oppen— Determination of Organic Nitrogen 365

That the liberation of the resulting ammonia may be quanti¬

tative, it becomes necessary in some cases to destroy heavy met¬

al-ammonium complexes formed during digestion. This is par¬

ticularly true when mercury compounds have been used in this

operation. Potassium sulfide, the precipitant most used today,

was introduced by Wilfarth (51). Following this came sodium

thiosulfate (98), now recommended in the current edition of

“Methods of Analysis” of the Association of Official Agricul¬

tural Chemists as an alternate to sodium sulfide, then mono¬

sodium phosphate (99), potassium xanthogenate (100), and

finally potassium arsenate (101), because its sponsor had found

reason to criticize the action of sodium phosphate. Be that as

it may, the first two of this list have been found to be satisfac¬

tory, provided a slight excess of the sulfide or a generous excess

of the thiosulfate is used (102).

Distillation of Ammonia.— Distillation and absorption prob¬

lems have received their measure of attention, too, the attack

upon them being more from the mechanical rather than the

chemical standpoint. Space does not permit a description of

the many spray traps that have been devised to correct the ten¬

dency, because of violent ebullition or foaming, of alkali to be

carried over into the condensing system, nor yet a discussion of

the evolution of the distillation apparatus from that in vogue in

KjeldahPs time to the highly efficient, multiple-unit commercial

type of the present day. Suffice it to state, however, that these

difficulties may be practically eliminated by the use of suitable

block tin condensers and distilling traps, and such remedial

measures for bumping and foaming as zinc, or graphite (102),

or even, as is still the practice of some, to follow the Biblical

admonition of pouring oil— in this case paraffin— upon troubled

waters (175).

It was not long after the appearance of the Kjeldahl method

that the suggestion was made that steam distillation (108) be

used for the recovery of the ammonia. This practice has met

with a certain measure of success (10A-106) although some in¬

vestigators (AO) have reported that it causes small amounts of

alkali to be entrained and carried over by the spray. In micro¬

analysis, especially, has this technique been widely used (17, AO,

107-111).

366 Wisconsin Academy of Sciences, Arts, and Letters.

Aeration methods have found not a little favor among anal¬

ysts, particularly in physiological and biological fields (112,

113). A comparison of results ( 10U) obtained on the one hand

by aeration and on the other by distillation has shown, in sev¬

eral instances, that at least as much as one per cent of the ni¬

trogen obtainable by the latter mode of recovery was not car¬

ried over by the former. It was found, however, that steam

distillation served to bring these low results up to the values

obtained by heat distillation, and that a large excess of strong

alkali materially aided in increasing the yields by aeration.

Determination of Ammonia.— There now remains the analy¬

tical determination of the ammonia. Here again many proced¬

ures are available and all of them have found a measure, but by

no means the same measure, of success. Of antiquarian interest

only is the extravagant precipitation as the double platinum

salt (26). If not wholly obsolete, the iodometric determination

of excess acid proposed by Kjeldahl has enjoyed a very limited

use not only because of its cost, but also because of the extreme

simplicity of the competitive acidimetric procedure which came

into use soon after Kjeldahl’s original communication appeared.

References to indicators pertinent to this titration are indeed

numerous. A discussion of them has been omitted in order to

make room for less common means of attaining the same end.

A variant of the acid-base procedure, proposed by Sors (111>),

consists in carefully neutralizing the digestion mixture, adding

a measured excess of standard alkali solution, boiling to expel

ammonia, and determining by titration the decrease in alkalin¬

ity of the solution because of this evolved gas. Satisfactory re¬

sults are claimed, distillation is avoided, but still the method

has not been much used. The thought suggests itself, although

it has not been verified experimentally, that the presence of such

large quantities of salts in solution would make for a sluggish

end-point.

A happy union of two old neighbors, so to speak, was

brought about when Nessler’s reagent was applied to Kjeldahl

distillates, for the history of this reagent can be traced back

nearly as far as that of the Kjeldahl digestion. As early as 1839

the fact was announced (115) that a dark brown precipitate is

formed when mercuric iodide is treated with ammonium hy¬

droxide, but no analytical use was made of this reaction until

Schuette & Oppen — Determination of Organic Nitrogen 367

Nessler described it (116) as an invaluable one for character¬

izing ammonia. Following this, it was widely accepted as a deli¬

cate qualitative and quantitative reagent for extremely small

amounts of ammonia (117, 118). Eventually it was applied to

biological materials (112), and to micro-analysis in general

(W, 119).

The reaction between formaldehyde and ammonium salts,

studied extensively by Schiff (120), has given rise to still an¬

other important method for the determination of ammonia. If

we treat a neutral ammonium salt with an excess of strong, neu¬

tralized formaldehyde solution, the products of reaction are the

free acid and, probably, hexamethylenetetramine. Since one

equivalent of acid combines with one of ammonia, the titration

of this liberated acid is equivalent to the amount of ammonia in

the original salt. This fact was first utilized by Ronchese (121),

and was subsequently employed by others for the determination

of ammonia in urine (122), as well as quarternary ammonium

bases and ammoniacal solutions (123). Its first application to

the analysis of a Kjeldahl distillate was apparently in 1910. Its

use has spread widely since then (124-129).

Closely related to the above work is that of Sorensen (130).

the so-called formol titration of amino acids. This method de¬

pends upon the fact that amino acids in general are neutral sub¬

stances because of a mutual compensation of the amino and

carboxyl groups. Upon treatment with formaldehyde, however,

the amino group is changed to a neutral methylene imino group

as shown below, causing the acidity of the carboxyl group to

become available by titration.

RCHNHUOOH + HCHO = RCHN:CH2.C00H + IFO

Since the reaction is reversible, an excess of formaldehyde is

employed to drive imine formation far towards completion.

Very wide application has been given this method, as for ex¬

ample in urine analysis (131), in studies on the proteolysis of

brewery products (132), in nitrogen metabolism investigations

(133), in determining the nitrogen content of bacteriological

media, and as an index of the falsification of honey (134-136).

The formol titration method is rapid and convenient, the

distillation involved in most determinations of basic nitrogen is

eliminated, yet it is not a precision method. Difficulties (136)

368 Wisconsin Academy of Sciences , Arts , and Letters.

hinging around concentration of solutions, concentration of in¬

dicators, and the small but unavoidable excess of alkali in the

neutralized formaldehyde make duplication of results difficult.

Very recently Levy (137) in a physical-chemical study of the

equilibria involved has observed that two types of reaction, in

which either one or two mols of formaldehyde per mol of amino

acid are involved, may occur. Then, too, the fact that certain

amino acids have a basic reaction is mentioned also as an addi¬

tional complication. In short, for almost every variation in ex¬

perimental conditions or nature of amino acids, there is a

change in pH of the end point. Moreover, each acid in a mix¬

ture-— they nearly always occur in mixtures — exhibits a differ¬

ent end-point when titrated under these conditions. The formol

titration is thus the concurrence of many separate and complex

phenomena and can hardly be regarded as a single definite op¬

eration.

As we have seen, the Kjeldahl method attained early popu¬

larity as a commercial analytical procedure because of its sim¬

plicity and adaptability to routine work. That it should become

the subject of constant investigation with a view to further sim¬

plification, in the hope of effecting even small economies of time

or materials, could hardly have been otherwise. In particular,

the need for two standard solutions as in the ordinary acidi-

metric procedure has been circumvented in a number of ways.

One of these, the formol titration, has already been discussed.

Another, proposed by Neumann (138), is based on the observa¬

tion (139) that, given a properly cooled water condenser, and a

sufficiently large volume of liquid in the receiver, no ammonia

is lost, even though an insufficient quantity of standard acid be

present. All that is necessary is to measure out a slightly in¬

sufficient amount of standard acid solution, perform the distilla¬

tion, and finish the titration in the presence of a suitable indi¬

cator. It seems a bit strange that no references pertinent to

this method, other than the two cited, appear in the literature.

In contradistinction to this lukewarm reception stands that

given the boric acid absorption method of Winkler (140). Its

principle may best be given in the author's own words, “Boric

acid is indeed such a weak acid, that its solution does not notice¬

ably cause color changes of certain indicators. Ammonia is,

however, completely fixed by it, provided that a suitable excess

Schuette & Oppen — Determination of Organic Nitrogen 369

of the acid be present.” Distillation is carried to completion,

whereupon the ammonia is titrated directly in the presence of

Congo red (141), bromphenol blue (142) or methyl red indi¬

cators (148). The method has been successfully applied to a

large variety of subjects.

The iodometric titration of the excess of standard acid takes

place according to the following equations :

5 KI + KIOs + 3 ILSCb = 61 + 3 RSO* + 3 ILO

h + 2 NaaSaOa = Na.S.0. + 2 Nal

This reaction, described in most of the older books on analytical

chemistry, is not much used today for the reason that it is ex¬

tremely sensitive towards carbon dioxide (144) .

Less than three decades ago there appeared a second iodo¬

metric procedure (145, 14 8) for the determination of ammonia

which has received a fair measure of recent recognition. It de¬

pends upon the fact that ammonia is oxidized quantitatively to

nitrogen by an alkaline solution of hypobromite according to the

following reaction :

2 NHa + 3 NaBrO = 3 NaBr + N* + 3 H*0

A measured excess of the latter is added, the unused portion

being subsequently determined iodometrically. Willard and

Cake (147) were the first to apply this procedure to the Kjel-

dahl determination. A number of others have done likewise

since (148-151). In a sense, oxidation of ammonia by hybo-

bromite is a reaction with a dual role, for, besides carrying out

the iodometric procedure as described, the nitrogen which is

evolved may be determined gasometrically. This was done some

time ago, in principle at least, on urea (152). The reaction has

been widely utilized since (101, 151-154).

Scope of Application of the Method— The great bulk of those

natural products, cereals, meat, milk, glue, tankage, or any of

their manufactured derivatives, in which the nitrogen content

is of vital significance, have a common fundamental character¬

istic, to wit : nitrogen is present there in the basic condition for

the determination of which the Kjeldahl method is universally

applicable.

370 Wisconsin Academy of Sciences, Arts, and Letters.

But there exists a second large class of substances, promi¬

nent among which are nitro-, nitroso-, and azo-bodies, hydra¬

zine, quinoline and pyridine derivatives, and inorganic nitrites

and nitrates which are characterized by a very refractory be¬

havior toward a Kjeldahl decomposition. To the numerous at¬

tempts made to subjugate these and so to universalize the meth¬

od, some attention must be paid. It has been pointed out that

the Kjeldahl treatment involves strenuous oxidation. It is quite

logical to expect that little could be accomplished with sub¬

stances unsatisfactorily decomposed merely by increasing the

severity of treatment. A search of the literature reveals, indeed,

that no such attempts were successful and that whatever has

been done by way of extending the method has been accom¬

plished by means of a preliminary reduction before oxidation.

At first rather mild reagents were used, for von Asboth

(155) claimed to have obtained satisfactory results in the analy¬

sis of nitro- and cyano- compounds by adding to the mixture, to

provide the necessary reducing environment, either sucrose or

benzoic acid. Arnold, however, found that the claims for this

mode of procedure were overstated (156).

In the same year Jodlbauer (157) introduced an important

fundamental principle which is still in use today for the de¬

termination of nitrates by the Kjeldahl method, viz: the addi¬

tion of easily nitrated phenols to the sulfuric acid which fixes

the nitrogen in a form lending itself to ready and complete

reduction. The value of this modification is especially significant

in view of the wide occurrence and application of nitrates in

fertilizers, etc. The reducing agent used here consisted of sev¬

eral grams of zinc dust with a small amount of platinum chlor¬

ide solution as catalyst.

Several years later Forster (97) employed a phenol-sulfuric

acid mixture of five to six per cent phenol content, with sodium

thiosulfate. Following this, after a lapse of twenty-seven years,

came the use of salicylic acid (158) in sulfuric acid solution as

the nitrate-fixing agent. This reagent now enjoys an “official”

status. A suggestion (159) that resorcinol or phloroglucinol be

used instead of phenol itself has apparently received little no¬

tice.

Zinc in acid solution is by far the most widely used reductant

for the resulting nitro-group. In addition, zinc and iron filings

Schuette & Oppen — Determination of Organic Nitrogen 371

(160), stannous chloride in the presence of metallic tin (161),

iron powder (162), elementary sulfur alone (66), and sodium

hydrosulfite (163, 164) have found application.

Reduction methods of an entirely similar nature also serve

to bring within range of the Kjeldahl decomposition aromatic

nitro-, nitroso-, hydrazine, and azo-substances, although low re¬

sults have often been recorded in recalcitrant cases (35, 106,

158, 160, 161, 163, 165,). Very recently Friedrich (166) de¬

veloped a micro method of universal applicability, involving

concentrated hydriodic acid (d. 1.7) as reducing agent, followed

by a Kjeldahl digestion. With volatile substances, the reaction

is carried out in a sealed tube under pressure.

Although it had been established that molecular structure,

as well as position and nature of substituents, plays a significant

role in determining the susceptibility of aromatic nitrogen com¬

pounds to a Kjeldahl decomposition, whether plain or modified,

nevertheless, attempt at generalization have proved futile. This

is also true of the sugar osazones (167).

It has been stated that accurate generalizations are futile;

still, the mention of one such attempt by Fleury and Levaltier

(168) may be helpful in giving a concise, if crude, picture of

the situation as it stands. By the application of their procedure

they find that most substances are completely decomposed by

one-hour's digestion. Difficultly attacked substances, such as

creatin, skatol, isatine, quinine, piperazine, morphine, betaine,

choline, atropine, tyrosine, and pyridine, require not more than

one and one-half hours. The addition of benzoic acid makes

analysis of the following possible: sodium nitrate, benzonitrile,

the oxime of acetophenone, and piperidine. Reduction with zinc

extends the method to picric acid, m-dinitrobenzene, phenylhy-

drazine, mannose hydrazine, and glucosazine. A combination

of benzoic acid and zinc opens the way to semicarbazids and

semicarbazones. No good decomposition procedure was found

for either antipyrine or pyramidone.

With this discussion of the extension of application of the

Kjeldahl method there is concluded the historical sketch of the

evolution of this great analytical tool. There remain a few

words about micro-analytical and biological methods.

Microchemical Methods Applied to Nitrogen Analysis. It

has become a matter of common knowledge that Fritz Pregl

372 Wisconsin Academy of Sciences, Arts, and Letters.

was the originator of the new branch of analytical chemistry

called micro-analysis. His contribution to combustion analysis

was far greater than mere reduction of the scale of operations

and time involved, as many suppose, for through his painstaking

research he succeeded in a fuller refinement of the method and

a more complete approach to theoretical accuracy than had pre¬

viously been achieved. This two-fold advance has been of par¬

ticularly great benefit to workers in biological chemistry, since

the tedious necessity of collecting workable samples of vital ma¬

terials is greatly lessened thereby. Historically, it is interest¬

ing to remember that the inception of micro-methods was due to

just this difficulty encountered by Pregl in 1910. While engaged

in his physiological researches at Innsbruck he was faced with

the choice of spending many weary months purifying quantities

if material, or devising analytical methods to fit the samples

available. Inasmuch as he was a fine laboratory technician, he

chose the latter expedient, continuing work after his transfer

to Gratz, until, in 1914, he gave the first demonstration lecture

in microanalysis before the Vienna Academy. But still the meth¬

od did not suit him for it was not until 1917 that the first edi¬

tion of his now famous book, “Die quantitative organische Mi-

kroanalyse”, appeared. Even here progress did not stop, as the

book was twice revised and enlarged, the third edition being re¬

leased in 1929, only one year before his passing. Because he

realized that successful micro-analyses are possible only through

meticulous attention to details, Pregl took great pains to avoid

being misunderstood. To this end he published no articles on

the subject in the scientific journals, and made his book almost

trivial in its attention to detail, so that those who sought in¬

formation had of necessity to go to the source and delve deeply.

Not only is a small sample of advantage in biochemical in¬

vestigations, but it also offers great possibilities in another and

less explored field, namely the study of side reactions occuring

in conjunction with a given main reaction. It has been said that

side reactions are the curse and the hope of organic chemistry.

The curse, because they lower the yield of desired product and

make it difficult to obtain it pure; the hope, because suitable

modification of conditions can make the erstwhile side reaction

attain predominance, thus increasing the scope of every syn¬

thesis and beating back yet farther the frontiers of synthetic

Schuette & Oppen — Determination of Organic Nitrogen 373

organic chemistry. Obviously, a systematic, accurate method of

characterizing small quantities of materials must be the foun¬

dation stone of any such investigation.

Concerning application of the principles of the Kjeldahl pro¬

cedure to micro-analysis, much has been written, though little

can be said in a review of this nature. The general tendencies

in this application have been first, the use of all-glass apparatus

for digestion and distillation in order to minimize the errors of

transfer or those occasioned by leaching of rubber connections ;

second, the use of colorimetric methods for determining the

evolved ammonia. Since each of these phases has been ade¬

quately treated in previous pages, they will not be discussed

again.

1 : ; I. I ■ I | ;

One may well question, from the standpoint of sampling,

the advisability of applying micro-analysis to agricultural prod¬

ucts in general where fine pulverization is so often impossible.

Always a difficult problem in this field, it becomes more acute

when one considers the responsibility involved in selecting three

or four grains of wheat as representative of a carload or a tiny

fragment of tankage as a true average of the heterogeneous

substances composing that product. Furthermore, in semi-solid

materials, the difficulty of representative moisture content would

be almost insurmountable. To conclude, then, micro-analysis is

probably more valuable than macro-analysis where one is deal¬

ing with chemically pure compounds of unquestioned homogene¬

ity because of economies of sample, reagents, apparatus, and

time. But agricultural products as they ordinarily occur pre¬

sent the very antithesis of an ideal sample for this work, and,

hence, the procedure must be applied with extreme caution, if

at all.

Nitrogen Determination in Relation to Biochemistry . Be¬

cause of the vital relationship that obtains between the nitrogen

content of living cells, their products, and life itself, as exem¬

plified by the familiar nitrogen cycle it is most natural and in¬

evitable that biochemists and physiologists should have seized

upon the analysis for nitrogen as a research method of the most

profound significance and universal application. It is natural,

also, that they should have developed their own methods for this

purpose. They are accurate enough to indicate the trends in

374 Wisconsin Academy of Sciences, Arts, and Letters .

which they were interested, but more convenient than the time-

consuming precision methods referred to above. The formol

titration of Sorensen is one such method ; it has been discussed

elsewhere. Again, the contributions of Folin and co-workers,

appearing in the Journal of Biological Chemistry from about

1910 to date, have done much to further the practical estimation

of ammonia, amines, amino acids, etc., in biological materials.

These last cannot be discussed here, however, for we would lose

the ocean for the water in the attempt.

The name of D. D. Van Slyke is another one well known

among workers in this field, for to him goes the credit for de¬

vising the gasometric determination of amino nitrogen based on

the familiar reaction of primary amines with nitrous acid

RNH* + HNCb = ROH + 1FO + N*

in which a molecular equivalent of gaseous nitrogen is liberated

for every mol of amine present. This was first introduced (169)

in 1910 and has been widely used since in the study of proteins,

urea, and enzymes. Since nitrous acid also reacts with urea,

it is necessary that this material be carefully removed before

analysis (170).

Urea is a particularly important substance because it is the

chief end-product of nitrogen metabolism in the animal body.

A method for its determination proposed by Marshall (171) in

1913 depends upon the fact that it is hydrolyzed quantitatively

to ammonium carbonate by the enzyme urease. The ammonia

may then be distilled off and determined iodometrically or in

any of various other ways. Apparently the selection (172) of

an active urease preparation is a matter of some concern for not

all such extracts are of equal quality. Attention must also be

given to the concentration of extract, since it has an effect upon

the amount of hydrolysis obtained (173). All in all, the pro¬

cedure is one of typical clinical accuracy, but serves very well

for comparative purposes.

The determination of nitrogen in water and sewage is prop¬

erly included here, again because of its animal origin and its

relation to the history of the water under examination. Ordi¬

narily a sample of water is examined quantitatively for so-called

“albuminoid ammonia”, free ammonia, nitrite, and nitrate ni¬

trogen. Of these, the last two involve color reactions with or-

Schuette & Oppen — Determination of Organic Nitrogen 375

ganic dyes which need not be taken up here. Ammonia is de¬

termined by Nesslerization, with or without distillation, depend¬

ing on the amount present. The first-named, or “albuminoid ni¬

trogen”, is defined as that nitrogen which is liberated as am¬

monia by digestion with alkaline permanganate solution after

expulsion of any ammonia originally present. Its determina¬

tion was first proposed by Wanklyn {17 1^) in 1867 and, although

only of qualitative significance, and incapable of precise defini¬

tion, has nevertheless persisted these sixty-odd years as a meas¬

ure of the loosely combined organic nitrogen content of waters

and sewage.

Conclusion

And now we have indeed reached the end of this review of a

century or more of progress of the analytical aspect of nitrogen

in organic combination. From the pioneer flights of “pure”

chemistry to the practical routine of technology, scientists are

rendered invaluable aid by the practice of organic nitrogen anal¬

ysis. The chemistry of nitrogen is inseparable from the clothes

we wear and the food we eat. The chemistry of organic nitro¬

gen is the chemistry of life, and he who understands it has gone

a long way towards fathoming the mysteries of bodily economy.

The analytical or research chemist who has in his mind some

picture of the century-and-a-quarter panorama of evolution pre¬

ceding even our present limited knowledge cannot but have a

deeper appreciation for those who have gone before and for his

own humble work in consequence. It is the hope of the authors

that this review will prove a contribution to such a picture.

876 Wisconsin Academy of Sciences , Arts , and Letters.

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93. Frey, R. W., Jenkins, L. J., and Joslin, H. Ml., J. Am. Leather Chem.

Assoc., 23, 397 (1928).

94. Stock, W. F. K., Analyst. 17, 109, 152 (1892).

95. Thuau, U. J., and de Korsak, P., Collegium, 1910, 364; Chem. Zentr.,

1910 II, 1247.

96. Kruger, M., Ber., 27, 1633 (1894).

97. Foerster, O., Z. anal. Chem., 28, 422 (1899).

98. Possi-Escot, E., Compt. rend., 149, 1380 (1910).

99. Neuberg, C., Biochem. Z., 24, 423 (1910).

100. Justin-Mueller, E., Bull. sci. pharmacol. 23, 167, (1916).

101. (Davis, C. F., and Wise, M., Cereal Chem., 8, 349 (1931).

102. Mach, F., and Lepper, W., Landw. Vers. Stat., 109, 363 (1929).

103. Dafert, F. W., Relat. Inst. Agron., Sao Paulo, Brazil, 1892, 107;

Expt. Sta. Record, 4, 461 (1892).

Schuette & Oppen— Determination of Organic Nitrogen 379

104. Falk, K. G., and Sugiura, K., J. Am. Chem. Soc., 38, 916 (1916).

105. Fearson, W. R., Dublin J . Med. Sci., [4] 1, 28 (1920).

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108. Luhrig, H., Pharm. Zentralhalle , 62, 437 (19121).

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Chem. Abst., 21, 30 (1927).

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111. Hoppe-Seyler, F. A. Mikrochemie, 10, 446 (1932).

112. Folin, P., and C. J. Farmer, J. Biol. Chem., 11, 493 (1913).

113. Kohler, P. A., and Graves, S. S., J. Am. Chem. S'oc., 35, 1594 (1913).

114. Sors, P., Chem.-Ztg., 56, 156 (1932).

115. Rammelsberg, Ann. Physik Chem. (Poggendorf), 47, 153 (1839).

116. Nessler, Julius, Inaugural Dissertation, Freiburg, 1856; Chem.

Zentr., 1856, 534.

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118. Holland, P., Chem. News (Am. Rpt.), 2, 243 (1868).

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120. Schiff, H., Ann., 310, 25 (1899); 319, 59, 287 (1901); 325, 348

(1908).

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122. Malfatti, H., Z. anal. Chem., 47, 273 (1908).

1213. Wilkie, J. M., J. Soc. Chem. Ind., 29, 6 (1910).

1214. Rona, P., and Ottenberg, R., Biochem. Z., 24, 354 (1910).

125. Mink, P. J., and Brown, E. W., U. S. Naval Med. Bull., 4, 69 (1910).

126. Wright, A. M., Trans. Proc. New Zealand Inst., 42, 224 (1910).

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128. deGraaf, W. C., Pharm. Weekblad, 52, 1777 (1915).

129. Shaw, W. S., Analyst, 49, 558 (1924).

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109 (1912).

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(19219).

135. Tillmans and Kiesgen, J., Z. Nahr. Genussm. 53, 131 (1927).

136. Schuette, H. A., and Templin, Vera, J. Assoc. Official Agr. Chem., 13,

136 (1930).

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139. Klinkerfues, F., Chem.-Ztg., 35, 1274 (1911).

380 Wisconsin Academy of Sciences , Arts , and Letters .

140. Winkler, L. W., Z. angew . Chem ., 28 I, 231 (1913).

141. Scales, F. M., and Harrison, A. P., J. 7ra£. Chem., 12, 350 (1920).

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143. Ingle, J. D., Chemist- Analyst , 21, No. 12, 8 (1932).

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145. Rupp, E., and Rossler, E., Arch . Pharm ., 243, 104 (1905).

146. Artman, P., and Skrabal, A., Z. cwtaL Chem., 46, 5 (1907).

147. Willard, H. H., and Cake, W. E., J. Am. Chem. Soc ., 42, 2646 (19210).

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149 Lewi, B. J., J. Chem . Ind. (Russia), 8, 393 (1931) ; Chem. Zentr

1931 II, 278.

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152. Ronehese, A., Bull. soc. chim., [4] 3, 1135 (1908).

153. Grigaut, A., and Brodin, P., Compt. rend. soc. biol., 73, 458 (1913).

154. Heyninx, Biochem. Z., 51, 355 (1913).

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157. Jodlbauer, M., Chem . Zentr., 57, 433 (1886).

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160. Edwards, V., Chem. News, 65, 241, 265 (1892).

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164. Simek, B. G., Chem. Listy, 25, 322 (1931) ; Chem. Abst., 25, 5871

(1931).

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(1930).

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Leipzig and Vienna, 1933, p. 205; Mikrochemie, 13, 115 (1933).

167. Dorfmuller, G., Z. Ver . deut. Zuckerind., 80, 407 (1930).

168. Fleury, P,, and Levaltier, H., J. Pharm. chim., [7] 30, 265 (1925).

169. Van Slyke, D. D., Proc. Soc. Exptl. Biol. Med., 1, 46 (1910).

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Chem. Abst., 26, 3000 (1932).

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[2] 5, 445 (1867).

175. Sherman, H. C., “Methods of Organic Analysis”, Macmillan Com¬

pany, New York, 1912, 2 ed., p. 292.

THE CUPRO- ALKALI METAL CARBONATE SOLUTION

IN THE DETERMINATION OF REDUCING SUGARS.

II. A MODIFICATION OF PELLET’S SOLUTION*

Chang Y. Chang and H. A. Scbuette

University of Wisconsin

Some sixty years ago the suggestion1 was made that a solu¬

tion of a lesser degree of alkalinity than that characteristic of

Fehling’s solution2 be used for the determination of reducing

sugars in the presence of sucrose because of the hydrolytic ac¬

tion of this reagent upon the latter. This end was to be met by

the use of both sodium hydroxide and sodium bicarbonate in

such proportions that the former fall short of the stoichiometric-

al relationship which exists between these two compounds. Pel¬

let,3 unable to verify the claims made by the sponsor of this solu¬

tion for a passivity towards sucrose and a satisfactory stability

after a six months’ aging period, then proposed a modification

thereof. The novel features of his reagent were the use of so¬

dium carbonate and a stabilizing agent, ammonium chloride,

whose presence was briefly dismissed with a statement of the

purpose of adding it to the mixture.

Except for the attention given the first of these proposals by

Pellet, both, it seems, passed unnoticed and were soon forgotten.

This situation seemed to warrant an investigation of these for¬

gotten techniques, and particularly so because of the compara¬

tively recent revival of interest in this type of sugar-oxidizing

solution. As the result of such a study in this Laboratory there

has come a modification of Pellet’s solution in turn. This new

reagent is one which, unlike that of the current form of Fehl-

ing’s solution, does not in the course of time give any evidences

of having undergone auto-reduction nor does it exert a hydro¬

lytic action upon sucrose under the experimental conditions set

up in the investigation. Furthermore, it is somewhat more sen¬

sitive than the latter towards the common reducing sugars.

* Condensed from a thesis submitted by Chang Y. Chang to the faculty of the Graduate

School of the University of Wisconsin in partial fulfillment of the requirements for the degree

Doctor of Philosophy, 1933.

381

382 Wisconsin Academy of Sciences , Arts, and Letters.

The data relevant to the experimental development of this

reagent and a record of its copper-sugar equivalents are pre¬

sented in this communication.

Experimental

Stability of Pellet's Solution. — Pellet's original directions35

for the preparation of his solution were followed, viz : — CuSO*.

5HaO 68.7 g., Rochelle salts 200 g., N^COa 100 g., NH*C1 6.87 g.,

water, q.s. 1000 ml. It was early apparent that a finely divided

precipitate, its quantity increasing with age, will form in this

solution irrespective of the method employed in making the lat¬

ter. The reaction which is responsible for its formation is not

a reversible one for the precipitate in question is stable at ele¬

vated temperatures. It availed nothing to heat the freshly pre¬

pared solution in the hope of hastening equilibrium inasmuch

as the formation of the precipitate continued after the filtered

reagent has been set aside. That it is obviously formed at the

expense of the strength of the reagent became apparent (Table

I) when, from time to time, the filtered reagent (20 ml. plus

30 ml. water) was treated for 45 min. at 65° with a dextrose

solution containing 80 mg. of this sugar in 50 ml.

Table I

Effect of Age upon the Dextrose-Copper Equivalent of Pellet’s Solution

Tentative Modification.— The objectionable characteristics

of the foregoing reagent were corrected in a measure by follow¬

ing the course of procedure prescribed by Soxhlet4 in his modi¬

fication of Fehling’s solution,2 that is preparing it in two parts

to be mixed just before use5 in this instance in the proportion

of one volume of the copper solution to four of the alkaline tar¬

trate mixture. Solution A consisted of 343.5 g. CuS(X5H20 and

34.35 g. NILC1 in water q.s. one liter, whereas solution B, simi¬

larly, contained 250 g. Rochelle salts and 125 g. NazCCk After

letting the latter stand for some time at room temperature, or

overnight at 60-70°, to permit the separation of impurities,

mainly calcium carbonate, it was ready for use. Equilibrium

Chang & Schuette — Determination of Reducing Sugars 383

apparently had been reached in this time for after removal of

the precipitate the solution no longer deposited a sediment.

On putting this reagent to use, an unexpected situation arose

in that it was found impossible to duplicate satisfactorily the

sugar equivalents of this reagent when prepared exactly under

the same conditions but at different times, unless one chose to

disregard four to ten-mg. discrepancies as inconsequential, the

inevitable experimental error chargeable to the “personal equa¬

tion” of the operator.

The causes for this erratic behavior of the reagent appar¬

ently lay in some fault in its composition and, perhaps too, in

experimental conditions incapable of bringing out the maximum

efficiency of the reaction. Further refinements, such as might

be suggested by a study of the effects of shifting some of the

factors governing the reduction of the copper solution, were

obviously necessary. A resume of the results of such a course

of procedure follows.

Temperature Factor: — The optimum temperature conditions

for reduction were studied with two solutions of different alka¬

linity. By reducing the tartrate content of solution B to 216.25

g. per liter, a concentration exactly comparable to that of the

Soxhlet Fehling solution (yet less than that recommended by

Pellet by 27 g.) it was possible, because of solubility considera¬

tions affecting all of the ingredients, to increase its sodium car¬

bonate content to 238.5 g. and still higher as was done in a later

phase of the whole study.

Working at thermostatically controlled temperatures (±

0.1°) and using dextrose (Bureau of Standards sample 41) as

the experimental sugar under the conditions noted in preceding

paragraphs, it was found (Figure 1) that the rate at which the

copper solution is reduced is a function of the degree of its alka¬

linity and, of course, the temperature at which the reaction is

carried out. The solution of greater alkalinity was found to be

preferable to the lesser because under these conditions reduc¬

tion is apparently completed at a lower temperature, or 90°.

This point was, therefore, selected as the best working tempera¬

ture for the studies which followed.

384 Wisconsin Academy of Sciences , Arts, and Letters.

70 80 90

temperature in °C,

Fig. 1. — Effect of temperature upon the reducing power of dextrose

Time Factor. — Without changing the conditions set up in the

preceding experiments with respect to concentration of reac¬

tants and nature of reducing agent, and using the copper solu¬

tion of higher alkalinity, the time required to reach equilibrium,

or substantially so, was determined. Observations were made

at the end of a 20-minute reaction period and at stated intervals

thereafter up to 50 min. inclusive. From the data so obtained

it was apparent (Table 2) that the reaction practically reaches

equilibrium within 40 min. and that the 45-min. time interval

previously adopted was well chosen.

Chang & Schuette — Determination of Reducing Sugars 385

Table II

Effect of Time upon the Dextrose-Copper Equivalent

of Pellet’s Solution

known6 that the concentration of hydroxyl ion influences the

rate and amount of oxidation of the sugars. In spite of the

fact that a carbonate content equivalent to 4.5 N had already

been found to make for a satisfactory performance of the re¬

agent, yet it remained to determine whether this concentration

of sodium carbonate actually represented the most effective al¬

kalinity. To that end the whole gamut from 1 N to 7 N carbon¬

ate content was studied with respect not only to the oxidation of

dextrose, but levulose, maltose and lactose* as well, besides a

fifty-fifty mixture of the first-named pair (Figure 2).

It was found in this instance, too, as is the case with Fehl-

ing’s solution, that the amount of copper reduced increases with

increasing concentration of hydroxyl ions up to a point where

oxidation of the sugar molecule appears to have reached its max¬

imum. For levulose this condition apparently lies in the use of

a reagent whose carbonate content is 2 N, yet at the 4.5 N point

of alkalinity the curve begins to straighten itself out. This sit¬

uation is significant, for not only does this concentration seem

to represent the effective alkalinity for the oxidation of all the

other sugars in this group but it is also at this point that dex¬

trose, levulose and an equal mixture of the two exhibit practi¬

cally the same reducing action when present in the reaction

mixture to the extent of 80 mg.

Pellet's Solution in its Modified Form . — The relative concen¬

trations of copper sulfate and ammonium chloride with respect

to the whole reagent have been retained, but the desirability of

increasing the carbonate content made necessary the use of less

* These three sugars were of the highest degree of purity obtainable and _ conformed, on

analysis, to the specific rotation characteristic of each. They were dried at 56° in vacuo before

use.

386 Wisconsin Academy of Sciences , Arts , and Letters .

Fig. 2. — The influence of concentration of sodium carbonate

upon the reducing powers of various sugars

(1) Levulose (2) Dextrose (3) LeVulose-Dextrose

(4) Lactose hydrate (S) Maltose hydrate

Rochelle salts than once suggested. sb By preparing the reagent

in two parts, as described in preceding paragraphs, the instabil¬

ity of the original form1 and its tendency to erratic behavior

have been overcome. That portion containing the copper is not

photo-sensitive— solutions left standing on the shelf twelve

months or more with no protection from the sun-light have

shown no signs of decomposition— and hence is not susceptible

to the forces of auto-reduction because of the presence of the am¬

monium chloride which dissolves the cuprous oxide as fast as

it is formed. The practical significance of this is that no “blank”

determination on the reagent is necessary.

Chang & Schuette — Determination of Reducing Sugars 387

The optimum conditions for its use are: (1) time and tem¬

perature of reaction, 45 min. and 90°, respectively; (2) 20 ml.

of reagent mixed just before use in the proportion of one volume

of copper solution to four of tratrate; (3) a sugar solution con¬

taining not more than 4 mg. of reducing sugar per cc. ; and (4)

the reaction mixture diluted with water, if need be, to 80 ml.

Finally, the reduction should preferably be carried out in stop¬

pered flasks in order that the original volume of the reaction

mixture be maintained and temperature fluctuations at the sur¬

face be eliminated.

Sucrose shows no reducing action towards this reagent under

the experimental conditions herein set up. This fact was dis¬

covered when no cuprous oxide was formed in an 80-ml. reaction

mixture containing in one instance 80 mg. of sucrose and 2 g.

in another, nor was any significant increase in the copper equiv¬

alent of dextrose, levulose, and an equal mixture of the two

noted when this sugar was admixed with each to the extent of

five per cent, in one series of experiments and fifty per cent, in

another. Similarly, the reducing power of maltose was not af¬

fected by the addition of an equal weight of sucrose, yet lactose

gained slightly in its ability to reduce the copper solution under

the same conditions and when it constituted only 17 per cent,

of such a mixture. In view of the passivity of sucrose towards

this solution, it is difficult to explain this phenomenon except,

perhaps, on the basis of promoter action of the former.

Sugar-Copper Equivalents . — The relationship existing be¬

tween four of the common reducing sugars and the amount of

cuprous oxide, in terms of copper, precipitated by them from

this reagent under the conditions described above is a straight

line function. The relevant mathematical expressions are:

388 Wisconsin Academy of Sciences , Arts , and Letters .

The copper equivalent per unit weight of sugar was found

to be 2.9 for dextrose or levulose and, similarly, 1.3 and 1.2 for

lactose and maltose, respectively. That these equivalents point

to a greater degree of sensitiveness of the modified Pellet solu¬

tion than that shown by the same sugars towards Fehling's solu¬

tion is shown by the following comparable data: in the order

named above they are 2.08, 1.88, 1.23 and 1.01. The practical

significance of this lies in the fact, for example, that discrepan¬

cies in duplicate determinations of dextrose of 2.9 mgs. are in¬

consequential inasmuch as the analyses will still agree within

one mg. of sugar equivalent.

Summary

The experimental development of a sugar-oxidizing reagent

from the forgotten foundations laid by others1,3 many years ago

has been described with such attention to detail as seemed de¬

sirable to point out the fact that the inherent faults in the com¬

position of the latter and the technique of using it have been

corrected. The reagent in its present form consists of two parts

one of which contains in one liter of solution 343.5 g. CuSO*.

5H2O and 34.35 g. NILC1, the other, similarly, of 216.52 g. so¬

dium-potassium tartrate and 283.5 g. Na^CO. The copper equiv¬

alents of four reducing sugars have been determined by a mode

of procedure which involves an 80-ml. reaction mixture 20 ml.

of which is reagent (one volume of copper solution mixed im¬

mediately before use with four of tartrate solution), and the

remainder reducing sugar solution containing not more than 100

mgs. of dextrose or levulose or 220 mgs. of lactose or maltose,

the reduction being carried out in a stoppered flask immersed

for 45 min. in a 90° bath. What is deemed to be a unique char¬

acteristic of this reagent is its passivity towards sucrose under

the experimental conditions herein set down.

Literature Cited

(1873).

2. Fehling, Arch, physiol. Heilkunde, 17, 64 (1848) ; Ann.. 72, 106

(1849).

3. (a) Pellet and Champion, Compt. rend., 80, 181 (1875) ; (b) Pellet,

ibid., 86, 604 (1878).

4. Soxhlet, Chem. Zentr., [3] 9, 218 (1878) ; J. prakt. Chem., 129, 227

(1880).

5. C. Y. Chang, Master’s thesis, University of Wisconsin, 1932.

6. Quisumbing and Thomas, J. Am. Chem. Soc., US, 1511 (1921).

A STUDY OF LIGNEOUS SUBSTANCES IN

LACUSTRINE MATERIALS

John F. Steiner1 and Y. W. Meloche

University of Wisconsin

In past years limnologists in their study of the environments

of plant and animal forms in inland lake and sea waters have

analyzed waters for dissolved solids and gases and have at¬

tempted to determine the composition of lake and sea bottoms

as well as that of the existing biological forms. Until recently

these analyses were confined largely to inorganic constituents,

little attention being paid to the organic material. Although

Schuette (16) studied the ether extract of Daphnia as early as

1918, it was not until recently that Birge and Juday and Krogh,

studying inland lakes, and Waksman studying sea water (20)

made investigations designed to obtain more information con¬

cerning the types of organic compounds made available for fur¬

ther plant and animal consumption by decomposition of already

existing dead plants and animals in the water and in bottom

muds.

Birge and Juday (S) in “Particulate and Dissolved Organic

Matter in Inland Lakes” discuss the suspended and dissolved

organic matter in inland lake waters. They describe a system

of analysis in which the organic material is divided into pro¬

teins, fats and carbohydrates. It may be well to summarize the

method here.

A determination is made of the total organic nitrogen, total

organic carbon (A) and ether extract in the residues under ex¬

amination. Nitrogen is calculated to protein (B) by means of

the factor 6.25, its carbon content being taken as 53 per cent.

The carbon content of the fat fraction (C), or ether extract, is

taken to be 75 per cent. Calculation of the carbon due to car¬

bohydrate (D) which is 452 per cent of the total carbohydrate

is then made with the aid of the following formula :

D — A — 0.53 B - 0.75 C

1 This study was supported in part by a grant from the Wisconsin Alumni Research Founda¬

tion. The analytical results contributed by Lester Brillman and Leslie Gerlach were made possible

by CWA funds.

a In view of results more recently obtained, SO per cent has been used as the carbon content

of the carbohydrate.

389

390 Wisconsin Academy of Sciences, Arts , and Letters .

More recent work on ligneous materials indicates that in the

above mentioned carbohydrate fraction is included the less

easily decomposable ligneous materials. It therefore seemed de¬

sirable to determine whether or not the ligneous materials con¬

stitute a significant portion of the organic material in the orig¬

inal samples. The limitations of the above procedure are

apparent since at least part of the calculation is dependent on

the use of average values. It seems unlikely that a better method

will be obtained until a more accurate characterization of the

organic constituents is made available.

In 1933 Ohle (11) reported stratification of the sediment in

the Ukleisee near Plon by determining the methoxyl and ligne¬

ous content of different strata. For our present investigation

the corrected residue isolated from various samples by the sul¬

furic acid method of Scherrard and Harris (18) has been used

as a measure of ligneous material. Methoxyl determinations

were made on the original samples and on the lignin fractions

isolated by the acid treatment. They are presented for purpose

of comparison.

Condition of Samples

Samples of bottom mud from the deep parts of six Wisconsin

lakes; a sample of mud from Lake O’Malley, an Alaskan fresh

water lake; flora and fauna from Wisconsin lakes including

Isoetes, Gloeotrichia, Lobelia, and Moss, and several catches

each of net plankton and nannoplankton from the waters of

Lake Mendota were supplied by the Wisconsin Geological and

Natural History Survey for this study. The classification net

and nannoplankton is that used in Bulletin 64 published by the

Survey.

All mud samples were air dried at the time they were ob¬

tained and have been preserved in stoppered bottles. Such a

treatment is necessary to prevent fermentation of the organic

matter in the moist sediment. Samples of net and nannoplank¬

ton were dried in a Hear son* oven at 60 °C. Before analysis

the samples were dried further in a vacuum desiccator at 60° C

until no loss of weight could be observed. All results reported

are on the dry weight basis.

Mud samples were obtained with an Ekman dredge and rep¬

resent only the upper 15 cm. of the deposit.

* In the Hearson oven outside air is passed over the samples by means of a motor driven fan.

Steiner & Meloche — Lacustrine Lignin

Methods of Analysis

891

Total nitrogen in original samples and special fractions was

determined by the micro-Kjeldahl procedure described by Kem-

merer and Hallett.8

We were primarily interested in the examination of organic

matter in the original residues and in the so-called ligneous frac¬

tion. The presence of carbonates and hydrous silicates inter¬

feres with the estimation of total organic matter in the original

muds by loss on ignition. Hence total carbon was determined

by the micro-combustion method of Kemmerer and Hallett7 and

the carbonate carbon by a micro carbonate method described by

the same authors.9 The carbonate carbon was subtracted from

the total carbon leaving the organic carbon. It should be noted

that a survey of many determinations of organic carbon and

total organic matter has shown that a reasonably constant ratio

exists between total organic matter and organic carbon, namely,

organic carbon X 2 = organic matter. Limnologists commonly

use this method in order to obtain a close approximation of the

total organic matter. A slightly different factor has been pro¬

posed by Waksman for use on marine deposits, this value being

1.887.

In the case of ligneous fractions isolated by the sulfuric acid

method to be described later it was possible to determine the

total organic matter (lignin and insoluble protein) by ignition.

The acid treatment decomposed all carbonates and most of the

hydrous clays, the constituents which interfere with the direct

determination of organic matter in the original sample by igni¬

tion.

ii ' .»

Ignitions were made in a muffle furnace at about 700° C, the

sample being heated to constant weight.

The Lignin Content of Lacustrine Substances

Previous researches have indicated that lignin is the most

abundant organic identity present in bottom deposits. Such a

condition exists because of all the organic types present lignin

is the least decomposable by either aerobic or anaerobic bacterial

action and lignin does not readily serve as a food substance for

macroscopic animal or plant life.

The acid extraction method recommended by Sherrard and

Harris for the isolation of lignin seemed well adapted to the sort

392 Wisconsin Academy of Sciences, Arts, and Letters.

of material under investigation. According to them cold con¬

centrated sulfuric acid dissolves practically all oxygenous and

nitrogenous compounds from naturally occurring organic mix¬

tures, leaving lignin mixed with a relatively small amount of

crude protein. The residual nitrogen is probably present in a

ligno-protein complex. Temperature and concentration of the

sulfuric acid must be such as to remove the whole cellulosic part

without causing caramelizing. Acid which is too strong or too

warm may also sulfonate some lignin.

Method :

Add a dry two gram sample of algal or other plant material,

or a dry ten gram sample of bottom mud, to 16 times its weight

of cold 72 per cent sulfuric acid (10°C). A large vessel is re¬

quired to retain mud samples high in carbonate ; a four hun¬

dred cc. beaker is large enough for a ten gram sample of mud

from Lake Mendota. Although the carbon dioxide liberated

upon addition to the acid causes considerable foaming it is pos¬

sible to keep the material in the beaker. Maintain the acid wet

sample at 10° C for sixteen hours, shaking occasionally. Then

pour the mixture quantitatively into twenty five times its vol¬

ume of water in a 3 to 5 liter round bottom flask rotating the

vessel meanwhile in order to prevent local overheating. Boil

the resulting solution and residue for from 4 to 5 hours under

a reflux condenser to hydrolyse any carbohydrate which might

have been precipitated by dilution. Upon cooling the solution

should be quite clearly wine or red brown, the color being deeper

in organic rich mud. Filter through a small hard paper filter

disk by aid of suction. Wash the residue until free of sulfate.

Carefully remove the residue from the filter paper after it has

become caked, transferring it to a tared watch glass. Dry in

the same manner as the original sample was dried (vacuum

desiccator at 60° C) and weigh. A sample of this dry residue

which contains lignin, together with small amounts of nitrogen¬

ous substance and varying quantities of silica and other inor¬

ganic materials, is then ignited in a muffle furnace at about

700° C for thirty to forty minutes. The loss in weight repre¬

sents the total organic matter in the residue.

After the prescribed acid extraction no samples showed any

evidence of charring or caramelizing. A second extraction with

Steiner & Meloche — Lacustrine Lignin 393

concentrated sulfuric acid did not remove any additional or¬

ganic matter ; hence a single treatment may be considered com¬

plete. The final product is dark brown yielding a pure white

ash on ignition. No sulfur trioxide fumes were detected during

the ignition ; indicating that little or no sulfonation of the lignin

occurred, and that inorganic sulfate had been completely re¬

moved.

The sulfuric acid digestion, besides removing all carbohy¬

drate and a large part of the protein, effected the solution of

most of the mineral constituents except silica. Analyses of the

ash remaining from the ignition of ligneous residues are given

in the following table :

Analysis of Ash from Ligneous Residues

In lake bottom detritus the polysaccharides and hemi-eellu-

loses have been almost completely removed by bacterial and

possibly hydrolytic action. Such a condition of the sample ob¬

viates a preceding acid hydrolysis for removal of pentosans or

similar carbohydrates which are alleged to contribute a poly¬

merization product to the lignin residue.*

Results

Analyses of the ligneous residues obtained by the sulfuric

acid extraction are reported in Table I. As previously noted

the organic fraction of this residue was determined by loss upon

ignition. At first glance it might appear that this value is ir¬

relevant. It must be noted, however, that it is necessary to

know the organic fraction in the ligneous residue before it is

possible to calculate the corrected lignin. In the calculations

represented in Table I nitrogen was determined on the ligneous

residue and this was converted to an approximate crude pro¬

tein value by the use of the factor 6.25 X N. The crude protein

subtracted from the total organic matter in the ligneous residue

gave the so-called corrected lignin. The carbon in the crude

* Norman (JO) and others have advised a dilute acid hydrolysis preceding the 72 per cent

sulphuric acid digestion in the process of isolating lignin from plant material and composts in

order to remove xylan and any other pentosans present which are converted in part by strong

sulphuric acid into so-called apparent lignin.

394 Wisconsin Academy of Sciences, Arts, and Letters.

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395

protein was calculated by the use of the average value of 53 per

cent. This carbon subtracted from the total organic carbon in

the ligneous residue gave the carbon due to lignin. Finally, the

carbon due to lignin divided by lignin gave the percentage of

carbon in lignin.

From the nature of the isolation of the ligneous residue and

from the method of calculation described it must be obvious that

the values given must be considered approximations. However,

if the values reported in this and following tables give the gen¬

eral order of magnitude of the carbon in lignin and the order

of magnitude of the lignin in the organic part of the original

samples, their purpose will be served.

From an examination of the organic content of the ligneous

residue in Table I it is obvious that the acid extraction pro¬

duces a ligneous residue which is strongly contaminated with

inorganic constituents, principally silica. The percentage of

carbon in lignin of many woods has been found to be somewhat

below 65 per cent. In fresh water lake muds the carbon in the

lignin as determined by the technique already described ranged

from 40.2 per cent for the mud samples from Lake O’Malley to

63.9 per cent for the mud from Lake Mary. Greater agree¬

ment is noted for the carbon in the lignin for other samples, the

values from net plankton and nannoplankton being 63.6 per

cent and 62.4 per cent, respectively. Four other samples, Lo¬

belia, Gloeotrichia, Isoetes, and Moss gave an average carbon in

lignin of 60.9 per cent. The relatively great divergence of re¬

sults for carbon in lignin in the lake muds might suggest that

the sulfuric acid treatment outlined, while suited for undecom¬

posed woods, was unsuited for the quantitative isolation of lig¬

nin in the heterogeneous bottom muds. At the same time it

must be remembered that in the list of lakes is represented both

drainage and seepage lakes as well as hard and soft waters, high

and low color, and high and low dissolved and suspended or¬

ganic material. It must also be noted that the organic matter

in lake muds represents material which has resulted from the

decomposition of plant and animal forms. Most investigators

who have reported more regular results have worked with orig¬

inal materials such as wood. Although there was this wide di¬

vergence in carbon in lignin between the various lake mud sam-

396 Wisconsin Academy of Sciences , Arts , and Letters .

pies from different lakes, no difficulty was experienced in

checking the results obtained on an individual sample.

Perhaps of more interest to the limnologist is the ratio of

lignin to total organic matter in these inland lake samples or

since according to the calculation already described the lignin

was formally included in the carbohydrate fraction of the or¬

ganic matter in lake residues it should be important to make

some estimation of the ratio of lignin to carbohydrate.

In Table II the weight of the original samples and the weight

of the sulfuric acid residues are given to illustrate the size of

the residues handled. With the exception of two columns the

remainder of the data is given in percentages. Although per¬

centage lignin in the original material is given this value has

less importance than the calculated lignin organic matter ratio

since the different samples contain different amounts of inor¬

ganic residue. The ratio between lignin and organic matter in,

the samples is calculated by dividing the corrected lignin in the

original material by the total organic matter in the original ma¬

terial, the total organic matter being obtained by multiplying

the total organic carbon by 2.00.

The highest lignin content of the muds examined was 29.67

per cent in Lake Mary mud; the lowest, 4.5 per cent in Lake

Mendota mud. The highest lignin to organic matter ratio oc¬

curs in Lake Mary mud where 47.76 per cent of the organic

matter was found to consist of lignin; the lowest ratio was

29.75 per cent in Lake Mendota mud. The other materials ex¬

amined which are the precursors to the organic substance of

the mud were all relatively lower in lignin, the average lignin

content of all being 17.5 per cent. These are results which might

be anticipated from a consideration of the course of decomposi¬

tion to which organic matter is subject after it reaches the

bottom.

Methoxyl Content of Lacustrine Lignin

In order to obtain additional information regarding the na¬

ture of lacustrine lignin and if possible to find a shorter means

of measuring the relative amounts of lignin in the original sam¬

ples, the methoxyl content of the original materials previously

studied was determined and compared with the methoxyl con¬

tent of the lignins obtained from them. It was hoped that the

Table II

LIGNIN CONTENT OF SOME LACUSTRINE SUBSTANCES

Steiner & Meloche — Lacustrine Lignin

397

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398

Wisconsin Academy of Sciences , Arts , and Letters,

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Steiner & Meloche — Lacustrine Lignin 399

methoxyl content of the original material might be an index of

the lignin in the sample. Clark's modification of the Vieboch

and Schwappach method for methoxyl was used for the samples

in question. The results obtained are recorded in Table III.

The ratio of the percentage of methoxyl in the sulfuric acid res¬

idue to percentage of lignin in this residue gives the percentage

of methoxyl in lignin, i.e. the methoxyl in the ligneous residue

from Lake Mendota mud was 0.26 per cent and the percentage

of lignin in the residue was 10.89 per cent; hence, 0.26/10.89

X 100 — 2.39 per cent or the methoxyl in lignin for this partic¬

ular sample. The assumption is made in these calculations that

all of the methoxyl in the sulfuric acid ligneous residue is due

to lignin ; that the crude protein left in the lignin fraction con¬

tributes no methoxyl. In examining the percentages of meth¬

oxyl in lignin and comparing these values to those reported in

the literature note that wood lignin is reported to contain 17

per cent to 26 per cent methoxyl while some straws are reported

to contain lignin with as little as 5 per cent methoxyl. In the

lignin recovered from lake muds the percentage of methoxyl in

lignin was considerably less than 5 per cent. The original wood

examined by Sherrard and Harris contained 5.3 per cent meth¬

oxyl while isolated lignin contained 82.4 per cent of this. An

examination of the data in Table III shows that no such recov¬

ery was experienced in the investigation of sub-aquatic types.

Not only was there little regularity in the ratio of the methoxyl

in lignin to that in the original sample for the bottom muds;

there was also considerable variation in the methoxyl content

of the lignin recovered from the bottom muds. The methoxyl

due to lignin constitutes only one third to two thirds of the total

methoxyl in the muds from Wisconsin lakes. In Lake O’Mallley

(Alaskan) mud, in which the decomposition of proteins, carbo¬

hydrates, fats and other organic types is presumably slower

than in the warmer Wisconsin lakes, the lignin portion of the

organic part of the sample is smaller and the proportion of

methoxyl due to lignin is correspondingly smaller. Only 0.04

per cent of the Lake O'Malley mud was methoxyl due to lignin

and this was only 0.15 of the total methoxyl in the mud. The

results for the net and nannoplankton show slightly greater

uniformity than do those for the lake muds. For the net plank¬

ton 7 to 11 per cent of the total methoxyl was in the ligneous

400 Wisconsin Academy of Sciences, Arts, and Letters.

residue. However, the corrected lignin content for the same

samples was 8 to 14 per cent. In the case of the three nanno-

plankton samples the methoxyl in the ligneous fraction was 6 to

12 per cent of the total in the sample.

An examination of the column, per cent CHsO in corrected

lignin, reveals that the lignin recovered by the acid process de¬

scribed contained a comparatively small amount of CHsO. The

question arises, is this small methoxyl content due to a too dras¬

tic treatment or is it due to the nature of the samples? Al¬

though the recovery of methoxyl is not high, i.e., the CHsO found

in the ligneous residue compared to total CHsO in the original

sample is low (column 7), it must also be noted that the CHsO

in the original sample is unusually low.

Conclusions

If one may assume that the sulfuric acid method for the iso¬

lation of lignin from sawdust may be applied with a reasonable

degree of success to samples of lake muds and miscellaneous

lake types, the data in this report give a fair approximation of

the relative amounts of lignin in lake muds, net and nanno-

plankton, and a few additional lacustrine types. Of more im¬

portance, data are given to show what portion of the total or¬

ganic matter of the samples is ligneous material. For lake muds

30 to 48 per cent of the total organic matter is lignin, for net

plankton 18 per cent of the organic matter is lignin, and for

the nannoplankton 10 to 20 per cent of the organic matter is

lignin. This information is of interest to the limnologist since

the ligneous fraction represents material which is not readily

available for plant and animal food.

Regarding the ligneous fraction isolated, it is of some signi¬

ficance that the percentage of carbon in the corrected lignin

(column 8, Table I) for samples which have undergone the

least decomposition approach the high value of 64 per cent, i.e.,

the carbon content of lignin from lake muds is low for the lake

muds and relatively high for the net and nannoplankton and

special samples. In considering the low methoxyl content of

the lignin from the lake muds, one may explain this in part by

the histories of the various samples. Particularly for the bot¬

tom muds, the organic matter has been exposed to extreme

chemical conditions. If the lignin undergoes gradual decom-

Steiner & Meloche — Lacustrine Lignin 401

position, the most labile functional groups might reasonably be

expected to be attacked first; hence, a slow hydrolysis or other

change may have caused the alteration in the nature of lignin

which has remained for years on the lake bottom.

From the relatively small number of samples examined it

does not seem likely that the methoxyl content of the samples

is any index of the amount of lignin present. As previously

noted, this may be due to the method of isolating the lignin or

to the previous history of the samples.

The above material is presented as a preliminary report. It

does not seem that conclusive results can be obtained until a

greater variety of samples is examined and a more satisfactory

technique perfected.

Literature Cited

1. Birge, E. A. and Juday, C. Organic Content of Lake Water. U. S.

Bur. Fisheries Bull., U2, 185 (1926).

2. Birge, E. A. and Juday, C. The Organic Content of the Water of

Small Lakes. Proc. Am. Phil. Soc., 66, 357 (1927).

3. Birge, E, A. and Juday, C. Particulate and Dissolved Organic Mat¬

ter in Inland Lakes. Ecology, U, 440 (1934).

4. Birge, E. A. and Juday, C. Inland Lakes. Wis. Geol. Nat. Hist.

Survey Bull. 6U, (1922).

5. Black, C. S. Chemical Analysis of Lake Deposits. Trans. Wis. Acad.

Sci., 2Jf, 127 (1929).

6. Clark, E. P. The Viebock and Schwappach Method for the Deter¬

mination of Methoxyl and Ethoxyl Groups. J. Assoc. Official

Agr. Chem., 15, 136 (1932),

7. Kemmerer, G. and Hallett, L. T. An Improved Method of Organic

Micro Combustion. Ind. Eng. Chem., 19, 173 (1927).

8. Kemmerer. G. and Hallett, L. T. Improved Micro Kjeldahl Ammonia

Distillation Apparatus, ibid., 1295.

9. Kemmerer, G. and Hallett, L. T. Micro Determination of Carbonate

Carbon, ibid., 1352.

10. Norman, A. G. and Jenkins, S. H. A New Method for the Deter¬

mination of Cellulose Based upon the Removal of Lignin and

Other Incrusting Materials. Biochem. J., 27, 819 (1933).

11. Ohle, W. Chemische-Stratographische Untersuchungen der Sedi-

mentmetamorphose eines Waldsees. Biochem. Z., 258, 420 (1933).

402 Wisconsin Academy of Sciences , Arts, and Letters.

12. Phillips, M„ The Quantitative Determination of Methoxyl, Lignin,

and Cellulose in Plant Materials. J. Assoc. Official Agr. Chem.,

15, 124 (1932).

13. Phillips, M,. Report on Lignin, ibid., 16, 476 (1933).

14. Ritter, G. J. Chemistry of Wood. VII. Relation between Methoxyl

and Lignin in Wood. Ind. Eng. Chem., 15, 1264 (1923).

15. Ritter, G. J., Seborg, R. M. and Mitchell, R. L. Factors Affecting

Quantitative Determination of Lignin by the 72 Per Cent Sul¬

furic Acid Method. Ind. Eng. Chem. Anal. Ed. 4, 202 (1932).

16. Robinson, R. and Kemmerer, G. The Determination of Kjeldahl Ni¬

trogen in Natural Waters. Trans. Wis. Acad. Sci., 25, 123 (1930).

17. Schuette, H. A. A Biochemical Study of the Plankton of Lake Men-

dota. ibid., 19, 594 (1918).

18. Sherrard, E. C. and Harris, E. E. Factors Influencing Properties of

Isolated Wood Lignin. Ind. Eng. Chem., 24, 103 (19132;).

19. Titus, L. and Meloche, V. W. A Microextractor. Ind. Eng. Chem.

Anal. Ed., 5, 286 (1933).

20. Waksman, S. A. Die Chemische Zusammensetzung von Torfbildungen

und Torfarten und ihre Untersuchungsmethode. Brennstoff-

Chem., 11, 277 (1930).

21. Waksman, S. A. A Critical Study of the Methods for Determining

the Nature and Abundance of Soil Organic Matter. Soil Sci.,

30, 97 (1930).

22. Waksman, S. A. On the Distribution of Organic Matter in the Sea

Bottom and the Chemical Nature and Origin of Marine Humus,

ibid., 36, 125 (1933).

23. Waksman, S. A. and Stevens, K. R. A System of Proximate Chemi¬

cal Analysis of Plant Materials. Ind. Eng. Chem. Anal. Ed., 2,

167 (1930).

PROCEEDINGS OF THE ACADEMY

Sixty-third Annual Meeting

The sixty-third annual meeting of the Wisconsin Academy of Sciences,

Arts and Letters was held in joint session with the Wisconsin Archeo¬

logical Society and the Midwest Museums Conference, at the University

of Wisconsin, April 7 and 8, 1933. The following program of papers,

special lectures and demonstrations was presented:

Friday morning , Section A.- — -Duane H. Kipp, Systematizing Wiscon¬

sin place names; Albert 0. Barton, The ransom of the Hall sisters; F. W.

Harris, The queriosites of coins; Joseph Schafer, The Stockbridge Indians;

Herman Kerst, Postage stamp printing; Louise P. Kellogg, A Wisconsin

Indian agent, Nicolas Boilvin; Kermit Freckman, Pleasant Lake Indian

mounds; Vivian Morgan, Old time “memory wreaths”; Harold A. Engle,

The “On Wisconsin” broadcasts; Donald Newton, An adult hobby show.

Friday morning , Section B. — -V. C. Finch, Characteristics of occu-

pance on the Mississippi delta fringe; E. F. Bean and A. R. Ostrander,

Hamilton Mounds quartzite in Adams County; Joseph Wanenmacher, W.

H. Twenhofel and Gilbert 0. Raaseh, Some new facts on the palaeozoic

rocks of the Baraboo region; Glenn T. Trewartha, Sequent occupance on

the Prairie du Chien terrace: a study in historical geography; Eric R.

Miller, Fluctuations of rainfall in Wisconsin, 1836-1932; L. R. Wilson,

Historical and climatic correlation of a Madison white oak log; O. A.

Mortensen, The cerebrospinal fluid and the cervical lymph nodes; H. W.

Mossman, Implantation, fetal membranes and placentation in the squirrels

(Sciuridae) ; L. H. Rubnitz and A. M. Katz, Changes in the testes and

accessory glands of the gray and fox squirrels during their seasonal re¬

productive cycle; R. C. Miullenix, Neuro-histological preparations: organ

of corti of monkey, retina of monkey, purkinje cells with pericellular

baskets, etc.; W. E. Tottingham and R. Nagy, Chemical changes associ¬

ated with discoloration of cooked potatoes.

Friday afternoon , Section A. — Theo. T. Brown, A fish-head effigy pipe;

Marguerite F. Stiles, Engraved catlinite tablets; John J. Knudsen, An

Indian flint implements file; Alice I. Vinje, Norwegian childhood tales;

E. Ralph Guentzel, Large native copper knives; Chas. L Emerson, Lieut.

James Gorrell, diplomat and strategist; W. F. Bauchle, The mound-build¬

ers; Charles E. Brown, Two Wisconsin animal pipes; Gilbert O. Raaseh

and Fred Wilhelm, A new and inexpensive type of museum display; Wm.

W. Bartlett, The Cornell University lands in the Chippewa valley; Bind¬

ley V. Sprague, Medicine among the North American Indians.

Friday afternoon , Section B. — Margaret B. Siler, Development of

spore walls in Sphaero cargos; Frank H. Smith, Chromosome associations

in Brodiaea ; Fred W. Tinney, Structure and behavior of chromosomes in

Sphaero cargos; Jeanette Jones, Ordovician starfishes of Wisconsin; Nor¬

man Schmeichel, Sex differentiation in the lined snake, Tropidoclonion

408

404 Wisconsin Academy of Sciences , Arts, and Letters.

lineatum (Hallowell) ; Olga A. Smith, The effect of salt on Bacillus mega¬

therium) A. H. Maslow, Psychoanalysis and mental hygiene as social

philosophies; Alfred Senn, The literary German language and its relation

to the German dialects; Ernest Voss, German Bible translations originat¬

ing in the territory of the Order of the German Knights about the middle

of the fourteenth century (by title).

Special lecture on Friday. M. E. Diemer, Wisconsin wild flower pho¬

tography.

Saturday morning , Section A. — Edgar P. Doudna, The banking prob¬

lem and the constitution of Wisconsin; Lewis Severson, The gold standard;

Edward Bennett, A plan for the restoration of employment.

Saturday morning , Section B. — N. C. Fassett, One day’s botanizing

near Madison; H. A. Schomer, Photosynthesis of algae at various depths

in the lakes of northeastern Wisconsin; L. R. Wilson, Lake development

and its effect on the quantity and quality of aquatic plant life; John

Steenis, Planting of walleyed pike in northern Wisconsin lakes; Edward

Schneberger, Age and growth of gamefish in Wisconsin waters; S. X

Gross, Studies on the parasites of northern Wisconsin fish.

Special lectures on Saturday. J. H. Mathews, Scientific methods of

crime detection; R. G. Mhllenix, Evolution and progress.

The annual business meeting of the Academy was held immediately

following the general session, April 8. The secretary presented the fol¬

lowing report on membership as of April 7, 1933: honorary members, 3;

life members, 12; corresponding members, 16; active members, 307; total,

339. Membership losses during the year: deceased, 5; resigned, 2:8; drop¬

ped for non-payment of dues, 24; dropped for loss of address, 3. Applica¬

tions for membership from forty-two individuals being presented, the sec¬

retary was unanimously instructed to cast the ballot of the Academy in

their favor. The list follows: Lawrence J. Berner, West DePere; Paul

Carroll, Milwaukee; Francis H. Clabots, West DePere; George Claridge,

West DePere; Hilary J. Deason, Ann Arbor, Mich.; L. A. Dobbelsteen,

Luxemburg; Sears P. Doolittle, Washington, D. C.; A. A. Drescher, Fen-

nimore; F. J. B. Duchateau, Green Bay; Wm. R. Duden, Ann Arbor,

Mich.; Paul L. Errington, Ames, la.; L. H. Halverson, Marquette, Mich.;

Hance F. Haney, Madison; Carl Hoffman, Appleton* V. K. Kapingen,

Milwaukee; Cornelius J. Kirkfleet, Somonauk, Ill.; T M. Langley, Super¬

ior; Jos. B. Layde, West DePere; Harold LeMahieu, West Allis; Jos.

McCaffrey, West DePere; David J. Mack, Madison; S. L. Mains, Milwau¬

kee; Sister M. Cassiana Marie, Green Bay; Jos. A. Marx, Green Bay;

H. J. Mellum, Kenosha; Mrs. A. C. Neville, Green Bay; Peter P. Pritzl,

West DePere; Paul P. Rhode, Green Bay; R. K. Richardson, Beloit;

Gregory R. Rybrook, West DePere; W. B. Sarles, Madison; Paul L. Sav-

ageau, West DePere; J. P. Schumacher, Green Bay; Lewis E. Severson,

Beloit; Olga A. Smith, Appleton; Mary E. Storer, Beloit; H. L. Traeger,

West DePere; M. J. Vanden Elsen, Brussels; R. P. Wagner, West De

Pere; E. J. Westenburger, Green Bay; P. W. Wilson, Madison; ML J.

Windt, West DePere. The nominating committee, consisting of E. B.

Proceedings of the Academy

405

Skinner, Arthur Beatty and E. M. Gilbert, reported nominations for the

various offices, and on motion the nominees were unanimously elected for a

term of three years: President , Rufus M. Bagg, Appleton; Vice-Presi¬

dent in the Sciences , Storrs B. Barrett, Williams Bay; Vice-President in

the Arts , A. M. Keefe, West DePere; Vice-President in Letters , A. R.

Hohlfeld, Madison; Secretary -Treasurer, H. A. Schuette, Madison; Li¬

brarian, Walter ML Smith; Curator , Charles E. Brown, Madison. The

treasurer’s report, as of March 31, 1933, was presented as follows:

Receipts

Balance in State Treasury, March 31, 1932 . $1478.39

State appropriation for 1932-33 . 1000.00

Dues received from members . . 526.00

Wis. Geol. & Nat. Hist. Survey for printing . 2)30.00

Annual allowance from A. A. A. S . 100.00

Endowment fund income . . . 116.43

University of Wisconsin for printing . 60.00

Academy publications sold . . . 40.45

Reprints sold to authors . . . 42.85

Miscellaneous . . . . . 1.52

Total . $3595.64

Disbursements

Printing of Vol. 27 of Transactions . $1426.54

Etchings and half-tones . 104.05

Reprints for Yol. 27 of Transactions . 373.66

Other printing (programs, announcements, etc.) . . 31.39

Secretary’s salary for the year . 200.00

Postage . 45.00

Express . 15.81

Total disbursements . . . $2196.45

Balance on deposit, March 31, 1934 . 1399.19

Total . .$3595.64

Endowment Fund

Trust agreement, Central Wis. Trust Co . . . $1000.00

City of Madison bond . 500.00

Rock County Highway bond . . . 500.00

Chapman Block bonds . 400.00

Commonwealth Telephone Company bonds . 400.00

Wisconsin Power and Light Company bonds . 200.00

Capitol Square Realty Company bonds . . . 200.00

Certificates of deposit . . . . . . 48.50

Cash on hand . 12.16

Total . . . . . . . . . $32)60.66

406 Wisconsin Academy of Sciences, Arts, and Letters .

The auditing committee, consisting of Arthur N. Bragg and N. C.

Fassett, reported that it had examined the accounts of the treasurer and

had found them correct.

The annual dinner was held at the University Club on Friday evening

with approximately fifty people in attendance. After the dinner Prof.

Charles E. Allen, retiring president of the Academy, presented an address

entitled “The Course and Significance of Sexual Differentiation”.

The following members of the Academy died during the past year:

W. B. Cairns, Aug. 2, 1932.

Carl Russell Fish, July 10, 1932.

C. Dwight Marsh, April 23, 1932.

Dana C. Munro, Jan. 13, 1933.

Huron H. Smith, Feb. 25, 1933.

Lowell E. Noland

Secretary-Treasurer

SIXTY-FOURTH ANNUAL MEETING

The sixty-fourth annual meeting of the Wisconsin Academy of Sci¬

ences, Arts and Letters was held, in joint session with the Wisconsin Ar¬

cheological Society and Midwest Museums Conference, at Lawrence Col¬

lege, Appleton, April 6 and 7, 1934. The meeting was formally opened by

an address of welcome by Dr. Henry M. Wriston, president of Lawrence

College, after which the following program of forty-two papers and lec¬

tures was presented:

Friday morning , Section A. — H. R. Holand, Mogachutes, a village of

the Stone Age; C. E. Brown, A large stone adze; W. A. Titus, Early

Wisconsin tribesi Louise P. Kellogg, The Winnebago visit to Washington

in 1828.

Friday morning, Section B. — L. F. Graber, The white grub menace in

Wisconsin; L. F. Graber, Leaf -hopper ( Empoasca fabae) populations as

related to cutting treatments of alfalfa; H. A. Schuette, A medieval honey

law.

Friday morning , general session. — Aldo Leopold, Wild life research in

Wisconsin.

Friday afternoon, Section A. — H. R. Holand, Michingan, the haven of

refuge; L. S. Buttles, The destruction of mounds in certain southern

states; John B. MacHarg, Stereoptican and bulletin board in museum

work; Wm. F. Raney, The Grignon family; Geo. Overton, Indian cooking

Proceedings of the Academy

407

stones; J. 0. Frank, Superstitions of the Fox River valley; Nile J. Behn-

cke, Two Indian legends; C. H. Hocking, Ridgeway ghost tales; Gene

Sturtevant, The dream dance of Keshena; Lorraine C. Brown, Mounds

of the University of Wisconsin arboretum; A. L. Kastner, Recent con¬

tribution to the origin and history of siphilis.

Friday afternoon , Section B. — Louis Kahlenberg, Neal Johnson and

A. W. Downes, On the activation of gases by metals; Eric R. Miller, Dust-

fall of November 12, 1933, in Southern Wisconsin; G. W. Woolley, (intro¬

duced by L. J. Cole), Early introduction of cattle into Wisconsin (before

1800); N. C. Fassett, Niagara limestone and its flora; L. E. Casida

(introduced by L. J. Cole), Endocrine stimulation of ovarian development

in farm animals; J. P. von Grueningen, Goethe in American periodicals

1860-1900; Arthur H. Weston (introduced by R. M. Bagg), The date of

Christmas ; V. W. Mleloche, The polarograph, a new tool in analytical

chemistry ; Ernest Voss, Goethe monuments in America (by title) .

Saturday morning, Section A. — A. H. Griffith, Abraham Lincoln ; Su¬

san B. Davis, The Wisconsin tercentennary celebration ; Francis S. Day-

ton, Indian fishing camps of the Wolf River ; H. E. Mansfield, Drama and

the folk spirit of Wisconsin; R. N. Buckstaff, Meteorological display for

museum purposes.

Saturday morning, Section B. — B. W. Roland (introduced by J. H.

Mathews) , The application of colloid chemistry to industrial problems ;

H. F. Lewis, The chemical behavior of the various components of wood

fiber; Loyal Durand, Jr., (introduced by V. C. Finch) , The geographic

regions of Wisconsin; K. Bertrand (introduced by V. C. Finch), A re¬

gional interpretation of woodland and forest land of Wisconsin! J. Riley

Staats, The geography of the central sand plain of Wisconsin ; L. P. Coo-

nen (introduced by A. M. Keefe), Seasonal variations in Sargassum fili-

pendula; J. C. McCaffrey (introduced by A. M. Keefe) , Some observations

on the true use of antiseptics ; J. J. Davis, Notes on parasitic fungi in

Wisconsin. XX. (by title) .

The Saturday morning sessions were preceded by a tour of inspection

of the buildings of the Institute of Paper Chemistry.

The annual business meeting of the Academy was held Friday, April

6, at 4:80 P.M. The secretary presented the following report on member¬

ship : honorary members, 3; life members, 12; corresponding members,

14; active members, 353; total, 382. Membership losses during the year:

deceased, 2; resigned, 7; dropped for non-payment of dues, 20. Applica¬

tions for membership from eighteen individuals being presented, the secre¬

tary was unanimously instructed to cast the ballot of the Academy in their

favor. The list of newly elected members, which includes four elected on

Nov. 18, 1933, by council action, follows : Donald Cameron, Racine ; Orville

Carey, Appleton ; Ethel Carter, Appleton ; George H. Conant, Ripon; Mil¬

ford A. Cowley, La Crosse ; John T. Curtis, Waukesha; John P. von Grue¬

ningen, Madison; Norris F. Hall, Madison; Erma N. Henry, Appleton ;

Loren C. Hurd, Madison ; M. R. Irwin, Madison ; Hilda C. Jorgensen,

408 Wisconsin Academy of Sciences , Arts, and Letters .

Appleton; Clement D. Ketchum, Appleton; Aldo Leopold, Madison; Harry

F. Lewis, Appleton; Ralph S. Nanz, Waukesha; Wm. F. Raney, Apple-

ton; E. Margaret Ritchie, Appleton; Isabel Schilling, Green Bay; George

F. Sieker, Madison; J. Riley Staats, Madison; John Voss, Peoria. The

status of both George C. Comstock, Beloit, and John R. Commons, Madi¬

son, was changed from that of active to corresponding members. Com¬

mittee appointments were announced as follows: Publication , the presi¬

dent and the secretary, ex officio, Arthur Beatty, Madison; Library, the

librarian, ex officio, A. W. Schorger, Madison, Mrs. A. C. Neville, Green

Bay, W. S. Marshall, Madison, A. L Barker, Ripon; Membership, the

secretary, ex officio, R. C. Mullenix, Appleton, J. O. Carbys, Milwaukee,

P. W. Boutwell, Beloit, G. W. Keitt, Madison.

The secretary-treasurer reported informally on the present condition

of the Academy’s finances and suggested, inasmuch as a final appeal for

a printing subsidy from the State had not yet been acted upon, that a

formal report be made at a later date. V. W. Mleloche and L. E. Noland

were appointed to audit the treasurer’s books. The question of the place

of meeting for next year was left to the Council for decision. A vote of

thanks was tendered the authorities of Lawrence College for placing the

facilities of the College at the disposal of the Academy.

Proceedings of the Academy

409

Receipts

Balance of receipts from State Treasurer . . . $ 308.31

Dues received from members . 557.64

Annual allowance from A. A. A. S . 94.50

Academy publications sold . 61.20

Reprints sold to authors . . 65.15

Interest on investments . . . . . . . 2:22.59

Wisconsin Archeological Society . 3.05

Certificates of deposit . 89.66

Securities matured . 1500.00

Total . $2902.10

Disbursements

Secretary’s allowance for the year . $200.00

Expenses of Appleton meeting . 83.40

Miscellaneous expenses . 25.28

Printing of programs, etc . 33.25

Securities purchased . 1545.00

Total . $1836.93

Balance in treasury . . . 1065.17

Endowment Fund

Home Owners Loan Corporation bonds . . $1050.00

Rock County Highway bond . 500.00

U. S. Treasury bond . 500.00

Commonwealth Telephone Company bonds . 400.00

Chapman Block bonds (in default) . . . 400.00

Wisconsin Power and Light Company bonds . 200.00

Capitol Square Realty Company bonds . 200.00

Cash . . 2591.87

Total . . . . . . . $3509.87

July 31, 1934.

The annual dinner was held in Ormsby Hall with seventy-five in at¬

tendance. President Bagg spoke informally on the history of the Academy.

Following this the assembled guests adjourned to Lawrence Memorial Cha¬

pel to listen to a public lecture by Prof. Laurence M. Gould of Carleton

College on the subject “Adventures in Antarctic Geology”.

The following deaths were reported:

Frank P. Hixon, Oct. 24, 1931.

A. R. Braun, June 21, 1933.

H. A. SCHUETTE

Secretary-Treasurer

esS*5