TRANSACTIONS

Si '

OF THE

WISCONSIN ACADEMY

OF

SCIENCES, ARTS AND LETTERS

VOL. XXXVIII

NATURAE SPECIES RATIOQUE

MADISON, WISCONSIN

TRANSACTIONS

OF THE

WISCONSIN ACADEMY

OF

SCIENCES, ARTS AND LETTERS

VOL. XXXVIII

NATURAE SPECIES RATIOOUE

MADISON, WISCONSIN

1946

The publication date of Volume 37 (1945) was April 10, 1947.

The publication date of Volume 38 (1946) is December 30, 1947.

PUBLISHED BY THE ACADEMY

OFFICERS OF THE WISCONSIN ACADEMY OF SCIENCES,

ARTS AND LETTERS

President

L. E. Noland, University of Wisconsin

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In Science: E. L. Bolender, Superior

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In Letters R. K. Richardson, Beloit

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The President

The Vice-Presidents

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E. A. Birge, past president

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Committee on Publications

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on the Council of the American Association

>r the Advancement of Science

Banner Bill Morgan

Representative

TABLE OF CONTENTS

Morphology of the Alimentary System of the Snail Page

Lymnaea Stagnalis Appressa Say. Melbourne Romaine Carriker 1

Maple Sugar: A Bibliography of Early Records. II.

H. A. Schuette and A. J. Ihde _ _ _ 89

f

Acidity of Soil and Water used in Cranberry Culture.

Neil E. Stevens _ _ ... _ 185

Preliminary Reports on the Flora of Wisconsin. XXXIII.

Ranunculaceae. Norman C. Fassett _ _ 189

A Cytological Study of the Development of the Oospore of Sclerospora

Macrospora (Sacc.). E. S. McDonough _ 211

Notes on Wisconsin Parasitic Fungi. VIII. H. C. Greene _ _ _ 219

Notes on Wisconsin Parasitic Fungi. IX. H. C. Greene _ 235

The Wisconsin Species of Peltigera. John W. Thompson, Jr. _ 249

Host-Parasite Relationships and Geographical Distribution of the

Physalopterinae (Nematoda). Banner Bill Morgan _ 273

Highest Abandoned Beach Ridges in Northern Door County,

Wisconsin. O. L. Kowalke _ _ _ 293

Artificial Hybrids between Muskellunge and Northern Pike.

John D. Black and Lyman 0. Williamson _ _ _ 299

Fox Hybrids. Leon J. Cole and Richard M. Shackelford _ 315

Copper in Lake Muds from Lakes of the Madison Area.

M. Starr Nichols, Theresa Henkel, and Dorothy McNall _ 333

Proceedings of the Academy _ _ _ 351

Constitution and By-Laws of the Academy _ _ 357

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Doe, J. H. 1934. The ecology of Wisconsin. Trans. Wisconsin Acad. Sci.

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120 State Historical Building, Madison, Wisconsin.

MORPHOLOGY OF THE ALIMENTARY SYSTEM OF THE

SNAIL LYMNAEA STAGNALIS APPRESSA SAY

Melbourne Romaine Carriker

Zoological Laboratory of the University of Wisconsin

Introduction

Knowledge concerning the details of the anatomy and his¬

tology of snails is becoming increasingly important as informa¬

tion accumulates regarding their role as vectors for parasites of

man, domestic animals, wild game and fish. Although the host-

parasite relations between the vertebrates and the helminths are

frequently well known, the corresponding relations between the

snail and the developmental stage of the parasite are less ade¬

quately understood. The pathological effect of the parasite on

the invertebrate host is frequently difficult to ascertain because

the normal microanatomy and histology of the organs of the mol¬

lusc are but poorly worked out.

In the present study an attempt has been made to assemble

and integrate the scattered and often incomplete information

available on the microanatomy of the alimentary system of the

snail Lymnaea stagnalis (Suborder Basommatophora, Order Pul-

monata), and by additional original investigation on L. s . ap-

pressa to reconstruct the form of this system in detail. Emphasis

has been placed on the proper selection of anatomical terms, and

where these have occurred in synonymy the synonyms have been

carefully considered in the selection of an acceptable term. The

introduction of some new terms has been necessary. The litera¬

ture presents but few quantitative descriptions ; for this reason

in this paper average measurements are given of living organs

wherever feasible. Emphasis is also placed on describing the

organs in their natural spatial relations within the shell and soft

parts. Studies already completed on the histology and physiology

of the alimentary system of this snail will appear in other

journals.

■JUKI !§f§

1

2 Wisconsin Academy of Sciences , Arts and Letters

Acknowledgments

This work was carried out largely at the University of Wis¬

consin (1939 to 1943) under the guidance of Prof. L. E. Noland

whose assistance is herewith gratefully acknowledged. Thanks

are due also to Prof. T. C. Nelson of Rutgers University under

whose direction the writer worked during the summer of 1942

at Rutgers University and the New Jersey Oyster Research

Laboratories.

Classification

According to the classifications of F. C. Baker (1902, 1911,

1928) there is question as to whether the subspecies of the snail

here investigated is appressa or jugularis. However, Dr. Henry

A. Pilsbry of the Academy of Natural Sciences has kindly clari¬

fied the matter of synonymy (by letter), explaining that the

original identifications of jugularis by Say are doubtful and that

such a dubious name is better discarded.

The following characters (of laboratory-cultured specimens

38 mm. or over in shell length) place this snail nearer appressa

taxonomically than to any other variety. As reported elsewhere

(Carriker, 1943b) a study of fifteen radulae disclosed the follow¬

ing limits of the number of teeth : laterals, 14-19 ; intermediates,

2-5 ; marginals, 21-30. Contrary to Baker's findings for this sub¬

species, the first laterals were found to be almost entirely tri¬

cuspid. Examination of 75 shells indicated that the spire length

varies from very much shorter than, to equal to, to longer than

the aperture length. The number of complete shell whorls varies

from 6.5 to 8, the average number being 7. The maximum length

of snails grown in the laboratory to date is 62.5 mm.

Historical Review

Of the references cited here only seven are the results of

research on American Lymnaeidae: Wetherby (1879), Whitfield

(1882), F. C. Baker (1900, 1911, 1928), Colton (1908) and Faust

(1920). These investigations, with the exception of that by

Faust which treats of the histology of the liver, describe only

the superficial gross anatomy of the alimentary system. The

more exhaustive researches are almost entirely those of Euro¬

pean investigators on European Lymnaeidae. Thus the Ameri¬

can lymnaeids are still an open field for investigation.

Carriker — Alimentary System of the Snail

3

The following are the more thorough morphological works

performed on the various parts of the alimentary system of

Lymnaea: on the raduia: Rossler (1885), Schnabel (1903), and

Hoffmann (1932); the radular cartilage: Dybowski (1885),

Loisel (1893), Schafer (1913) and Baecker (1932) ; radular col-

lostyle: Baecker (1932); salivary glands: Prenant (1923);

stomach region: Heidermanns (1924); liver: Barfurth (1880,

1883), Frenzel (1886), Enriques (1902) and Faust (1920).

Thus the buccal mass, the esophagus and intestine have received

little attention. With the exception of some preliminary obser¬

vations on the buccal musculature by Amaudrut (1898) and the

thorough work of Heidermanns on the innervation and vasculari¬

zation of the stomach region, the gross and detailed musculature,

innervation and vascularization of the system also have been

only superficially investigated.

The following reviews were very helpful on the morphology

of the Lymnaeidae: Moquin-Tandon (1885), Griffiths (1892),

Cooke (1913), Pelseneer (1906), F. C. Baker (1900, 1928), Sim-

roth and Hoffmann (1908-1928), Pelseneer (1935) and Yonge

(1936).

The research on the alimentary system of the remaining

Basommatophora is represented by only a few papers. The more

detailed contributions are those of Hurst (1927) on Physa;

H. B, Baker (1925), Lcrnx; Schumann (1911) Gadinia ; Hackel

(1911), Chilina ; and Hoff (1940) , Ferrissia. Older minor papers

are those of Andre (1893) on Ancylus; Buchner (1891) on

Planorbis; and Hutton (1879) on Amphibola.

The outstanding investigations on the Pulmonata have been

performed principally on the stylommatophoran Helix. The more

recent of such works which have a bearing on the pulmonate ali¬

mentary system are mentioned here : on the raduia : Spek (1921) ,

Pruvot-Fol (1926) ; alimentary canal: V. Haffner (1923) ; mor¬

phology of musculature: Trappmann (1916) ; morphology of

vascular system: Schmidt (1916) ; morphology of nervous sys¬

tem: Schmalz (1914); body membranes: Kisker (1923). In

other stylommatophorans the alimentary canal is also treated at

some length by Richter (1926) in Cerion; Hoffmann (1925),

Vaginulidae ; Argaud and Bounoure (1910), Arion; and Awati

and Karandikar (1940), Oncidium.

4 Wisconsin Academy of Sciences, Arts and Letters

MATERIALS AND METHODS

L. s. appressa was selected for the study in part because of

its excellent response to laboratory culture. Additional advan¬

tages are found in the relatively large size of the snail as com¬

pared with other fresh-water pulmonates, its short life-cycle and

its relatively thin, semitransparent shell.

Over 1,000 snails used in the investigation were raised

entirely in the laboratory. The original snails were collected in

Fox Lake, Wisconsin, in 1939. As some of these snails were

probably parasitized, the following method was utilized to obtain

parasite-free animals : snail egg masses, soon after oviposition,

were isolated into separate aquaria. Each new culture was

started in this way, hence transmission of infection was rendered

very improbable. Repeated dissections of snails of succeeding

generations have never revealed the presence of parasites.

The soft parts of the animal were most easily removed un¬

injured from the shell by breaking the whorls away from the

columella by means of slight outward-twisting pressure exerted

on curved forceps. The origin of the heavy columellar muscle

thus exposed is readily dislodged with the points of the forceps,

and the soft parts of the mollusc then may be gently twisted out

of the remaining shell spire.

Relaxation of the soft parts through anesthetization was

produced most satisfactorily by the use of nembutal. The variety

used was the commercial veterinary product, pentobarbital so¬

dium, Abbott ; list number 8612 ; concentration : 1 grain per cc.

dissolved in 10% alcohol. Injections were made with a 1-cc. hy¬

podermic syringe. The site of injection causing the minimum

contraction and distributing the hypnotic most rapidly through¬

out the body of the snail is the heart. A minute hole was picked

on the left side of the body whorl over the lung. If the snail is

held before a strong beam of light the outline of the organs

within the shell can be distinguished. The point of the hypo¬

dermic needle was directed through the renal organ and into the

pericardium. The injection was most effective when made di¬

rectly into the auricle or ventricle, both of which are clearly

seen dilating. The quantity of nembutal per unit of body weight

required to produce hypnosis is over fifty times that used in

human administration. The average net weight of adult L. s.

appressa is 5 gms. (maximum wet weight found to date: 8.39

Carriker— Alimentary System of the Snail 5

gms.) and the approximate dose for complete hypnosis is about

0.1 cc. Quantities below 0.03 cc. had little visible effect. Within

the limits of 0.03 and 0.20 cc. the promptness with which

hypnosis was obtained increased with increase in dosage. Doses

of 0.5 cc. per snail were used for the relaxation of snails for

morphological study. With a 0.1-cc. dosage of nembutal about an

hour was usually required for hypnosis of the more easily anes¬

thetized snails, and as long as two hours for the more refractory

specimens. Smaller snails required proportionately less nem¬

butal. At high temperatures (e.g., 30-35° C.) hypnosis occurred

more rapidly.

Magnesium sulfate was tried but with less success. Urethane

(ethyl carbamate), water at 60° C. and glacial acetic acid gave

only moderately i^elaxed animals. Chloroform, ether and similar

anesthetics produced violent contracture of the snail, even when

the anesthetic was floated in watch glasses on the water over the

snails.

Snails anesthetized with nembutal and injected with 2M mag¬

nesium sulfate did not give animals as well relaxed as with nem¬

butal alone, but the animals did loose the irritability which is

generally noted about the tentacles and the velum of snails anes¬

thetized with nembutal alone. Cardiac administration of 0.2 cc.

of nembutal and immersion in a small volume of water with a

few crystals of chloretone produced a more deeply anesthetized

snail in two to three hours than did the nembutal alone. Out of

25 snails thus treated only one had contracted into the shell.

Excellent relaxation of the snails was obtained by asphyxiation

in boiled water to which chloretone had been added, but it took

two to three days to completely asphyxiate them. Concurrent

asphyxiation of animals injected with nembutal is not advisable,

as under the anaerobic conditions of the method, the tissues

about the site of injection autolyze badly.

General dissection was carried out in both living and pre¬

served snails. In the former, the animals were anesthetized with

nembutal for about an hour and then dissected under a special

Lymnaea physiological solution developed to approximate more

closely the ionic concentrations of Lymnaea blood. An account of

this solution will appear in a separate paper on the physiology

of the alimentary tract of L. s . appressa. The above procedure

left the organs very nearly in their natural form. Crystals of

6 Wisconsin Academy of Sciences , Arts and Letters

chloretone added to the dissecting medium kept the animals

under anesthesia. For dissection of preserved specimens, snails

were killed by asphyxiation in boiled water to which chloretone

had been added, and after killing were preserved in a mixture

of dilute alcohol and formalin. Neutral red and methylene blue

in varying dilute concentrations were used to differentiate the

tissues during the process of dissection.

Study of the shape of the lumen of the alimentary canal was

facilitated by injecting the canal with pigmented latex or pig¬

mented vinylite. The canals of some 50 snails were cleared of

food material by five days of starvation. They were then relaxed

by the nembutal-chioretone method described above. Turtox latex

injection medium, diluted with tap water to a creamy consist¬

ency, was injected in a rubber-metal syringe. Similarly Synllex

vinylite, diluted with acetone, was injected in a dry glass syringe.

As water immediately sets the vinylite, excess water was blotted

from tissues and objects touched by the syringe. The dorsal

region of the heads of the anesthetized snails was opened longi¬

tudinally to expose the buccal mass ; this made possible the pas¬

sage of the needle through the mouth into the esophagus without

puncturing these delicate structures. The buccal mass was

pressed tightly about the needle with a pair of fine forceps dur¬

ing the actual injection. Pressure was exerted on the plunger of

the syringe until the injection medium poured out the anus. Not

infrequently injections were ruined by the rupturing of the

delicate pyloric region of incompletely anesthetized snails. Speci¬

mens injected with latex were allowed to set in dilute acetic acid.

With vinylite, water was used. The tissues of the vinylite-

injected specimens were dissolved away from the injectant by

the use of concentrated KOH or strong HN03. Some total speci¬

mens were cleared by the method described by Guyer (1936),

p. 117. The method is satisfactory if air bubbles are removed

by suction during the clearing.

For a study of the topography of the inner walls of the lumina

of the alimentary canal, starved, anesthetized snails were in¬

jected orally with a deep blue solution of methylene blue. This

technic distended the walls of the canal and accentuated the pat¬

tern of the inner walls. For a study of the relationship of the

alimentary lumina to the alimentary tract the carmine-staining

method advised by Lathrop (1937) was applied. The presence of

Carriker— -Alimentary System of the Snail

7

chitin in the alimentary system was detected by the method of

Campbell (1929). Portions of the tract were removed from nar¬

cotized snails and tested separately.

Methods for the preparation of the radulae for study have

been described elsewhere (Carriker, 1943b).

By far the most satisfactory injection medium for the vas¬

cular system was Emery's aqueous carmine (Lee, 1921), supple¬

mented by the use of approximately 0.5% of the vasodilator

sodium nitrite (NaN02). Snails were relaxed by the cardiac in¬

jection of 0.3 cc. of nembutal or by immersion in chloretone solu¬

tion. After about an hour the snails were sufficiently relaxed to

take the injectant. A 1-cc. syringe was filled with the carmine

solution and laid aside with the needle in damp cloth ; a shallow

support which would hold the needle at the level of the heart

during injection was also made ready. The shell of the relaxed

snail was removed as far back as the liver, exposing the heart

and the aorta, care being taken that the columellar muscle was

not freed from the remaining shell and that no portions of the

snail body were ruptured. The pericardium was cut open expos¬

ing the heart. With fine forceps a short piece of fine thread was

slipped under the ventricle and a loose overhand knot was tied

about it ; the syringe needle was then pressed into the ventricle

and down the aorta a short distance. The thread was tied tightly

about the ventricle and needle. Injectant was then slowly forced

into the vascular system until the minute arteries of the ventral

surface of the foot started showing red. The syringe was then

withdrawn and the ventricle tied off tightly. The snail, though

under the effect of the hypnotic, is stimulated chemically and

probably mechanically by the injectant, and moves slightly, thus

furthering the penetration of the injectant into the most minute

arterioles. The snail was then placed in approximately 5% alco¬

hol until it ceased movement, and then into 20-30% alcohol for

hardening of the carmine. Stronger alcohol hardens the carmine

more effectively, but so also the tissues, which is not desirable in

dissection. The injections were carried out entirely under the

binocular microscope with the snail lying on paper toweling and

under a strong beam of light. It was necessary to moisten the

animal repeatedly during injection. Even though anesthetiza¬

tion with chloretone was more time-consuming (approximately

six hours), it was found more expedient, as there is always

8 Wisconsin Academy of Sciences , Arts and Letters

danger of puncturing the ventricle or aorta with the syringe in

cardiac injection. Once the animal is punctured, it is of no fur¬

ther use for total vascular injection. In some instances well-

injected snails were dissected to expose the arteries, then dehy¬

drated in alcohol and cleared in xylol. Injection of the vascular

system was also made with undiluted India ink, with strong con¬

centrations of methylene blue, and with pigmented vinylite, but

these methods were not as effective as the first method outlined

above.

The topography of the gross nervous system was studied in

(1) narcotized snails dissected under Lymnaea physiological solu¬

tion, (2) relaxed snails immersed for several hours in 5% nitric

acid, (3) in snails killed by asphyxiation in chloretone and pre¬

served in dilute alcohol and formalin, (4) and in snails treated

as in (3) but cleared in glycerine and dissected under water after

removal of the excess glycerine. In all of these methods meth¬

ylene blue was sometimes applied during dissection to further

differentiate the tissues. Minute innervation of the organs of the

alimentary tract was studied by the technic of Alexandrowicz

(1932). Submerging of the tissues in the dye was decidedly

more fruitful than injection of the dye into the living animal.

Small pieces of tissue from anesthetized snails were spread flat

on thin pieces of paraffin (approximately 2 mm. thick) and

secured there by means of very small insect pins (“Minuten

nadeln”). These preparations were then immersed in solutions

of Lymnaea physiological solution to which the methylene blue

had been added (2 drops of 0.5% methylene blue and 1 cc. of

translucent paraffin plates, were placed under the compound

microscope from time to time to note the extent of staining. The

nerves of the salivary glands stained well in about six hours ; the

remaining parts of the tract took as long as 24 hours. Ganglionic

cells of the alimentary system were followed in histological sec¬

tions and by means of the method suggested by Heidermanns

(1924).

GENERAL ORGANIZATION OF INTERNAL ORGANS

All measurements given in this paper will be those normal

to 48 mm. snails, the average of adult laboratory animals : shell

length, 48 mm. ; greatest width, 23 mm. ; length of aperture, 25

mm.; and width of aperture, 15 mm. F. C. Baker (1911, 1928)

Carriker— Alimentary System of the Snail

9

describes the shell and the external appearance of the soft parts

of L. s, appressa in detail. Throughout the paper, right and left

refer to the animal's right and left.

The general organization of the internal organs is illustrated

in Figs. 1, 2, 3 and 4. The body cavity of the snail is divided

into three major spaces: the cephalic and the visceral hemocoels

and the lung. The cephalic hemocoel lies within the head and

neck regions (fig. 2). The visceral hemocoel is placed dorso-

posteriad the cephalic hemocoel and ventrad the lung and lies

within the shell (fig. 1). The lung saddles the middorsal region

of the animal, lying over the anterior third of the visceral hemo¬

coel and in the posterior dorsal limits of the seventh body whorl.

That portion of the diaphragm constituting the anteroventral

boundary of the lung is considerably depressed and makes con¬

nection with the ventral body wall by means of a diffuse fibro-

cellular septum. This separates the cephalic from the visceral

hemocoel. The ducts of the reproductive system, nerves and

arteries which pass between the two hemocoels are firmly bound

in the septal connective tissue ; the esophagus is permitted free

egress through a small rounded passageway remaining in the

left ventral portion. Lacaze-Duthiers (1872) first referred to

the relationship of these “cervical” structures in L. stagnalis

but did not define them very clearly. Moquin-Tandon (1885) in

Gastropoda, Robson (1922) in Paludestrina, Brown (1933-34)

in Philine, and Fretter (1939) in Tectibranchia make passing

reference but do not describe similar septa. Kisker (1923) in a

description of a similar structure in Helix suggests the name

“Membrana transversa” but does not consider the esophageal

opening. It is desirable to designate the interhemocoelic septum

as the cervical septum and the esophageal conduit as the eso¬

phageal iter.

The buccal mass occupies the anterior third of the cephalic

hemocoel (fig. 2) and is bounded on all but the posterior side by

the strong muscular walls of the head. A delicate laminated

membrane screens the buccal mass from the remainder of the

hemocoel. This membrane passes from the central nervous sys¬

tem forward completely around the buccal mass, attaching cir¬

cumferentially in the body wall a short distance behind the

mouth. This lamination is generously perforated and appears to

function in part as a baffle membrane in connection with the cir-

10 Wisconsin Academy of Sciences , Arts and Letters

. ctors at maud ib /<?

ouccoi moss

oetbr groove

Foot

hermaphroditic duct

posterior /obe of the Huer

poster/or ostg/e of bodg cohort

'osphrodtum

'gizzard

a er> trorisc erot orterg

Carriker — Alimentary System of the Snail

11

PLATE I

Explanation of Figure

Figure 1. Perspective line draw¬

ing of an entire L. s. appressa re¬

moved from the shell and opened

along the left side. The dorsum has

been flattened to the right and the

organs spread for the sake of clar¬

ity. The stomach region and the

posterior lobe of the liver have been

turned on the longitudinal axis about

one half turn to the right (compare

with fig. 3). The midventral portion

of the diaphragm and the tunica pro¬

pria have been omitted. In the na¬

tural position the reproductive or¬

gans are closely grouped over the

postesophagus, the lung covers the

reproductive glands and some of the

organs in the visceral hemocoel, the

heart makes connection on the left

ventral side with the aorta, the colu-

mellar muscle attaches to the colu¬

mella of the shell. The length of the

postintestine has been shortened. The

configuration and nomenclature of

the reproductive glands illustrated

here are taken from Holm (1946).

X 2.

Abbreviations

aur.— auricle

cen. ner. sys.-— central nervous sys¬

tem

col. mus. — columellar muscle

cr. spt. — cervical septum

diap. — diaphragm

g. p. 9 —female genital pore

g. p. S — male genital pore

pe. — penial nerve

per. — pericardium

pneu. — pneumatopore

pre. ret. m. — retractor muscle of the

preputium

ren. o. — renal organ

ren. or. — renal orifice

salivary gl. — salivary gland

t. pro. — tunica propria

ven.—ventricle

12 Wisconsin Academy of Sciences , Arts and Letters

oe/ar groove

monHs

Carriker— Alimentary System of the Snail

13

PLATE II

Explanation of Figure

Figure 2. Dorsal aspect of the na¬

tural position of the contents of the

cephalic hemocoel. A median sagittal

incision has been cut through the

dorsum from the dorsal mandible to

the mantle. Thus the velum has been

bisected and the two resulting por¬

tions spread laterad. X 5.

Abbreviations

1 — dorsolateral protractor muscle

2 — preventral protractor muscle

4 — dorsolateral retractor muscle

6-— preventral levator muscle

ao — aortic nerve

cap m. — capitocerebral membrane

cerebral g. — cerebral ganglion

g. p. $ — male genital pore

g. p. $ — female genital pore

Is — superior labial nerve

li — inferior labial nerve

parietal g. — parietal ganglion

pla— anterior pallial nerve

pll — left pallial nerve

plm — median pallial nerve

plr — right pallial nerve

plv — ventropallial nerve

pneu. — pneumatopore

pre. prot. — protractor muscle of the

preputium

pre. ret. — retractor muscle of the

preputium

S — salivary artery

sp — splanchnic nerve

tt — tentacular nerve

14 Wisconsin Academy of Sciences , Arts and Letters

bucco / mass

reao/ organ

'recta/ s/nus

‘ectam

recta/ ar/er/es

'Pepat/c c/acts

post/n test/no/ Qrterg

major Pepat/c Qrterg

post//? test we

■ Pepat/c Qrterg

poster/or /ode of/foer

4

Carriker — Alimentary System of the Snail

15

PLATE III

Explanation of Figures

Figure 3. Outline of the organs of

the alimentary tract as seen in their

natural spatial relationship through

the shell. X 1.

Figure 4. Detailed illustration of

the alimentary system. The stomach

region and the posterior lobe of the

liver have been turned leftward one

half turn so that the ventral sur¬

faces of these parts are illustrated;

the dorsal surfaces of the remainder

of the tract are shown. The intestine

has been much shortened. Vasculari¬

zation of the posterior lobe of the

liver is drawn; only the openings of

the major hepatic ducts are indicat¬

ed. The portion of the body wall

about the rectum and anus has been

retained. The portion of the liver

which has been drawn has been un¬

coiled. The dorsal junction of the

salivary glands is severed and the

glands are spread laterad. X 5.

5 — buccal retractor muscle

5r — right buccal retractor muscle

51 — left buccal retractor muscle

29 — salivary retractor muscle

A — aorta

a. h.d. — hepatic ducts, anterior

of the liver

bl — laterobuccal nerve

b. m.-— buccal mass

bv — ventrobuccal nerve

b.w. — body whorl of shell

C— cephalic artery

cec. — cecum

cr. — crop

gr — right gastric nerve

GV — ventrogastric artery

giz. — gizzard

gz. — gizzard

H — hepatic artery

li — inferior labial nerve

Abbreviations

Is — superior labial nerve

m.i. — midintestine

pe — penial nerve

PE — penial artery

pli — internal pallial nerve

plr — right pallial nerve

pneu. — pneumatopore

pr.e. — proesophagus

pr.i. — prointestine

pt.e. — postesophagus

pt.i. — postintestine

pyl. — pylorus

re.— -rectum

s — salivary nerve

S— salivary artery

sp. — spire

tt — tentacular nerve

VD— dorsovisceral artery

VV — ventrovisceral artery

lobe

16 Wisconsin Academy of Sciences, Arts and Letters

culation of the body fluids. Many of the large nerves passing to

the velum and oral structures extend peripherally in the mem¬

brane. The term used for an analogous sheet in Helix by Kisker

(1923), capitocerebral membrane, is adopted here for this

membrane in Lymnaea .

The bulky preputium reaches posteriad from behind the base

of the right tentacle, and crowding the esophagus and related

organs to the left, occupies the middle third of the hemocoel.

The large prostate gland lies in the right posterior two thirds

of the cavity, and is pressed against the diaphragm and cervical

septum. The ducts of the reproductive glands curve to the left

behind the prostate gland and slip under the diaphragm through

the connective tissue of the cervical septum into the visceral

hemocoel beyond. The esophagus proceeds unhampered from the

one hemocoel to the other by way of the esophageal iter, and lies

to the left and partly under the left portion of the prostate gland.

The banded longitudinal muscles of the floor of the cephalic

hemocoel run posteriad and dorsally into the shell clustering

under the right side of the diaphragm as the columellar muscle

(fig. 1). This passes slightly beyond and under the posterior

boundary of the lung to its origin midway up the columella of

the shell. The visceral coils within the shell are clothed by a thin

transparent laminated muscular tunic, the tunica propria, which

makes connection with the remainder of the body wall at the

mantle collar.

The bulk of the reproductive glands are clumped in the ante¬

rior portion of the visceral hemocoel, somewhat to the left and

partly over the columellar muscle. In the sexually mature animal

they bulge prominently into the lung cavity under certain por¬

tions of the diaphragm.

Immediately behind the esophageal iter the esophagus runs

parallel with and to the left of the columellar muscle, depressed

between the oval spermatheca to the left and the large oo thecal

gland to the right. Past these glands it drops gradually below

the columellar muscle to join the crop, gizzard and pylorus ven-

trally. These names are those applied by English writers to the

three regions of the middle section of the tract. The gizzard and

pylorus are probably comparable to what some authors refer to

as the “stomach.” These digestive organs lie cupped in the

midventral concavity outlined by the sixth and seventh shell

Carriker — Alimentary System of the Snail 17

whorls, at a point dorsad the angle of the shell aperture and

partly under the columellar muscle. The albumen gland in adult

snails noticeably crowds the stomach organs to the left ; most of

the other reproductive glands lie above and anteriad the stomach

organs.

Shell whorls 1 to 5 and the greater part of 6 are filled by the

massive dark brown liver which is located „ behind the stomach

organs. The posterior half of the pylorus curves to the right and

passes a short distance alongside the gizzard (fig. 3). It is fol¬

lowed by three coils of the intestine for which the terms pro-,

mid- and postintestinal loops are offered. The prointestinal loop

completely encircles the crop, gizzard and pylorus. The intestine

then moves across the ventral surface of the viscera close behind

the pylorus towards the right side of the sixth body whorl and

mounts the external right contour of the dorsal surface of the

liver. Here it bends back on itself in a graceful U curve, forming

the midintestinal loop. This lies still in the sixth shell whorl.

From this loop the intestine descends over the right contour of

the liver, parallel to and but slightly posteriad the ascending

portion. The descending portion is confined to the angle of the

shell whorl. It turns under the liver again ventrally, still run¬

ning alongside and posterior to the earlier lengths of the intes¬

tine. At the junction of the sixth and seventh shell whorls it

passes immediately behind the pylorus and the prointestinal loop.

Its course persists in the angle of the body whorl forming one

last turn completely around the body whorl, the postintestinal

loop. As the intestine passes behind the lung it gradually en¬

larges to form the rectum which terminates in the anus . The

ovotestis lies embedded in the central spiral course of the liver,

generally bordering on the columella of the shell.

The lung is an empty chamber unornamented by accessory

respiratory structures (fig. 1). Midventrally the diaphragm is

very thin and transparent, becoming more strongly muscled as it

joins the cervical septum anteriad. The bright yellow-orange

renal organ spreads thinly across the roof of the respiratory

chamber from the pneumatopore on the right side to the peri¬

cardium on the extreme left ventral side of the body whorl. The

pericardium is located over the parietal wall of the shell slightly

behind the umbilical chink and encloses a single bright-yellow

ventricle and light-yellow auricle.

18 Wisconsin Academy of Sciences, Arts and Letters

GENERAL CONFIGURATION OF THE ALIMENTARY

SYSTEM

Buccal mass (figs. 5-12) . This is a pyriform muscular organ

whose muscles are colored a bright reddish-chestnut allegedly

due to the presence of hemoglobin (Lankester 1872, Griffiths

1892, Bowell 1924, Hoffmann in Simroth and Hoffmann 1908-

1928, Pelseneer 1985). It is 6.6 mm. long; 5.2 mm. through its

greatest dorsoventral dimensions at the posterior end; 8.5 mm.

through a dorsoventral plane directly behind the mandibles ; 2.9

mm. wide at the dorsal mandible ; and 4.9 mm. wide at the great¬

est width posteriorly.

In its simplest outline the buccal mass consists of two sup¬

porting structures, a large central odontophoral cartilage and a

smaller cylindrical posterior radular collostyle (fig. 6).

The odontophoral cartilage is an ovoid, spoon-shaped,

vesiculo-muscular organ whose longer axis in the resting position

runs dorsoventrad in the buccal mass. The sides curve back to

such an extent that from the front it appears semicircular in

cross section. When spread flat it measures 4.64 mm. in its

greatest length, 4.06 mm. at the greatest width basally and 2.90

mm. at the dorsal width where it narrows to a truncated lip

under the radular membrane. In thickness it varies from 1.16

mm. through the thickest lateral pillar-like portions to 0.70 mm.

in the middorsal central portion, and 0.29 mm. in the thinner

midventral central part. Thus a concavity is shaped in the poste¬

rior central dorsoventral plane of the cartilage which in part

supports the radular sac (figs. 6, 8, 9, 12). The lateral swellings

of the cartilage taper at each end, particularly in the dorsal posi¬

tion under the radula where the ends are approximately of the

same thickness as the midventral portion. The odontophoral

cartilage is not cartilagenous. Because analogous structures in

some of the marine snails are composed of tissue very similar to

mammalian cartilage and because of the continued application

of this term for such radular supportive structures by malacolo-

gists, the term is now generally accepted for the Gastropoda.

The number of so-called cartilages in different Gastropoda varies

from a single one to many: in Urosalpinx cinerea (Carriker,

1948a) there exist two separate and distinct cartilages; in Helix

Carriker — Alimentary System of the Snail 19

pomatia (Trappmann, 1916) the single cartilage appears com¬

posed of two lateral parts which are held intimately together by

the horizontal cartilage muscle; in L. s. appressa the cartilage

exists as a single structure with a shallow groove running dorso-

ventrally in the rear midventral portion and is pigmented a

faint reddish-chestnut color.

Nestled in the posterior hollow of the cartilage lies the

radular sac. This is strengthened centrally by a cylinder of

translucent milky-white supportive tissue, the radular collostyle ,

which is firm and of a gelatinous nature. The term collostyle is

introduced here to replace the inaccurate name “papilla” which

with many other terms has been applied to this structure. The

collostyle is topped dorsally by a cap of epithelium and slightly

pigmented muscle fibers. The term collostylar hood is suggested

for the cap. The nascent portion of the radula passes around and

envelopes most of the cylindrical collostyle, and is clothed ex¬

ternally by the radular sac. The latter measures 3.78 mm. from

the ventral end to the collostylar hood. Ventrally it is 1.22 mm.

in diameter; halfway between the ends, 0.98 mm.; and at the

hood, 1.57 mm. (figs. 6, 9, 11).

The cartilage, radular sac and attendant muscles bulge as the

odontophore under the buccal cavity epithelium in a gracefully

rounded protuberance and amply fill most of the buccal cavity

(fig. 7). The cartilage and muscles of the odontophore function

synchronously as a cushion upon which the radula is carried,

longitudinally directed. The buccal cavity epithelium is affixed

to the subepithelial tissues on all but the ventral surfaces of the

cavity. Here it rises freely over the cartilage and is connected

with the odontophoral musculature principally at the sides and

behind the odontophore. Consequently the cartilage is permitted

considerable play under the cavity epithelium. Behind the crest

of the odontophore the epithelium makes a deep tubular evagina-

tion which passes down into the odontophore between the car¬

tilage and the radular collostyle to the ventral extremity of the

collostyle. This tubular evagination is collapsed on itself and

forms a double-layered trough which has come to fit closely and

almost completely around the collostyle. It thus forms a two-

layered cellular envelope around all but a narrow longitudinal

band on the posterior surface of the collostyle (figs. 6, 9-11).

20 Wisconsin Academy of Sciences, Arts and Letters

Carriker — Alimentary System of the Snail

21

PLATE IV

Explanation of Figures

Figure 5. Right lateral face of

the buccal mass. Emphasis is placed

on the external musculature, inner¬

vation and external vascularization.

X 10.

Figure 6. Internal left half of the

buccal mass which has been cut in a

longitudinal sagittal plane. The ar¬

rangement of the intrinsic buccal

muscles about the odontophoral car¬

tilage and the radular sac is illus¬

trated. The radula and radular sac

have been left entire. X 10.

Abbreviations :

Muscles :

1 — dorsolateral protractor

2 — preventral protractor

3 — postventral protractor

4 — dorsolateral retractor

5 — buccal retractor

6 — preventral levator

7 — postventral levator

8 — suboral dilator

9 — dorsomandibular dilator

10 — suspensor of the radular sac

13 — anterior jugalis

14— posterior jugalis

15 — buccal sphincter

16 — mandibular approximator

17— dorsal odontophoral flexor

18 — ventral odontophoral protractor

19 — infraventral odontophoral pro¬

tractor

20 — supramedian radular tensor

21 — supralateral radular tensor

22 — inframedian radular tensor

23— infralateral cartilage tensor

27 — superior suspensor of the radu¬

lar sac

28 — tensor of the hood

Others :

arb.b. — buccal vascular arborescence

b.c. — buccal cavity

bcc— buccocerebral connective

BD— dorsobuccal artery

BT — postbuccal artery

bd — dorsobuccal nerve

bl — laterobuccal nerve

br— buccal retractor nerve

bv — ventrobuccal nerve

bt — postbuccal nerve

G- — cephalic artery

cart. — cartilage

c. h.— collostylar hood

d. f.c. — dorsal food channel

d.l.r. — dorsolateral ridge

d.m. — dorsal mandible

EP — pr oesophageal artery

es.gl. — buccal glands of proesopha¬

gus

es.l. — ledge at entrance to proesoph¬

agus

gb— buccal ganglion

gr — right gastric nerve

jp — posterior jugalis nerve

l.m.— lateral mandible

P — pedal artery

pr.es. — proesophagus

rad. — radula

rd.s. — radular sac

s— salivary nerve

s.d.o. — salivary duct opening

sal. d. — salivary gland duct

sal.g. — salivary gland

sr.e. — subradular epithelium

VI— inner buccal valve

22 Wisconsin Academy* of Sciences , Arts and Letters

8

Carriker — Alimentary System of the Snail

28

PLATE V

Explanation

Figure 7. Front view of the odon-

tophore, surrounded by muscles of

the buccal mass and portions of the

buccal cavity epithelium. X 10.

Figure 8. Musculature of the left

internal half of the buccal mass. The

of Figures

radular sac, distal portions of the

odontophoral cartilage muscles, por¬

tions of the odontophoral cartilage

and the epithelium of the buccal

cavity have been removed. X 10.

Abbreviations: Iff miMIIWliM'

Muscles :

2 — preventral protractor

3 — postventral protractor

5— buccal retractor

6 — preventral levator

7 — postventral levator

15 — buccal sphincter

16 — mandibular approximator

17 — dorsal odontophoral flexor

18 — ventral odontophoral protractor

19— infraventral odontophoral pro¬

tractor

20 — supramedian radular tensor

21 — supralateral radular tensor

22 — inframedian radular tensor

23— infralateral cartilage tensor

Others :

b.c.ep. — buccal cavity epithelium

b.c. — buccal cavity

cart. — cartilage

d.m. — dorsal mandible

esoph. — esophagus

l.m. — lateral mandible

rad. ch. — chitinous membrane of the

radula

subrad.ep. — subradular epithelium

Carriker— Alimentary System of the Snail 25

PLATE VI

Explanation of Figures

Figure 9. Posterior lateral view of

the radular sac lying in its normal

position against the posterior de¬

pression of the odontophoral cartil¬

age. The left half of the odontophore

and parts of the radula and the

radular sac have been removed to

expose the sinuses existing between

these structures. X 12.

Figure 10. Posterior view of the

radular sac and radula in the nor¬

mal relation to the musculature of

the odontophore. The radula has been

lifted off of the anterior face of the

odontophoral cushion and drawn in

the same plane as that of the radu¬

lar sac. The subradular epithelium

has been cut away from the radular

sac and the heavy supralateral radu¬

lar tensor muscles. X 12.

Figure 11. Posterior aspect of the

buccal mass. A segment of the right

posterior half of the buccal cavity

epithelium has been cut out ; the lam¬

inated posterior jugalis muscle over

the posterior portion of the buccal

mass has been dissected to the sides;

and the right posterior portion of

the radular sac has been opened to

expose the collostyle. X 10.

Figure 12. Posterior view of the

odontophoral cartilage and some of

its attendant muscles. X 10.

Abbreviations :

Muscles :

10 — suspensor of the radular sac

14 — posterior jugalis

17 — dorsal odontophoral flexor

20 — supramedian radular tensor

21 — supralateral radular tensor

22 — inframedian radular tensor

25 — lateral suspensor of the radular

sac

26 — inferior suspensor of the radu¬

lar sac

27— superior suspensor of the radu¬

lar sac

28— tensor of the collostylar hood

Others :

b.c.ep.— buccal cavity epithelium

bt — postbuccal nerve

btr — right postbuccal nerve

buc.gl. — buccal glands

C— cephalic artery

cart. — cartilage

col. — collostyle

col.h.— collostylar hood

esoph. — esophagus

rad.m. — radular membrane

subrad.ep. — subradular epithelium

s.r.ep.— subradular epithelium

26 Wisconsin Academy of Sciences, Arts and Letters

Of these epithelia Hoffmann (1932) and many previous workers

name the membrane under the radula the basal epithelium and

that over the radula the covering epithelium. The terms sub -

radular and supraradular epithelia , respectively, as used by some

workers, are thought here to give a clearer picture of the rela¬

tionship of the parts. A final, external, thin connective tissue

sheath forms a casing over the combined collostyle and epithelial

layers.

The radula is produced by and between these two evaginated

epithelial layers. As the radula passes out of the epithelial sac

it unfurls and spreads anteriad over the buccal cavity epithelium,

or subradular epithelium as it is better named. The radula thus

roofs the crest of the odontophore, and in the resting position

reaches a short distance over the anterior side of the odonto-

phoral cushion. Throughout, the radula is affixed quite firmly to

the subradular epithelium by means of the subradular chitin

which is secreted by the epithelium. At its exit from the radular

sac, the supraradular epithelium, before passing back into the

buccal cavity epithelium, elevates and arches over the radular

collostyle as the hood. This rests tightly against the unfurling

radula (figs. 6, 9, 10). The entire radula is a chitinous ribbon

approximately 6.0 mm. long by 2.5 mm. wide, surfaced by a uni¬

form distribution of some 130 transverse and 100 longitudinal

rows of glass-like posteriorly directed denticles. The posterior

half lies securely fastened about the radular collostyle (fig. 9).

The epithelial covering of the mouth is lined dorsally by the

heavy, hardened dorsal mandible, and at the sides by the two

depressed lateral mandibles. The dorsal mandible is 1.57 mm.

wide and 0.64 mm. high; and the laterals, 0.18 mm. wide and

0.87 mm. long. Ventrally the aperture is walled by deep longi¬

tudinal folds which permit expansion of the mouth to the diam¬

eter of the odontophore. The inner proximal margin of the

dorsal mandible is projected its full width as a relatively thick

layer some distance back under the roofing epithelium of the

buccal cavity. A narrower extension of the layer continues far

back directly over the path of the radula some 3.2 mm. and ter¬

minates at a point over the crest of the resting odontophore. The

extension paves the anterior two thirds of a wide deep depres¬

sion, the dorsal food channel , which is 0.75 mm. wide and 5.2

mm. long and passes from the dorsal mandible to the tapering

Carriker — Alimentary System of the Snail 27

of the dorsal part of the cavity into the esophagus (fig. 6). The

dorsal portions of the sides of the buccal cavity bulge inwardly

in the form of a shallow flattened elevation on each side, which

may be called the dorsolateral ridges . The two salivary gland

ducts make their entrance to the buccal cavity in the dorsolateral

region, at a point in the posterior origin of the dorsolateral

ridges (fig. 6). At the ventral boundary of the longitudinally

folded opening of the esophagus there is present also a promi¬

nence or esophageal ledge which extends a short distance ante-

riad over the hood. This is usually deeply impressed by two or

three longitudinal furrows which permit considerable expansion.

The living epithelium of the buccal cavity and its protective

covering, with the exception of the radula and the mandibles,

is very soft and of a velvety consistency. The mandibles, the

entire radula and the cuticular lining of the anterior two thirds

of the cavity gave a marked positive test for chitin. The radular

denticles when stained with orange G after the treatment with

the alkali appeared identical to normal untreated teeth, indicat¬

ing the presence of a very high concentration of chitin.

The greater part of the external walls of the buccal mass

consists of muscle, connective tissue and buccal glands which lie

between the muscle layers and the epithelium, and are distributed

over most of the buccal mass and anterior portions of the

esophagus. Over the rear dorsal portions of the mass the glands

present a rough surface raised unevenly in irregularly distrib¬

uted pustules (figs. 5, 11).

Esophagus (figs. 1-6, 8, 11, 15). On the basis of form, histol¬

ogy and function, the esophagus is distinctly divisible into an

anterior and posterior esophagus. The anterior esophagus, for

which is suggested the term proesophagus, passes out of the buc¬

cal mass as a smooth, straight, thin-walled, light cream-yellow

tube approximately 1.16 mm. in diameter. It courses posteriad

some 4.7 mm. from the buccal mass and suddenly widens into the

posterior esophagus under the salivary glands. The posterior

esophagus, here designated the postesophagus, retains its in¬

creased diameter of 1.35 mm. the full 16.5 mm. of its length. It

is capable of distension to 5.0 mm. Its coloration varies from a

translucent light cream to a gray-brown, furrowed by darker

longitudinal stripes.

Internally the esophageal walls present a heavily corrugated

surface (fig. 15). The folds run longitudinally. In the proesoph-

28

Wisconsin Academy of Sciences , Arts and Letters

Carriker — Alimentary System of the Snail

29

PLATE YII

Explanation of Figures

Figure 13. Detailed view of the

internal contours of the cecum, he¬

patic duct openings into the pylorus,

atrium and some of the pyloric folds

and passages. The cecum has been

bisected in the longitudinal axis of

the alimentary tract, thus in the

drawing only the right cecal fold is

evident. The pattern of the atrial

and hepatic vestibular corrugations

is accented. X 6.

Figure 14. Detailed drawing of the

anterior portion of the prointestine,

showing the pellet forming area. The

prointestine has been cut down the

middorsal line and spread. X 5.

Figure 15. Detailed illustration of

the dorsal internal surfaces of the

esophagus, crop, gizzard, pylorus and

a portion of the prointestine. A me¬

dian longitudinal incision has been

made to permit opening of the tract.

X 5.

Abbreviations :

cec.f. — cecal fold

d.pyl.p. — dorsal pyloric passage

exc.c.t. — excurrent cecal tubule

hep.cor. — hepatic corrugations

hep.d. — hepatic duct

hepatic duct, a — hepatic duct of an¬

terior lobe of liver

hepatic duct,p — hepatic duct of pos¬

terior lobe of liver

hep.vest. — hepatic vestibule

inc.c.t. — incurrent cecal tubule

m. pyl.f. — major pyloric fold

n. pyl.f. — minor pyloric fold

proc.p. — procurrent pyloric passage

retr.p. — retrocurrent pyloric passage

ventral f. — ventral fold

30

Wisconsin Academy of Sciences , Arts

and Letters

py /or us '

Carriker — Alimentary System of the Snail

31

PLATE VIII

Explanation of Figures

Figure 16. Detailed drawing of the

ductal system of the liver. The pos¬

terior lobe of the liver has been

partly uncoiled. Those hepatic ducts

lying in the first plane of the draw¬

ing and the cecum and the pylorus

have been opened by an incision in

the longitudinal axis of the struc¬

tures. X 5.

Figure 17. Detailed drawing of the

external dorsal aspect of the crop,

gizzard, pylorus, cecum and proin¬

testinal loop. The details of inner¬

vation and vascularization are ac¬

cented. X 6.

Abbreviations :

33 — gizzard compressor muscle

34 — gizzard tensor

A — aorta

C — cephalic artery

CC — cecal artery

CP — compressor artery

ET — postesophageal artery

G — genital artery

GD — dorsogastric artery

gl — -left gastric nerve

gr — right gastric nerve

h — hepatic nerve

HP — prohepatic artery

i — ingluvial nerve

ID — dorsoingluvial artery

ip — prointestinal nerve

IP — prointestinal artery

pd — dorsogastric plexus

PM — major pyloric artery

PN — minor pyloric artery

R — rectal artery

VD — dorsovisceral artery

VT — vestibular vascular arboresence

VV — ventro visceral artery

ZD — dorsal gizzard artery

32 Wisconsin Academy of Sciences, Arts and Letters

agus four such folds are present : two lie in the dorsolateral and

two in the ventrolateral portions of the tube. The two dorso¬

lateral folds pass into the buccal cavity as the dorsolateral buccal

cavity ridges, and the two ventral lateral folds fade into the

posteroventral buccal wall. At the initial flaring of the post¬

esophagus the four folds of the proesophagus branch to give

approximately fifteen smaller folds. These run back through the

crop as far as the gizzard. The postesophagus is capable of great

distension in which state the folds level out considerably ; the pro¬

esophagus does not appear capable of such marked enlargement.

Salivary glands (figs. 1, 2, 4). The pair of salivary glands

are conspicuous in the cephalic hemocoel in their brilliant yellow

color, palmatifid form and very finely and uniformly textured

surface. When spread flat each gland measures 6.9 mm, in length

and 5.2 mm. in width. In their normal relationship to the ante¬

rior portion of the posteosphagus, each gland spreads posteriad

over one lateral half of the esophagus in finger-like lobes. These

adhere firmly to the esophageal walls by means of short strands

of muscle and the vascular ramifications. The two glands are

connected over and under the esophagus by bridge-like exten¬

sions of the marginal gland lobes, and thus completely encircle

the esophagus.

The two salivary gland ducts are straight, smooth-walled,

light-yellow tubes 7.0 mm. in length and 0.35 mm. in diameter.

They pass, one to each side of the esophagus, from the rear dorsal

surface of the buccal mass through the aperture in the central

nervous system in company with the esophagus. Each duct

branches many times in each gland ; the diameter of each of the

first branch ducts is 0.20 mm.

Yonge (1936) has suggested that the salivary glands in the

Gastropoda be designated topographically, those emptying into

the buccal cavity being called buccal glands. The fact that such

a change in terms would confuse the true buccal glands in the

walls of the buccal mass (see Hackel 1911, and later in the pres¬

ent paper) with the salivary glands, and the additional fact that

the present term of “salivary” has been so long in use, indicate

that no advantage is to be gained from such a change.

Crop (figs. 1, 4, 15, 17). The crop is indistinctly separated

from the postesophagus and is distinguished from it only by its

increased radial dimensions. It is a thin-wailed bulbular pocket

Carriker— Alimentary System of the Snail

33

which connects the postesophagus with the gizzard and measures

2.3 mm. in length, 3.5 mm. in width and 4.4 mm. in depth. Inter¬

nally the ventral portion is depressed in a rounded, shallow

hollow below the floor of the gizzard and the walls bear continua¬

tions of the longitudinal folds from the postesophagus. There is

no pronounced constriction of the postesophagus as it enters the

crop, but the diameter at this junction usually remains small

(fig. 15), Posteriorly the crop joins the gizzard with only a

slight narrowing of its lumen, a constriction caused by the pull¬

ing in of the crop walls to meet the inner margins of the gizzard

lobes. Externally and internally a sharp boundary is evident be¬

tween the shallowly corrugated internal surface of the crop and

the smooth, slightly elevated cuticulated surface of the gizzard

pads. The external coloration of the crop varies in different indi¬

viduals from ivory yellow to slate gray, accented with more

intense stippled markings ; a velvet-gray lining covers the inter¬

nal surfaces of the crop as well as those of the esophagus.

Hoffmann (Simroth and Hoffmann, 1908-1928) was uncer¬

tain as to whether the crop should be considered a part of the

stomach or a true esophageal crop. Structurally the crop of L. s.

appressa appears more closely allied to the postesophagus.

Gizzard (figs. 1, 3, 4, 15, 17). This is a crushing organ con¬

sisting essentially of two very strong, laterally placed muscular

cushions, the gizzard lobes, which are joined dorsally and ven-

trally by tough bands of silvery luster, the gizzard tendons. Both

muscular lobes are colored a strikingly deep reddish-chestnut

and in some individuals the surface bears added black stippled

designs. The dimensions of the organ are as follows: length,

4.9 mm. ; width, 6.5 mm. ; and depth, 5.3 mm. Each lobe is globu¬

lar and is rounded and narrowed medially on the dorsal and ven¬

tral sides to about one third of its length, where the two are

strongly connected by the non-contractile gizzard tendons. The

left lobe lies slightly behind the right.

Internally the lobes are represented by two ovoid, distinctly

concave, slightly projected, flattened pads which face each other.

These lie parallel aiid almost meet when the gizzard lumen is

’ void of food and/or sand particles. The pads are firm and hard

and protected by a relatively thick layer of cuticle which, by

reason of the deeply pigmented musculature beneath, appears

red. This cuticle when tested for chitin reacted negatively to all

34 Wisconsin Academy of Sciences, Arts and Letters

of the tests! Nor was chitin found in any other part of the

digestive system with the exception of the buccal mass.

Possibly the first to describe this organ, but only superficially,

were Cuvier (1806, 1817), and Stiebal (1815). More recent rec¬

ords are those of Goodsir (1840), Lacaze-Duthiers (1872), Gar-

tenauer (1875), Wetherby (1879), Whitfield (1882), Moquin-

Tandon (1885), F. C. Baker (1900, 1911), Colton (1908) and

Heidermanns (1924). In the illustrations of some of these in¬

vestigators the stomach parts are shown upside down (Lacaze-

Duthiers 1872, Wetherby 1879, F. C. Baker 1900). According to

F. C. Baker, Whitfield was the first to correctly describe the

stomach region in American Lymnaea. Heidermanns believes

that the gizzard tendons are to be compared in their functions

to the tendons of vertebrate muscle.

Pylorus (figs. 1, 4, 15-17). The morphology of the basom-

matophoran pylorus has been neglected, except for the work of

Heidermanns (1924), which was principally on cellular physi¬

ology. He mentioned briefly, but did not name, the three longi¬

tudinal folds which partially block the opening into the liver and

which swell to partly occlude the opening into the gizzard lumen.

In the following paragraphs an attempt is made to describe and

name those structures most significantly correlated with the

function of the pylorus.

Immediately behind the gizzard the alimentary canal con¬

tinues as a tapering tube, the pylorus, which curves to the right

around the right gizzard lobe. The two hepatic ducts and the

cecum empty into the rear middorsal part and make the site of

greatest constriction (fig. 15). At the junction of the pylorus

and the gizzard, the pyloric walls narrow slightly, accommo¬

dating the posterior opening of the gizzard. Directly behind the

opening of the cecum into the pylorus and existing as a modified

prolongation of the pylorus, the digestive canal widens into a

flaring rounded chamber whose dorsal and lateral walls appear

palmately ribbed, the main stem passing from the proximal pos¬

terior base of the cecum. The term atrium first applied to a

similar structure in Ferrissia by Hoff (1940) is adopted here

for this structure in L. s. appressa, but because of its function

it is more rightfully considered as a part of the pylorus than

of the intestine. The efferent ends of the two hepatic ducts

approach each other from opposite sides and converge broadly

Carriker— Alimentary System of the Snail

35

over the rear dorsum of the pylorus. Here a common ante¬

chamber, for which the term hepatic vestibule is offered, some

0.46 mm. across, is interposed. The cecum passes out of its poste¬

rior portion and the vestibule itself makes connection with the

pyloric lumen (figs. 15, 17). The dimensions of the pylorus. vary

considerably with the extent of muscular dilation and with the

content of nutriment. A relaxed, moderately filled, straightened

pylorus measures about 2.3 mm. in its greatest diameter just

behind the gizzard; 1.5 mm. in its least diameter immediately in

front of the hepatic vestibule; 4.9 mm. from the gizzard to the

hepatic vestibule; and 7.5 mm. from the gizzard to the atrium.

The atrium is 2.5 mm. across and 2.4 mm. in length from its

posterior margin to the posterior base of the cecum. The pylorus

is usually translucent and colored a straw yellow.

Two principal channels pass down the length of the pylorus,

for which the terms dorsal and ventral pyloric passages are pro¬

posed (figs. 13, 15, 16). The dorsal passage leads along the

dorsal third of the pylorus from the gizzard to the hepatic

vestibule. It is a smooth, deep, narrow groove making connection

between the gizzard lumen and the liver by way of the hepatic

ducts and the hepatic vestibule. The ventral pyloric passage runs

in the ventral two thirds of the pylorus. It is the more spacious

channel and constitutes the connection between the gizzard

lumen and the intestine. The two passages are almost completely

separated by two ridges of tissue, the major and the minor py¬

loric folds. These are located in the dorsolateral walls of the

pylorus and meet along their distal crests. The major fold ex¬

tends along the right dorsolateral wall from the right gizzard

pad to the atrium, passing under and to the right of the hepatic

vestibule. It increases somewhat in bulk as it advances, and ter¬

minates in a flattened protruding bulge partly under the opening

of the hepatic vestibule. The minor fold reaches posteriad along

the left dorsolateral portion of the pylorus parallel to the major

fold. It is a flattened, less conspicuous, fold which becomes

reduced and disappears slightly to the rear of the left wall of the

hepatic vestibule. The major and minor folds thus may effect

complete closure of the hepatic vestibule from the ventral pyloric

passage. Passing down over the midventral wall of the ventral

passage is found the low, wide, ventral pyloric fold. This extends

from the ventral junction of the gizzard pads to the left ventral

36 Wisconsin Academy of Sciences , Arts and Letters

margin of the atrium and the excurrent cecal ridges located

there. It is thickened into a prominent muscular elevation di¬

rectly behind the gizzard pads and partly occludes the ventral

passage from the gizzard cavity. The fold gradually diminishes

in bulk posteriad. In the living animal the folds are relatively

turgid and smooth; in the dissected specimen they become

wrinkled and convoluted ; and in histological sections they shrink

markedly.

The ventral pyloric passage is thus partly divided into two

smaller passages by the ventral pyloric fold. Because of the di¬

rection of the ciliary currents in these two conduits, the terms

procurrent and retrocurrent passages are suggested for them.

The former lies to the right and the latter to the left of the

ventral fold. The procurrent passage is narrower than the retro-

current and becomes smaller further down the pylorus, finally

passing out into the intestine ventrally beneath the atrium. The

retrocurrent passage has no direct outlet to the intestine, and its

walls are capable of marked distension. The three pyloric folds

are faintly pigmented a light reddish-chestnut color, as was first

observed by Heidermanns (1924). No references can be found

on the investigation of the hemoglobin content of these folds or

of the gizzard lobes.

The atrium flares widely to the rear of the hepatic vestibule

and the cecum, as a flattened inflation in the dorsal and lateral

portions of the rear of the pylorus. The procurrent pyloric pas¬

sage after passing down the right lateral side of the pylorus,

swings gradually to the midventral side and empties into the

intestine across the vacancy left by the discontinuation of the

atrium in the ventral pyloric wall. The connection between the

atrium and the anterior portion of the pylorus may be completely

occluded by constriction of the pylorus at this point and the

consequent approximation of the major and minor pyloric folds.

The efferent ends of the hepatic ducts are lined internally by

a corrugated epithelium. The furrows so formed commence in

the hepatic ducts as minute longitudinally directed grooves which

coalesce progressively into large gutters in their course toward

the hepatic vestibule. In the hepatic vestibule there are finally

present only some twelve such grooves and ridges, where their

longitudinal axes are pointed in the direction of the cecal open¬

ing (fig. 13). Two or three furrows of the same proportions as

Carriker— Alimentary System of the Snail 37

those In the vestibule arise In the passage leading from the vesti¬

bule to the atrium and run into the latter, in a direction at an

obtuse angle from that of the grooves in the vestibule. The few

larger grooves branch into an extensive system of progressively

smaller depressions which flare out fan-like over the inner walls

of the atrium. This ramiform pattern originates at the opening

of the cecum into the vestibule and produces the palmate design

visible externally.

Cecum (figs. 4, 13, 15-17). For some time this organ was

entirely overlooked by morphologists and was considered to be

absent in the Lymnaeidae. By Wetherby (1879) and F. C. Baker

(1900) it was referred to as the “pancreas.” Heidermanns

(1924) calls attention to the cecal folds but does not describe in

any detail the structure of the organ.

The cecum in L. s. appressa arises as a dorsal evagination of

the hepatic vestibule and extends horizontally anteriad over and

between the right lobe of the gizzard and the early part of the

prointestinal loop. It is a cylindrical, slightly tapering, blind tube

4.4 mm. long, with a proximal diameter of 1.2 mm. and a distal

diameter of 0.6 mm. Numerous branching blood vessels secure

it to the intestine and gizzard and it is well hidden by the ante¬

rior lobes of the liver. This fact, combined with the fact that the

cecum is an inconspicuous, watery, translucent, straw-yellow

structure, readily explains why early workers entirely missed it.

The cecal lumen is divided longitudinally into two channels by

the cecal folds . By reason of the function of these two conduits,

the terms incurrent and excurrent cecal tubules are offered for

them. The cecal folds exist as two flattened ledges of tissue

which extend from the posterior borders of the hepatic vestibule

along the lateral walls of the cecum. The projected folds meet

along their distal extremities and pass almost to the distal tip

of the cecum where they flatten out. Thus they leave there a

small opening which connects the distal portions of the incur¬

rent and excurrent tubules. The incurrent tubule originates in

the posterodorsal portion of the hepatic vestibule. One of the

minute ridges in the epithelium of the vestibule passes about one

third of the distance into the incurrent tubule midway between

the two cecal folds ; and two others accompany it to either side

about half this distance. The remainder of the tubule is smooth-

walled. The excurrent tubule is entirely smooth-walled and is

38 Wisconsin Academy of Sciences, Arts and Letters

more spacious than the counterpart. At the base of the cecum

the excurrent tubule narrows slightly, turns posteriad and emp¬

ties by way of a furrow running across the atrium into the pro¬

intestine, a groove outlined by the cecal folds which at the same

time thicken basally (fig. 15).

Liver (figs. 1, 4, 16). This exists as two distinct, racemose,

compound-tubular outpocketings of the pylorus, the anterior and

posterior liver lobes, whose terminal ducts end blindly in minute

clustered follicles. The whole branching complex is supported by

a meshwork of connective tissue, blood vessels and nerves. Ex¬

ternally it is enveloped in the thin muscular tunica propria.

Slight tenacious strands of connective tissue and muscle and

some of the peripheral blood vessels bind the tunica to the liver.

The coloration of the liver varies from a delicate salmon pink in

partly starved animals to a greenish-brown in animals feeding

heavily on lettuce. According to Sorby (1876) the reddish pig¬

mentation of the liver of Lymnaea is caused by a secreted fluid

composed in part of hematin. A white, irregularly distributed

mottling is evident over the surface of the liver, as well as over

all the other organs of the body. This is caused by the presence

in the connective tissue of white calciferous concretions similar

to those found in Helix by Kisker (1923) and by v. Haffner

(1923).

In newly matured snails the liver is roughly one third the

size of the soft parts of the animal. In old snails, however, it

atrophies markedly. The anterior (morphologically the right

one) lobe is roundly triangular and measures about 4.5 mm. by

5.0 mm. on its peripheral surface and is 3.5 mm. deep. It lies

ventrally, closely bounded by the contours of the gizzard and the

pylorus. The posterior (morphologically the left) lobe consti¬

tutes the greater part of the first six whorls of the shell and

when coiled within the shell measure 25.0 mm. in length and

12.0 mm. in its greatest diameter anteriorly. When removed

from the shell and uncoiled, the lobe is fully 37 mm. long. Both

lobes are thickly penetrated by a multibranching ramification

of blind hepatic ducts. The anterior lobe possesses only one such

many-branching conduit; the posterior lobe is filled by several.

The spire is reached principally by one large median duct which

spirals in the gland across the winding contour of the columella ;

a second large tube passes spireward along the periphery of the

Carriker — Alimentary System of the Snail 39

whorls; two smaller duets push anteriad into the forward pro¬

jections. These ducts coalesce in the middle anterior portion of

the lobe, behind the flexure of the pylorus, forming a large,

short vessel which in turn discharges into the left end of the

hepatic vestibule. A similar smaller duct from the anterior lobe

enters the vestibule from the right end. The hepatic ducts, with

the exception of the short corrugated portions emptying into the

hepatic vestibule, are smooth, thin-walled vessels. The openings

of the larger ones are about 1 mm. in diameter — but slightly

smaller than that of the intestine. The liver, when considered in

terms of internal surface area of its ramiform tubular complex,

exposes a relatively vast surface of epithelium which in com¬

parison dwarfs the inner surface dimensions of the remainder

of the alimentary canal.

Little attention has been given to the macroscopic morphol¬

ogy of the liver. The work of Pelseneer (1906) and Faust

(1920) may be mentioned, but their descriptions are cursory.

The liver of the gostropods has been known by so many different

appellations that the present use of the term “liver” deserves

explanation. In the gastropod literature of the past century such

English terms as liver, digestive gland, digestive diverticulum,

hepatic gland, hepatopancreas, gastric gland, and liver mass are

extant. German names encountered are “Leber” and “Mittel-

darmdriise” ; French designations, “foie”, etc. Liver is most gen¬

erally accepted. It is of course obvious that the gastropod liver

and the vertebrate liver are not homologous either structurally

or functionally, but there is similarity in general position and

function, and the term liver has been used so universally for

this organ in the gastropods that acceptance of the designation

is justified.

Intestine and rectum (figs. 1, 3, 4, 14, 15, 17). The intestine

and rectum together measure 70 mm. in length. The entire ali¬

mentary canal is 115 mm. long. If the total length of the evagina-

tions of the canal (salivaries and ducts, liver and ducts and

cecum) were considered, the total length would exceed 170 mm.

Thus the intestine and rectum together constitute about 41% of

the total length of all the parts of the alimentary system. The

intestine varies roughly from 1.3 to 1.5 mm. in diameter. The

rectum is some 15 mm. long and 1.5 to 2.0 mm. in diameter.

Both the intestine and rectum are very thin-walled and easily

40 Wisconsin Academy of Sciences , Arts and Letters

ruptured in dissection. The former passes imperceptibly into

the latter. The rectum, after passing to the right behind the

lung, reaches the anus by means of a tubular passage, the rectal

sinus, made for it through the heavy musculature of the posterior

angle of the mantle.

No mention of the specialization of the prointestine in the

Basommatophora were encountered in the literature examined.

The anterior half of the prointestinal loop may be referred to

as the pellet-molding region which has developed two structures

for this function: The first is the intestinal typhlosole and the

second may be named the pellet-compressor . The typhlosole

arises as an infolding of the ventral muscular wall of the intes¬

tine and projects at right angles about 0.6 mm. into the intestinal

lumen. It is 0.4 mm. wide proximally and projects gradually

from the intestinal wall some 2.4 mm. behind the atrium. It is

3.4 mm. long, is placed along the longitudinal axis of the pro-

intestine and slowly reaches its maximum size at the posterior

extremity (figs. 14, 15). This end stops abruptly as a rounded

prominence. The pellet-compressor consists essentially of a spe¬

cialized circular band of the intestine, 3.4 mm. wide. It lies adja¬

cent to and at the anally directed end of the typhlosole and con¬

sists of a slightly thickened wall of circular muscle which is lined

internally by approximately 40 distinctly outlined, thick epi¬

thelial annulations. These are conspicuous in their yellowish

brown coloration and circular uniformity. They are not always

parallel, branching and coalescing at intervals (fig. 14). The

compressor generally lies in that portion of the prointestinal

loop passing over the postesophagus.

The typhlosole persists as a slight ridge of muscle beneath

the annulations of the pellet-compressor. It is not evident there

unless the epithelial layer is removed. Posterior to the pellet-

compressor the ridge is visible again and can be followed a short

distance down the intestine as a low, narrow elevation covered

by a raised band of epithelium which runs down along the same

side of the intestine. The term intestinal raphe is offered for

this raised band of epithelium. The inner lining of the intestine

and rectum, beyond the pellet-compressor, is ribbed by bands

which pass obliquely around the intestinal circumference from

the raphe (fig. 14). The bands are low, narrow, flattened eleva¬

tions of the epithelium, similar in width and height to the raphe,

Carriker — -Alimentary System of the Snail 41

and meet opposite the raphe, tending to form a continuous longi¬

tudinal band down the intestine and raphe. This second band

opposite the raphe may be referred to as the pseudoraphe. The

term intestinal costae is suggested for the obliquely running

bands.

H. B. Baker (1925) describes “plicae” in the intestinal epi¬

thelium of Lanx. The intestinal costae of L. s. appressa could

hardly be called plicae as they are not true folds. Hackel (1911)

noted that the entire intestine of Chilina is covered within by epi¬

thelial “folds,” which according to his description, are dis¬

tributed similarly to those in its near relative, L. s. appressa,

MUSCULATURE OF THE ALIMENTARY SYSTEM

Buccal mass (figs. 5-12, 27). The musculature of this spe¬

cialized organ is the most complicated in the entire animal, and

indeed, one of the most intricate muscular structures in the in¬

vertebrate group. Loisel (1893) gave an early, though rather

confused, description of some of the muscles of L. stagyialis.

Amaudrut (1898) offered the first really comprehensive study.

His work was concerned mostly with Helix , with but occasional

references to Lymnaea , and has formed the basis for some of the

terminology of the more recent investigators. He states that the

buccal retractor muscles are missing altogether in Lymnaea.

They are always present in L. s. appressa, however, and F. C.

Baker (1900, 1911, 1928) also found these muscles in all the

Lymnaeidae investigated. As Hoffmann (Simroth and Hoff¬

mann, 1908-1928) states, little accurate work has as yet been

done on the basommatophoran buccal musculature. The best

work yet published on the musculature of the pulmonate buccal

mass is that by Trappmann (1916) on Helix pomatia. Other

good gastropod papers in this connection are those of Herrick

(1906) on Sycotypus, Richter (1926) on Cerion, and Crofts

(1929) on Haliotis. The terminology adopted for the muscles of

the buccal mass of L. s. appressa is based wherever homology

warrants it on the works cited above, principally on that of

Trappmann. The intrinsic musculature of L. s. appressa has

been found to be almost identical with that of Helix pomatia,

although Helix , possibly as a result of its terrestrial habitat,

possesses stronger, bulkier, more developed muscles. Amaudrut

42 Wisconsin Academy of Sciences , Arts and Letters

43

Carriker— Alimentary System of the Snail

PLATE IX

Explanation op Figures

Figure 18. Typical ostiole in the

surface tissue of the postesophagus.

X 200.

Figure 19. Ventral view of the

buccal mass showing the mode of at¬

tachment of the ventrobuccal artery

and extrinsic muscles, and the loca¬

tion of the arterial valves. The loca¬

tion of the inner buccal valve is in¬

dicated by the dark oval mark (VI)

and is seen only in dissection of the

mass. X 10.

Figure 20. Inner buccal valve.

X 12.

Figure 21. Anteroventral aspect

of the inferior suspensor muscle of

the radular sac. The outer, flaring,

hollow portion of the suspensor has

been opened medially. X 12.

Figure 22. Penetration of the right

gizzard lobe by the right dorsal giz¬

zard artery. The proximal portion

of the artery has been cut away.

X 10.

Figure 23. Capillary, showing its

terminations in the surface of the

postesophagus in ostioles. Before it

plunges into the tissue, the capillary

is thickly covered with vesicular

cells. X 100.

Figure 24. Central nervous system

and principal nerves of the buccal,

cerebral, parietal and abdominal

ganglia. X 4.

Abbreviations :

Muscles :

2 — preventral protractor

3 — postventral protractor

7— postventral levator

14 — posterior jugalis

19 — infraventral odontophoral pro¬

tractor

20- — supramedian radular tensor

25 — -lateral suspensor of the radular

sac

28— -inferior suspensor of the radu¬

lar sac

27 — superior suspensor of the radu¬

lar sac

Others :

ao — aortic nerve

art.— -arteriole

bd — dorsobuccal nerve

bcc— buccocerebral connective

bl — laterobuccal nerve

bt — postbuccal nerve

bv — ventrobuccal nerve

BV — ventrobuccal artery

c — columellar nerve

c.c. — calciferous concretion

cap. — -capillary

cr — -cardiac nerve

cs — splanchnic complex

f — frontal nerve

g — genital nerve

ga — abdominal ganglion

go — cerebral ganglion

go — osphradial ganglion

gpa — parietal ganglion

gpe — pedal ganglion

gpl— pleural ganglion

gr — right gastric nerve

in— intestinal nerve

Id — dorsolabial nerve

li — inferior labial nerve

Is— superior labial nerve

Iv — ventrolabial nerve

nu— nucleus of vesicular cell

o— gonadal nerve

pe — penial nerve

pla— anterior pallial nerve

pll — left pallial nerve

plm — median pallial nerve

pli— internal pallial nerve

plr — right pallial nerve

plv — ventropallial nerve

s— salivary nerve

sp — splanchnic nerve

to — optic nerve

tt — tentacular nerve

VB— outer buccal valve

v.c. — vesicular cell

vl-— velar nerve

VI— inner buccal valve

VO — odontophoral valve

44 Wisconsin Academy of Sciences, Arts and Letters

go/ c-—

py /or us

c/orsgyas/r/o

p/oxos

or ter/or /obe of

//ver

uer/ro gas tr/c

p/ezus .

Pepot/'c c/uc/s

/ oos/er/or /o6e of

//oep —

Carriker— Alimentary System of the Snail

45

PLATE X

Explanation of Figure

Figure 25. Innervation of the ali¬

mentary system. Portions of the

postesophagus and of the intestine

have been omitted. The stippled sur¬

faces indicate areas of relatively less

developed musculature. Arrows indi¬

cate the continuity of the nerves.

X 6.

Abbreviations :

Muscles :

5 — buccal retractor

13— -anterior jugalis

14 — posterior jugalis

15 — buccal sphincter

16 — mandibular approximator

17 — dorsal odontophoral flexor

18 — ventral odontophoral protractor

19 — infraventral odontophoral pro¬

tractor

20 — supramedian radular tensor

21 — supralateral radular tensor

23 — infralateral cartilage tensor

Others :

ano — anastomosis of dorsogastric

plexus and gonadal nerve

anp — anastomosis of gastric nerves

over pylorus

anz — anastomosis of gastric nerves

over gizzard

ao— aortic nerve

bcc — buccocerebral connective

bd — dorsobuccal nerve

bl — laterobuccal nerve

blr — branch of laterobuccal nerve

br — huccal retractor nerve

bt — pcstbuccal nerve

btr — right postbuccal nerve

buccal gls.— -buccal glands

bv — ventrobuccal nerve

c-— columellar nerve

cin — intestinal complex

ea — anterior esophageal nerve

ep— proesophageal nerve

f — frontal nerve

ga — abdominal ganglion

gal. c. — ganglionic nerve cell

gb — buccal ganglion

gl — left gastric nerve

gr — right gastric nerve

gpa — parietal ganglion

h — hepatic nerve

i — ingluvial nerve

in — intestinal nerve

ip — prointestinal nerve

jp — posterior jugalis nerve

id — dorsolabial nerve

li— inferior labial nerve

Is— superior labial nerve

lv — ventrolabial nerve

o — gonadal nerve

p — pyloric nerve

pla— anterior pallial nerve

pil — left pallial nerve

plm— median pallial nerve

plr — right pallial nerve

plv — ventropallial nerve

s — salivary nerve

sp — splanchnic nerve

t— typhlosolic nerve

vl — velar nerve

46 Wisconsin Academy of Sciences , Arts and Letters

observed a great uniformity throughout the Pulmonata as re¬

gards the buccal mass.

In the following compilation each muscle is given a number

which corresponds to similar numbers designating the muscles

in the illustrations. In addition to the number, there is listed for

each muscle: the name (in italics), the origin and insertion, and

a brief description including function where the name is not

self-explanatory.

Extrinsic muscles, from body wall to buccal mass:

1. Dorsolateral protractors (tig. 5). Origin : body wall at the

sides of the oral aperture. Insertion: external posterior dorso¬

lateral surfaces of buccal mass. Two wide, thin bands on either

side of the buccal mass.

2. Preventral protractors (tigs. 5, 6, 8, 19). Origin: anterior

ventral surface of the body wall immediately behind oral aper¬

ture. Insertion: anterior ventral external wall of buccal mass

directly about buccal valve of the ventrobuccal artery (see sec¬

tion on vascularization). A pair of thin bands, passing forward

somewhat obliquely.

3. Postventral protractors (tigs. 5, 6, 8, 19). Origin: ventro¬

lateral body wall just behind oral aperture. Insertion: postero-

ventral external wall of buccal mass in ventral surface of and to

either side of buccal artery. Usually two thin bands.

4. Dorsolateral retractors (tig. 5). Origin: lateral body wall

on level with rear of buccal mass. Insertion: anterior dorso¬

lateral external surface of buccal mass. A pair of narrow, thin

ribbons to either side of the mass.

5. Buccal retractors (tigs. 2, 4-6, 8). Origin: columellar

muscle in ventral wall of cephalic hemocoel just in front of cer¬

vical septum. Insertion : external lateral surfaces of buccal mass

over the cartilage. Two slender muscle bands passing between

the ganglia of the central nervous system. The left buccal re¬

tractor, because of the salivary glands (tigs. 2, 4), where the

salivary muscle detaches, passes along the right ventral side of

the esophagus. The right buccal retractor extends posteriorly

somewhat farther to the right of the esophagus and does not

branch.

6. Preventral levators (figs. 5, 6, 8). Origin: lateral body

wall opposite the middorsal portion of the buccal mass. Inser-

Carriker — Alimentary System of the Snail 47

tion: anterior ventral external surface of buccal mass. Flat¬

tened broad bands on each side of the buccal mass which elevate

the anterior portion of the buccal mass on the rasping stroke of

the radula.

7. Postventral levators (figs. 5, 6, 8, 19). Origin: usually

same as preventral levators. Insertion: posteroventral external

surface of buccal mass beneath the cartilage, between the odon-

tophoral and the buccal valves of the ventrobuccal artery. Usu¬

ally two flattened broad bands on each side of the ventrobuccal

artery. The pre- and postventral levators suspend the buccal

mass as in a double sling ; the latter in part raise the rear of the

mass on the forward stroke of the odontophore, thus helping

to bring the odontophore into the position for rasping.

8. Suboral dilators (figs. 5, 6). Origin: across the ventral

body wall, midway under the buccal mass. Insertion : across the

anterior ventral wall of the buccal mass, immediately behind its

attachment to the body wall. A number of fine bands which

depress the anterior wall of the buccal cavity during the protru¬

sion of the odontophore.

9. Dorsomandibular dilators (figs. 5, 6). Origin: across the

dorsal body walk Insertion : across the dorsal attachment of the

dorsal mandible. A large number of fine bands which function,

in conjunction with the suboral dilators, in opening the mouth,

and which draw the dorsal mandible upwards and backwards.

10. Suspensor of the radular sac (fig. 5). Origin: commis¬

sure of the buccal ganglia at base of esophagus. Insertion : ex¬

ternal distal end of radular sac. A stout muscle serving to sus¬

pend the distal end of the radular sac, and passing over the stout

postbuccal nerve.

11. Labial retractors . Origin: within body wall. Insertion:

lip, entirely around oral aperture. These exist as a large number

of small fibers within the body wall which radiate from the lip.

They draw the lip away from the mouth during protrusion and

early retraction of the odontophore.

12. Labial sphincter . A circular diffuse band of a large num¬

ber of fine fibers located in the body wall at that point where the

buccal mass attaches to it, and which entirely surround the oral

aperture, serving to close the mouth.

48 Wisconsin Academy of Sciences , Arts and Letters

Intrinsic muscles in buccal wall, from cartillage to mandible :

13. Anterior jugalis (tig. 5). Origin: median dorsal surface

of buccal mass and corners of dorsal and lateral jaws. Insertion:

ventrolateral ends of cartilage. The conspicuous external mus¬

cular yoke passing over the anterior two thirds of the top and

sides of the buccal mass from the base of the cartilage, and con¬

tinuous over the top of the mass. Functions in suspending, ele¬

vating, and protracting the odontophore.

14. Posterior jugalis (figs. 5, 6, 11). Origin: along attach¬

ment of dorsal mandible in the buccal mass and body wall. In¬

sertion: distal tip of radular sac and over the ventrolateral ex¬

ternal surfaces of cartilage. A bursiform, thin, muscular sheet

enveloping the rear of the buccal mass and extending forward

under the anterior jugalis and the buccal sphincter over the dor¬

sal wall of the mass. The envelope thickens to the sides of the

radular sac and as it passes forward over the sides of the esopha¬

gus. This muscle pulls the ventral end of the odontophore pos¬

tered, upward and forward, while others draw the dorsal end

anteriad, downward and forward thus shifting the central axis

of the odontophore from a dorso ventral resting direction to the

anteroposterior one which it must assume before passing out of

the mouth.

15. Buccal sphincter (figs. 6-8). A broad, thick band of

muscle encircling the buccal mass, and lying between the anterior

margins of the cartilage and the mouth. It is covered by the

anterior jugalis, and dorsally covers the posterior jugalis.

16. Mandibular approximator (figs. 6, 8). Insertion: lateral

corners of dorsal mandible and attachment of lateral mandibles.

A broad band of muscle lying directly in front of and adjacent

to the buccal sphincter, and passing between the mandibles by

way of the ventral wail of the buccal mass. In contraction it

depresses the dorsal mandible and approximates the lateral

mandibles about the receding radula.

17. Dorsal odontophoral flexor (figs. 6-8, 12). Origin: lat¬

eral corners of dorsal mandible and anterior dorsum of buccal

mass. Insertion: dorsal third of the lateral extremities of the

cartilage. A pair of inconspicuous, wide, flat bands which flare

dorsoanteriad on either side of the cartilage. The muscles lie

next under the anterior portion of the posterior jugalis and the

buccal sphincter, and partly adjacent the buccal cavity epithelial

Carriker— Alimentary System of the Snail

49

lining. They function, in conjunction with the posterior jugalis,

in orienting the longitudinal axis of the odontophore in an an¬

teroposterior direction ; in keeping the cartilage spread and the

radula tensed, and in drawing the odontophore forward to and

partly out of the mouth.

18. V entral odontophoral protractor (figs. 6-8). Origin: lat¬

eral mandibles and ventral lip. Insertion: across the anterior

ventral base of the cartilage. A wide, thin sheet, thicker later¬

ally, which spreads under the epithelial lining of the ventral wall

of the buccal cavity, and meets the dorsal odontophoral flexor

laterally. It serves to draw the odontophore forward.

19. Inf raventral odontophore protractors (figs. 6-8). Origin:

ventral floor of the mouth, and mandibular approximator. Inser¬

tion : ventral base of cartilage. A pair of flat bands constituting

the external wall of the buccal mass, and continuous with the

anterior jugalis laterally. The ventral portions of the circularly

directed mandibular approximator, buccal sphincter, and infra¬

lateral radular tensor (see below) lie sandwiched between the

infraventral and the ventral odontophoral protractors.

Intrinsic muscles, from cartilage to radula and radular sac :

20. Supramedian radular tensors (figs. 6, 8-12). Origin:

ventral outer extremities of the cartilage. Insertion: dorsal an¬

terior face of the radular sac. Are two strong, flattened bands

which fit snugly in the posterior depression of the cartilage

between the radular sac and the cartilage.

21. Supralateral radular tensors (figs. 6, 8-12). Origin:

outer ventral and lower lateral extremities of cartilage. Inser¬

tion: (1) middle posterolateral surface of the radular sac, (2)

the under lateral margins of the subradular epithelium along

those portions at the mouth of the radular sac where the radula

begins to curve inward, (8) the under surfaces of the buccal

cavity epithelium at areas over, to the sides, and behind the

dorsolateral portions of the cartilage. This pair is the bulkiest

in the odontophore. They lie on either side of the supramedian

radular tensors and may be seen externally at the rear of the

buccal mass through the posterior jugalis. Dorsoposteriorly, a

laminated outer layer of muscle fibers pass medially from the

supralateral radular tensors over the radular sac, and thus bind

50 Wisconsin Academy of Sciences , Arts and Letters

the supralateral muscles, the cartilage, and the radular sac

together (fig. 11).

22. Inframedian radular tensors (figs. 6-8). Origin: ante¬

rior ventrolateral base of cartilage. Insertion : under side of sub-

radular epithelium, a pair over each lateral third of the cartilage.

These reach over the curved anterior surface of the cartilage,

each lateral muscle band branching once before attaching to the

epithelium. The anteriormost muscles draw the buccal epithe¬

lium posteroventrad when the odontophore is protruded ; and the

other posterior pair are larger and function more in pulling the

epithelium over the cartilage, in antagonism to the supramedian

radular tensors. The supramedian, supralateral and inframedian

radular tensors hold the radula tightly over the surface of the

cartilage and draw the radula back and forth over it during

rasping.

23. Infralateral cartilage tensor (figs. 6-8). An arc-like,

semicircular muscle band passing over the curved ventral ante¬

rior surface of the cartilage and fastening in the lateral edges

of the cartilage. Anteroventrally it is set adjacent to the buccal

sphincter. It serves in spreading the cartilage, especially on the

retractor stroke of the odontophore, when the supramedian and

supralateral radular tensors, in exerting the necessary pull to

stretch the radula, tend to collapse inward the outer margins of

the cartilage.

24. Intracartilage tensors. A mat of fibers passing through¬

out the cartilage at right angles to the anterior and the posterior

surfaces of the cartilage. The fibers situated in the lateral pillar¬

like portions of the cartilage are arched outward slightly. The

cartilage spaces between the muscle fibers are filled with the

turgid vesicular cells (see later) which act in part as antagonists

to the fibers. When the straight fibers in the medial portion of

the cartilage contract the result is a flattening out, compressing

and consequent slight laterad expansion of this part; with the

contraction of the curved fibers in the lateral pillars the result is

a drawing mediad and a slight bulging of the lateral extremities

of the cartilage. This specialization is of the utmost importance

in rasping, where it is the dorsal truncated tip and the posterior

dorsal surfaces of the cartilage, under the effective rasping por¬

tion of the radular, which function in transmitting the forces

exerted by the muscles of the buccal mass to the radula.

Carriker — Alimentary System of the Snail 51

Intrinsic muscles, passing outward from the radular sac :

25. Lateral suspensors of the radular sac (figs. 10, 11, 21).

Origin: median posterior surface of the supralateral radular

tensors. Insertion: distal base of the radular sac. Two short,

thin bands horizontally directed.

26. Inferior suspensor of the radular sac (figs. 10, 12, 21).

Origin: posteroventral median surface of the cartilage in the

cartilage groove. Insertion: distal base of the radular sac. A

short, laminated sheet which flares from its origin to its inser¬

tion, and whose slender anterior portion lies directly in the

opening of the odontophoral valve of the buccal artery.

27. Superior suspensor of the radular sac (fig. 11). Origin:

posterior ventral boundary of the buccal cavity epithelium, down

behind the odontophore. Insertion: distal base of the radular

sac. Thin, flattened band, vertically directed. The lateral, infe¬

rior and superior suspensors of the radular sac hold the distal

end of the radular sac securely suspended in the posterior ven¬

tral portion of the buccal mass, between the bulky supralateral

radular tensors.

28. Tensor of the hood (figs. 6, 10, 11). Origin: posterior

ventral surface of radular sac. Insertion: under surface of the

collostylar hood, A small hour-glass-shaped, muscle serving in

the holding of the collostylar hood during the rasping stroke

and, indirectly, also probably in preventing the forward passage

of the hood with the radula as it grows out of the radular sac.

Homologies for some of the muscles described above have not

been described in the pulmonate literature. The extrinsic mus¬

cles, with the exception of the suboral and the dorsomandibular

dilators and suspensor of the radular sac, are but slight varia¬

tions of similar muscles described for other pulmonates. The

suboral and the dorsomandibular dilators (Nos. 8, 9, 10), how¬

ever, are not mentioned before. Trappmann gives the name of

“anterior protractor muscles” to muscles in the same region in

Helix , but there is no such function for these muscles in L. s.

appressa. The suspensor of the radular sac has also gone un¬

described. Of the intrinsic muscles, the labial retractors and

labial sphincter (Nos. 11, 12), the mandibular approximator

(No. 16), the ventral odontophoral and the infraventral odonto¬

phoral protractors (Nos. 18, 19), the intracartilage tensors

(No. 24), the lateral suspensors of the radular sac (No. 25),

52 Wisconsin Academy of Sciences , Arts and Letters

and the tensor of the hood (No. 26), appear to be described here

for the first time. In Helix Trappmann seems to include the

mandibular approximator as a part of the buccal sphincter;

however, in L. s. appressa they are distinct. He also considers

the infraventral odontophoral protractors as a ventral continua¬

tion of the anterior jugal is.

The muscle of the dorsum of the buccal mass is a composite

of the muscles passing dorsad from the ventrum and sides, and

of those connecting the rear of the mass with the dorsal mandible

and of the short radial muscles which extend peripherad from

the basement membrane of the cuticulated epithelium. A con¬

spicuous portion is composed of the large vesicular cells and

other connective tissue which lie between the muscle fibers and

provide the turgidity which characterizes the dorsum.

Salivary glands. A slender muscle for which the term sali¬

vary retractor (No. 29) is suggested, has its insertion in the

ventral surface of the salivary glands. It extends to and becomes

a part of the left buccal retractor (fig. 4). At its insertion, the

salivary retractor also sends a smaller muscle anteriorly and

another posteriorly over the esophagus. The glands are fastened

to the esophagus by numerous slender strands, which may be

named the salivary connectives (No. 30).

Esophagus. The entire tube is composed of an outer layer of

very thin circular muscle and an inner layer of equally thin

longitudinal muscle which rises into and thickens in the longi¬

tudinal folds. The muscle layers of the proesophagus are slightly

thicker than those of the postesophagus. The longitudinal mus¬

cular ridges become slightly larger in the posterior region of

the postesophagus and do not flatten out with extreme dilation

of this part. The two longitudinal muscular ridges lying under

the two gastric nerves are conspicuously larger than the others.

They swing around to the topographically dorsal side of the

crop and gizzard in company with the nerves and other folds,

suggesting torsion of this part of the tract.

Crop. The outer circular muscle layer of the esophagus be¬

comes the inner layer in the crop, and passes to the gizzard, still

bearing, though in modified form, some of the longitudinal ridges

of the esophagus. The crop walls are strengthened primarily by

a new external layer of muscle passing over it from the gizzard.

A slightly thickened, diffuse band of muscle, for which the term

Carriker— Alimentary System of the Snail 53

ingluvial constrictor (No. 31) is suggested, encircles the open¬

ing of the postesophagus into the crop. On each side of the crop

slightly thicker bands run from the ventral to the dorsal ends

of the gizzard lobes, following the curvature of the gizzard lobes.

The designation of right and left anterior gizzard constrictors

(No. 32) is offered for these muscular bands. Ventrally the

bands arise partly in their respective lobes, and partly in the

opposite lobes, interweaving medially where they cross. The

remaining wall of the crop bears loose, somewhat more slender,

fibers which spread from the ventral junction of the gizzard

lobes to the crop constrictor and permit considerable inflation

of the crop.

Gizzard (fig. 17). The lobes of this organ are composed of

strong muscular layers running at right angles to each other:

four circularly directed layers and four layers running in the

long axis of the alimentary tract. The terms gizzard compres¬

sors (No. 33) and gizzard tensors (No. 34), respectively, are

suggested for these layers. The compressors reach circumferen¬

tially from the middorsal to the midventral tendons, and extend

laterally to the anterior and posterior margins of the lobes. The

tensors are sandwiched between the compressors, and their ends

terminate among the fibers of the compressors anteriorly and

posteriorly. Dorsally and ventrally the tensors decrease in bulk,

finally disappearing as the compressors merge into single mus¬

cles connecting with the gizzard tendons.

Heidermanns (1924) found but six muscular layers in the

European L. stagnalis , and points out that the number cannot

be ascertained with consistency as it shows a tendency to dimin¬

ish with increase in the size of the animal. His description of the

gizzard musculature of the European L. stagnalis , with the ex¬

ception of the number of compressor and tensor layers, fits that

of L. s. appressa very closely.

Pylorus . The same thin sheet of circular muscle found in the

crop is continued in the pylorus as the inner lining. Also present

are the companion muscle bands to the anterior gizzard con¬

strictors, the posterior gizzard constrictors (No. 35), which are

placed as are the anterior pair, although the interweaving of the

ventral portions of the muscles is more marked. The remainder

of the pyloric wall is composed mostly of loose, spongy, circular

muscle, with the exception of the folds which are strengthened

by small fibers running in all directions in connective tissue. The

54 Wisconsin Academy of Sciences, Arts and Letters

muscular wall of the retrocurrent passage is markedly thinner

than that of the procurrent passage. The external wall of the

minor passage is thin and is supported by relatively little muscle.

The circular muscle remains of uniform thickness as far as the

atrium, where strong fibers are particularly evident in the

boundary between the hepatic vestibule and the atrium. In the

latter the muscular wall becomes exceedingly thin and trans¬

parent, with faint thickenings suggestive of the atrial cor¬

rugations.

Liver, The hepatic ducts are supported by a relatively strong-

muscular wall which passes about one third of the distance into

the liver in the principal ducts. The fibers in the ducts are

found to run in several directions, although the circular ones

predominate. The corrugations of the principal hepatic ducts are

supported by corresponding, slight ridges of muscle and connec¬

tive tissue. The thickness and frequency of the fibers decrease

as the ducts pass peripherally in the liver and decrease in size.

Muscular elements are distributed even as far as the terminal

liver follicles. Under high magnification single fibers are evident

circumventing the liver follicles, and in living mounts, movement

of small groups of such liver cells may be observed.

Cecum. This is composed of a relatively thin outer layer of

circular muscle and some inner longitudinal fibers, most of which

are found in the cecal folds.

The intestine and rectum are encircled principally by a very

thin layer of circular muscle. The circular muscle layer in the

pellet-molding region and in the rectum is slightly stronger,

particularly in the pellet-compressor and anus. The typhlosolic

muscle (No. 86), as the muscular support of the intestinal

typhlosole may be called, is composed principally of circular

muscle which has invaginated in a thick longitudinal fold. In

the direction of the anus, the muscular wall of the rectum

thickens and the circular and longitudinal muscular elements

become more distinct. At the anus the circular muscle reaches

considerable size as the strong anal sphincter (No. 37). Longi¬

tudinal muscle passes radially out of the rectum and into the

musculature of the pneumatoporal lip, and because of its service

as the anal dilator (No. 38), may be so named. In the anal wall

and in the surrounding pneumatoporal lip, the longitudinal and

circular muscles are closely intertwined and further strength¬

ened by minor muscles running obliquely.

Carriker— -Alimentary System of the Snail

55

In its entire course over the surface of the liver the intestine

is held firmly to it by strong slender strands. In the rectal region

these become bulkier and more numerous, and extend from the

rectal walls, by way of the rectal, sinus which surrounds the

rectum, to the walls of the rectal sinus. For the former strands

the term intestinal connectives (No. 39) is offered; and for the

latter, rectal connectives (No. 40).

Summary of the Muscles of the Alimentary System of

L. s. appressa

(new terms in italics)

I. Extrinsic muscles: buccal mass to body wall:

1. dorsolateral protractors

2. preventral protractors

3. postventral protractors

4. dorsolateral retractors

5. buccal retractors

6. preventral levators

7. postventral levators

8. suboral dilators

9. dorsomandibular dilators

10. suspensor of the radular sac

11. labial retractors

12. labial sphincter

II. Intrinsic muscles: in buccal wall from cartilage to man¬

dibles :

13. anterior jugal is

14. posterior jugalis

15. buccal sphincter

16. mandibular approximator

17. dorsal odontophoral flexor

18. ventral odontophoral protractor

19. infraventral odontophoral protractors

III. Intrinsic muscles : from cartilage to radula and radular sac :

20. supramedian radular tensors

21. supralatera! radular tensors

22. inframedian radular tensors

23. infralateral cartilage tensor

24. intracartilage tensors

56 Wisconsin Academy of Sciences , Arts and Letters

IV. Intrinsic muscles: outward from the radular sac:

25. lateral suspensors of the radular sac

26. inferior suspensor of the radular sac

27. superior suspensor of the radular sac

28. tensor of the hood

V. Muscles of the remainder of the tract :

29. salivary retractor

80. salivary connectives

31. ingluvial constrictor

32. anterior gizzard constrictors

33. gizzard compressors

34. gizzard tensors

35. posterior gizzard constrictors

36. typhlosolic muscle

37. anal sphincter

38. anal dilator

39. intestinal connectives

40. rectal connectives

VASCULARIZATION OF THE ALIMENTARY SYSTEM

In L. s. appressa the organs are supplied with blood by means

of an extensive multiramous system of arteries which ultimately

branch richly as minute capillaries over the organs. From the

capillaries the blood empties directly into the large hemocoels

where it bathes the organs. The venous system has lagged far

behind the arterial system in specialization, and consists only of

series of connected passages. These lack also the special mus¬

cular coat which lines the arteries. The alimentary system re¬

ceives the greatest volume of arterial blood; the reproductive

system, the next largest amount.

The blood itself is a moderately clear, faint milky-colored,

watery fluid lacking hemoglobin, but containing hemocyanin and

amebocytes. The wandering cells display considerable activity,

are moderately numerous, and float freely in the body fluid ; in

the contracted state they are delicate highly translucent spheres

about 8 ii in diameter; when expanded they extend variably

shaped pseudopodia and may cover an area fully five times the

original. Cuenot (1892) described the phagocytic activity of

these cells in L. stagnalis.

Carriker — Alimentary System of the Snail 57

Thorough work on the topography of the circulatory system

of the Basommatophora, with the exception of that of Heider-

manns (1924), is entirely lacking; and the latter paper is con¬

cerned only with the circulation in the gizzard lobes of some

Lymnaeidae. Faust (1920), F. C. Baker (1900), and Lacaze-

Duthiers (1872) make but passing mention of the principal ves¬

sels to be found in various Lymnaea. In the stylommatophoran

pulmonates the detailed work of Schmidt (1916) on the topog¬

raphy of the vascular system of Helix pomatia is still the best

account. Jourdain (1879) described the terminal openings of

the capillaries in Avion rufus .

In the first part of this description the general topography

of the arteries leading to the various organs of the alimentary

system will be presented. The arteries, where homologous, will

be named in accordance with previous pulmonate nomenclature,

particularly that of Schmidt (1916) on Helix pomatia. Each

artery where designated by specific name, will also be denoted

by a parenthetical capital letter which corresponds with similar

designations in the illustrations. Later, the specific vasculariza¬

tion of each of the organs of the system will be considered.

General topography of the arteries (figs. 1, 2, 4-6, 17). The

two-chambered heart is located in the left ventral wall of the

lung, and lies directly over and slightly to the left of the um¬

bilical chink in the shell. The aorta (A) leaves the ventricle and

passes the short distance to the prointestinal loop and is readily

seen lying over the tunica propria ventrally (%. 4). It runs

under the foremost portion of the prointestinal loop and branches

immediately into the three principal arteries of the body: the

cephalic (C) which supplies the organs of the cephalic hemocoel

and the foot (figs. 1, 2, 4) ; the dorsovisceral (VD) which pene¬

trates the dorsal areas of the organs in the visceral hump (fig.

17) ; and the ventrovisceral (VV) which runs through the ven¬

tral portions of the visceral hump (fig. 4).

From its origin just behind the prointestinal loop and to the

right of the esophagus, the cephalic artery (C) passes over the

loop and turns sharply leftward to make a complete turn around

the esophagus. This brings it back to the right side of the

esophagus again, whence it continues cephalad in company with

the esophagus. The artery passes through the bulk of the genital

organs beneath the lung, and at the cervical septum plunges

58 Wisconsin Academy of Sciences , Arts and Letters

through and is bound securely by the connective tissue there. In

the cephalic hemocoel it passes under the esophagus and finally

courses through the central nervous system to branch into sev¬

eral smaller arteries under the base of the buccal mass (fig. 2).

The following arteries are given off by the cephalic artery in its

course forward :

Over the anterior arc of the prointestinal loop, one or two

compressor arteries (CP), in company with, two compressor

arteries given off by the aorta, are passed to the pellet-compressor

of the prointestine. The typhlosolic artery (T) spreads over the

typhlosole. Minor vessels, the prointestinals (IP), pass to other

portions of the prointestine (fig. 17).

Just in front of the junction of the crop and the posteso¬

phagus one or more post esophageal arteries (ET) are given off

to the postesophagus.

Three genital arteries (G) are extended to the spermatheca,

the upper prostate, and the lower prostate from the cephalic

artery as it passes beneath the diaphragm of the lung. The

cephalic artery is here bound firmly to the reproductive glands

by connective tissue.

In the cephalic hemocoel the large salivary artery (S) passes

off to the left supplying the midventral portion of the salivary

glands. It there divides into five smaller vessels, two salivary

arteries (S) and three midesophageal arteries (EM). The for¬

mer branch richly over the inner surfaces of the salivary glands,

one to each gland, and send some smaller vessels to the esophagus

(%. 4).

At the central nerve ring the cephalic artery passes first

under the abdominal and parietal ganglia and then over the

pedal ganglia. Minute cerebral arteries (CE) are extended out

around the ganglia (figs. 2, 4) . Just in front of the central nerve

mass and at the posterior base of the buccal mass, the cephalic

artery forks many times, forming a cephalic arborescence. One

fork, the penial (PE), passes to the right over the preputium. A

right and left cephalic artery (CR, CL) extend forward under

the sides of the buccal mass to the anterior cephalic body wall.

Here each sends a vessel, the probuccal artery (BP) , mediad and

around the oral aperture ; the principal vessels of the right and

left cephalics pass to the velum. Two unpaired branches, the

ventrobuccal artery (BV) and the pedal artery (P) , run straight

Carriker — Alimentary System of the Snail 59

forward under the buccal mass ; the pedal plunges into the foot,

and the ventrobuccal turns dorsad, penetrating the buccal mass.

Finally, the paired dorsobuccal arteries (BD) pass dorsally over

the rear of the buccal mass from the lateral extremities of the

arborescence (fig. 5). They follow, one on each side, the path

of the buccocerebral commissures, pass along the outer sides of

the buccal ganglia, and run anterior at the sides of the pro¬

esophagus, over the buccal mass. One proesophageal artery

(EP) is given off on each side to the esophagus and one post-

buccal artery (BT) extends on each side over the rear of

the buccal mass and under the proesophagus. The last tribu¬

taries of the dorsobuccals are a pair of salivary duct arteries

(SD) which run posteriad over the ducts of the salivary glands.

Occasionally only one duct will be supplied from the dorsobuccal

to one of the lobes of the salivary glands, in which case the other

will be vascularized by a branch from the salivary arteries.

The dorsovisceral artery (VD) arises just behind the ante¬

rior arc of the prointestinal loop, runs posteriad over the mid¬

dorsal surface of the gizzard and pylorus and closely alongside

the cecum as far as its base (fig. 17). The following arteries

branch from the dorsovisceral artery:

The dorsogastric (GD) passes to the left over the midregion

of the gizzard and sends two dorsoingluvial arteries (ID) ante¬

riorly over the dorsolateral portions of the crop; two principal

dorsal gizzard arteries (ZD) into the lobes of the gizzard; the

minor pyloric artery (PN) posteriorly over the dorsal flexure

of the pylorus; one of the largest of the genital arteries (G) to

the left and upward into the genital organs; and a large rectal

artery (R) which passes also to the left, then swings dorsad

and to the right over the coiumellar muscle which partly roofs

the stomach organs. At this muscle the rectal artery branches

thickly, then continues to the right and upward where it meets

the posterior intestine. Here it bifurcates and the two rectal

arteries then pass along the rectum within the rectal sinus

almost to the anus. The genital artery passes a single esophageal

artery (IT) to the crop and esophagus.

In the vicinity of the cecum two or more cecal arteries (CC)

pass from the dorsovisceral artery to the cecum (fig. 17).

Branches of these extend to the right portion of the prointestinal

loop as prointestinals (IP) and into the anterior lobules of the

60 Wisconsin Academy of Sciences, Arts and Letters

posterior liver lobe as hepatics (H). At the base of the cecum

the dorsovisceral artery branches extensively, forming a dense

vestibular arborescence over the junction of the hepatic ducts in

the hepatic vestibule. From this center of ramification one or

more atrial arteries (AT) spread over the atrium branching pro¬

fusely. One of these branches passes on as a prointestinal over

the prointestine. Many minute vestibular arteries (VT) vascu¬

larize the dorsal walls of the hepatic vestibule. The principal

branches from this arborescence, some four main prohepatics

(HP), run posteriorly into the anterior lobules of the posterior

liver lobe by way of the hepatic ducts ; smaller arteries are also

passed into the anterior liver lobe. Many of the prohepatics send

prointestinals out of the anterior liver lobules over the posterior

portions of the prointestinal loop.

The ramifications of the ventrovisceral artery (VV) are more

extensive than those of the dorsovisceral artery. It continues as

a straight extension of the aorta into the spire as follows (fig. 4) :

First, several small prointestinals (IP) advance to the lateral

portions of the prointestinal loop from the area under the

gizzard.

Next, the ventrogastric artery (GV) arises under the right

lobe of the gizzard and proceeds to the left under the gizzard,

giving off two ventroingluvial arteries (IV) to the gizzard lobes;

one major pyloric artery (PM) over the right side of the pylorus ;

and one prepyloric (PP) over the left anterior bulge of the

pylorus. A branch of the major pyloric artery, the minor hepatic

(HN), penetrates the anterior lobe of the liver.

A short distance behind the origin of the ventrograstric

artery, the ventral pyloric artery (PV) forks from the ventro¬

visceral, and presses over the ventral surface of the pylorus

(fig- 4).

Beyond the ventral pyloric artery, the ventrovisceral artery

continues as the major hepatic (HM) which vascularizes the

bulk of the posterior liver lobe and penetrates to the very tip of

the spire. It follows closely over the internal curvature of the

columella of the shell, and directs numerous hepatic (H)

branches to the various lobules of the liver. Many of these rami¬

fications pass to the dorsointestinal loop where they spread about

the intestine as the midintestinal arteries (IM) . Some of the an¬

terior hepatic arteries send prointestinals to the posterior por-

Carriker — Alimentary System of the Snail 61

tion of the prointestinal loop. The largest branch artery of the

major hepatic, the postintestinal (IT) , arises in the midregion of

the liver and passes peripherally to the posterior part of the

intestine at that point where the intestine runs off of the liver

into the postintestinal loop. The postintestinal artery follows the

postintestine as far as the rectal artery and the rectum. One or

two prointestinal arteries are found jumping from the post¬

intestinal artery onto the prointestine at that point where the

postintestine circles parallel to a portion of the prointestinal

loop. An artery to the ovotestis, the gonadal artery (GO), arises

from the major hepatic artery near the origin of the post¬

intestinal artery.

Some variability is found in the origin of the minor arteries,

but the particular portion of tissue being vascularized remains

constant. Of the terms employed for the arteries, the following

are adopted from those used in previous works : aorta, cephalic,

salivary, penial, pedal, right and left cephalics, rectal and

hepatic. The terms visceral, esophageal, intestinal and buccal,

also used in earlier works, have been modified for use in the

present description.

Specific Vascularization

Buccal mass (figs. 5, 6, 19-21). This is vascularized by the

paired probuccal, single ventrobuccal, paired postbuccal and

paired dorsal buccal arteries (fig. 5). The probuccals pass about

the oral structure, and especially between the fibers of the buccal

sphincter and the mandibular approximator muscles. The dorso-

buccal penetrates the dorsal part of the mass, passing beneath

the anterior jugalis muscle alongside the posterior jugalis mus¬

cle, and sends special arterioles into the buccal glands in the

dorsolateral wall. The postbuccal vascularizes the rear dorsal

part of the mass, in particular the portions of the buccal gland

there. Throughout the glandular tissue and adjacent connective

tissues these arteries pass terminally into minute and thickly

distributed capillaries, some of which measure only three micra

in outer diameter (carmine injected).

The principal passage of blood into the mass is by way of

the ventrobuccal artery which enters directly under the cartilage,

at that point where the extrinsic ventral buccal muscles have

their insertion (figs. 5, 6, 19). The artery enlarges broadly at

62 Wisconsin Academy of Sciences , awc£ Letters

its site of attachment. Blood penetrates the buccal mass by way

of two valves which exist as two longitudinal slits in the surface

musculature of the mass and within the anterior and posterior

limits of the attachment of the artery (fig. 19) . There is no spe¬

cialization of the valves in L. s. appressa as there appears to be

in the single buccal valve described for Helix pomatia by Schmidt

(1916). In L. s. appressa the valves consist each of two slight

lateral ledges of tissue which project mediad and meet tightly

when force is brought to bear on the surrounding tissues. The

term odontophoral valve is suggested for the posterior valve.

This leads into the space lying between the cartilage and the

radular sac and the subradular epithelium. For the anterior

valve the designation of outer buccal valve is offered ; this leads

into the spaces lying between the muscles in the anterior ventro¬

lateral portions of the mass, and is surrounded on all but the

anterior border by the insertion of the two extrinsic preventral

protractor muscles. The insertion of the postventral levator

muscles lies between the two valves, and that of the postbuccal

protractor muscles in the anterior borders of the odontophoral

valve. These extrinsic muscles are all so placed with respect to

the valves, that their contraction along with the contraction of

other principal muscles in the forward stroke of the odonto-

phore forces the valves to gape open. On the other hand, the

intrinsic supramedian and supralateral radular tensor, the infra¬

lateral cartilage tensor, the buccal sphincter, and the extrinsic

buccal retractor muscles and possibly others, are so oriented

with respect to the valves that the valves close upon the con¬

traction of these muscles. This occurs on the retracting stroke

of the odontophore when the rear ventral portion of the buccal

mass is constricted to the point where the lateral pillars of the

cartilage are partly collapsed on themselves, clamping the valves

shut.

The outer buccal valve leads into extensive, broadly con¬

tinuous spaces lying about the ventral and lateral portions of

the mandibular approximator, buccal sphincter, infralateral

cartilage tensor, and other muscles. The name buccal sinuses is

offered for these spaces. They pass in the lateral walls about the

muscles as far as the more compact tissues lying over the dorsal

channel of the buccal cavity, and back in the lateral walls to the

rear of the mass. An extensive honey-comb system thus sup-

Carriker — Alimentary System of the Snail 63

plies these muscles with body fluid. The cavity lying about the

cartilage and the radular sac and delimited by the subradular

epithelium dorsally, the circular muscles anteriorly, the lateral

radular tensors laterally, and the posterior jugalis muscle pos¬

teriorly, may be called the odontophoral sinus.

A third valve, which may be designated the inner buccal

valve, makes connection between the odontophoral and the buc¬

cal sinuses. This valve is merely a smooth transverse slit-like

opening in the midventral portion of the buccal sphincter muscle.

It is about the same size as the other valves and closes readily

upon the contraction of the circular muscles of the buccal mass.

All of the muscles of the mass, particularly the buccal

sphincter and mandibular approximator and the supralateral

radular tensor, are perforated by small numerous slit-like pas¬

sages which extend between the muscle fibers from the sinuses.

These vary much in length, average about 70/a, and behave very

much like the inner buccal valve in controlling the passage of

blood. Thus the blood trapped in the odontophoral and buccal

sinuses passes out into the hemocoel by way of these openings

during the relaxation or early contraction of the muscles. Por¬

tions of the posterior jugalis covering the rear of the mass are

composed in part of a laminated musculo-connective membrane

which is also perforated. These openings are rounded or oval

and the smaller measure about 15/a. Emery's aqueous carmine

when injected into the relaxed buccal mass by way of the ventro-

buccal artery readily extravasates through these openings. In an

injected animal which has partly recovered from anesthesia,

these same openings become tightly closed. The aqueous car¬

mine solution does not appear immediately toxic to the injected

snail, as muscles of such snails when retained in Lymnaea

physiological solution remain irritable for over twenty-four

hours. When carmine is suspended over the external valves from

which the ventrobuccal artery has been removed, it may be seen

to enter the valves upon relaxation of the buccal musculature;

but it does not pass back out again when the musculature con¬

tracts. The likeness of the odontophoral, the inner and outer

buccal valves, to the less specialized slits in the musculature is

striking. Possibly these valves were developed from the simple

slit-like openings in the evolution of the buccal musculature.

The distal portion of the radular sac which contains the

radula-forming odontoblasts is supplied with fresh blood from

64 Wisconsin Academy of Sciences, Arts and Letters

the ventrobuccal artery through a notable specialization of the

inferior suspensor muscle of the radular sac (fig. 21). This

muscle originates as a slender thread-like structure in the ven¬

tral portion of the posterior cartilage groove. A short distance

away from its origin it flares broadly into a small triangular¬

shaped inclosure of spongy connective tissue making its inser¬

tion in the distal extremity of the radular sac. The insertion

passes about and partly surrounds the insertion of the superior

suspensor muscle of the radular sac in the central distal tip of

the radular sac. At the initial flare of the inferior suspensor

of the radular sac, a relatively large opening is evident, the

anterolateral margins of which make connection with the car¬

tilage. Thus the suspensor acts as a baffle membrane intercept¬

ing a fraction of the blood passing into the odontophoral sinus

and directing it to the radular sac. In injected specimens this

sac is always heavily charged with the injectant.

A study of the minute anatomy of the buccal mass, combined

with observations on its movements, suggests the following

course of the blood through the mass : In the resting position of

the odontophore, the odontophoral and outer buccal valves are

open and the blood flows freely throughout the mass. During the

forward stroke of the odontophore, the extrinsic ventral muscles

in their contraction continue to hold the valves in an open state.

At the first phase of rasping the entire musculature of the mass

tenses, in particular that of the rear ventral part, and effects

complete closure of the valves, preventing resurgence of the

blood from the sinuses. It is quite probable also that, with the

initial tensing of the muscles, the slit-like openings leading from

them are also pressed shut. Thus the blood in the odontophoral

sinus becomes imprisoned and affords a turgor pressure about

the cartilage and under the radula at the rasping and retracting

strokes of the radula when the greatest expenditure of energy

occurs. In the retraction of the odontophore, the mandibular

approximator and the buccal sphincter muscle constrict mark¬

edly after the receding odontophore, forcing outward some of

the blood in the buccal sinuses. At the termination of the retract¬

ing stroke the musculature relaxes and the remaining blood in

the odontophoral sinus, being under pressure, is forced out

through the openings in the muscles. The rate of beating of the

heart is faster than the rate or protraction of the odontophore

Carriker — Alimentary System of the Snail 65

(at ordinary summer temperatures), so that the blood tends to

accumulate in the ventrobuccal artery while the buccal valves

are closed.

Esophagus (figs. 4, 5). The proesophagus is vascularized by

a pair of proesophageal arteries arising from the dorsobuccal

arteries, and by some four proesophageal arteries passing ante-

riad from the salivary and from the midesophageal arteries.

The principal site of penetration of the tissue by both sets of

vessels is along the four proesophageal folds, into which the

larger arteries pass and fork many times.

The postesophagus receives, on the average, eleven mid¬

esophageal arteries from the salivary artery. Each of these

passes posteriad embedded in one of the postesophageal folds,

specifically vascularizing these more muscular regions and send¬

ing smaller vessels out into the thinner tissue between the folds.

The main arteries extend posteriad almost to the crop. The

posterior portions of the postesophagus are penetrated by one

or more arteries from the cephalic artery, by a branch of the

genital artery passing from the dorsovisceral branch, and by

terminal ramifications of the ventro- and dorsoingluvial arteries

in the crop.

Capillaries (figs. 18, 22, 23). Just as Jourdain (1879) de¬

scribes for Arion, the arterioles in L. s. appressa ultimately ter¬

minate in capillaries which spread out over the tissues in a fine

ramification. These then end at the external surface of the

organs in truncated oval openings through which the blood

empties into the hemocoels. This applies to the entire alimentary

canal with the exception of the very muscular portions of the

buccal mass and the gizzard. In these, as already described for

the buccal mass, the body fluids empty outward through slit-like

passages between the muscle fibers. These passages and slits

appear to be but modifications of the condition existing in the

remainder of the tract. Because of the thinness and slight mus¬

cularity of the walls, the postesophagus affords a good place in

which to observe the capillaries.

As represented in Figure 23 the smaller arteries usually ex¬

tend over the external surface of the organs before plunging into

the tissues. The capillaries spring from these arterioles and

reach in all directions through the connective tissue, muscula¬

ture and along the epithelium for variable distances. They vary

66 Wisconsin Academy of Sciences , Arts and Letters

greatly in diameter, those of 10^ being common, but both larger

and smaller ones being almost as abundant. Beyond its final

branching, each capillary retains a moderately uniform diam¬

eter, and ultimately directs its course towards the external sur¬

face of the organ, there terminating in a slightly flaring aper¬

ture. These capillary ostioles, as the capillary exits may be des¬

ignated, are also extremely variable as to size, those as small as

6fi having been observed. Each ostiole is surrounded by a dense

network of fine muscle fibers (fig. 18) which are so oriented

that upon contraction the ostiole may be completely occluded.

Muscle fibers are found along the entire length of the capillary

situated in the wall of the organ. The carmine solution, when

injected intravascularly in thoroughly anesthetized snails, was

never observed to pass any appreciable distance down those

lengths of capillary lying in the tissue ; it was stopped abruptly

probably by constriction of the muscle fibers surrounding the

vessels. Conversely, when snails recently killed by asphyxiation

were injected, the injectant was sometimes found to pass con¬

siderable distances down the capillaries and extravasate into the

surrounding tissues. This extravasation was undoubtedly made

possible by the complete relaxation and the initiation of autolysis

in the cells, as it never occurred in anesthetized snails. However,

the fact that the carmine solution, even though heavily charged

with a vasodilator, is blocked by constrictions in the capillaries

of anesthetized snails, indicates that the muscle fibers surround¬

ing the tubules are extremely sensitive to chemical stimulation,

and probably play a significant role in the control of the passage

of blood through the tissues.

The course of the arteries and capillaries everywhere may

be readily followed under low magnification by the presence

along the course of the vessels of large numbers of white calci-

ferous concretions. These are discussed under varying names

in the pulmonates by such authors as Cuenot (1892), Kisker

(1923), v. Haffner (1923), Nold (1924), Hoffmann (Simroth

and Hoffmann, 1908-1928) and Baecker (1932). Some of the

concretions lie in the large vesicular cells (cells of Leydig) and

others, intercellularly. It would seem from the fact that all

transitional stages from a few small to many large concretions

are observable in the vesicular cells, that the concretions are

formed within them. These cells measure about 50>i in diameter

and their nuclei are readily seen in living tissues stained with

Carriker— Alimentary System of the Snail 67

methylene blue (fig. 28). The outer, relatively thick wall of the

larger arteries is composed almost entirely of these vesicular

cells, the majority of which do not contain the calciferous

concretions.

In cross section the arteries exhibit two distinct layers of

tissue: a thick outer sheath of vesicular cells and a thin inner

tube of muscle fibers. In the cephalic and visceral arteries the

outer sheath may be composed of as many as three layers of the

large vesicular cells. The number of cell layers decreases in the

smaller arteries and in the smallest arterioles the arterial lumen

becomes encircled by a single layer of only three or four cells

in cross section. In capillaries lying within the tissues, vesicular

cells no longer form the usual external sheath but lie sparsely

and irregularly scattered over the course of the tubule (fig. 23).

In the largest arteries the muscular layer is approximately

5 fi in thickness and is composed almost entirely of circularly di¬

rected muscle fibers. Very few longitudinal fibers are evident.

In the arterioles the circular layer is composed of only a one- or

two-cell layer of muscle and the longitudinal fibers are entirely

absent. In many of the smaller capillaries buried in the tissues,

the capillaries are devoid of any special capillary wall and give

the appearance of tubular spaces tunneled in the muscle fibers

and connective tissue, delimited only by the cells around them.

Mold (1924) and Baecker (1932) describe a similar condition

for the arteries of such Stylommatophora as Helix , with the

exception that in the larger arteries the longitudinal layer of

muscle is well developed, and the smaller arteries bear an in¬

ternal lining of endothelium-like, flattened muscle cells.

The type of capillary vascularization described above for the

postesophagus is found in all parts of the alimentary system

where the organ walls are relatively thin, as in the proesophagus,

crop, pylorus, intestine and rectum. The number of fibers sur¬

rounding the capillary ostioles varies with the muscularity of

the tissues : for example, those ostioles in the wall of the anterior

portion of the proesophagus, which is relatively less muscular,

are not delimited by as many fibers as those in portions of the

crop constrictor muscle. Then again, the degree to which the

capillaries taper in diameter is correlated with the relative

thickness, the complexity, and the glandular nature of the tissues

in question. Thus in the tissues of the buccal glands, the salivary

68 Wisconsin Academy of Sciences , Arts and Letters

glands and the liver, the capillaries are found to diminish to an

outer diameter of 8/x. In these glandular regions the capillaries

spread over individual groups of cells. In the liver the capillary

network embraces the individual liver follicles. In the salivary

glands, where the cells are so large, sometimes the capillary

network covers groups of only a few cells. Even in the glandular

areas, however, the capillaries open to the hemocoels by way of

capillary ostioles.

Salivary glands (fig. 4). These are supplied by the two prin¬

cipal salivary arteries which spread out fan-shape over the inner

surfaces of the glands and send minute capillaries throughout

the glands between the cells.

Crop and Gizzard (figs. 4, 17). The crop is vascularized by

the four ingluvial arteries and a part of the postesophageal

artery extending from the genital branch of the dorsovisceral

artery. These branch most abundantly over the crop constrictor

and anterior gizzard constrictor muscles. The four large dorsal

and ventral gizzard arteries pass almost directly from the dorso-

and the ventrovisceral arteries into the dense musculature of

the gizzard lobes. They penetrate the dorsal and ventral por¬

tions of the lobes near the tendons, and pass through slit-like

openings (fig. 22) about the muscle layers within. In well-

injected specimens dehydrated in alcohol and cleared in xylol,

the clearly outlined arteries may be seen to pass first through

the outer gizzard compressor to the gizzard tensor muscle layer,

and so on into the innermost layers, en route passing branches

to all areas of the lobes between the muscular layers. Smaller

arteries also penetrate the lobes along the anterior and posterior

sides of the gizzard and in the tendons. Within the lobes the

arteries open into small sinuses which lie about the muscle fibers,

particularly between the muscle layers. These sinuses in turn

lead to the exterior by way of the usual slit-like ostioles between

the muscle fibers. The ostioles are most numerous in the ante¬

rior and posterior margins of the gizzard along the anterior and

posterior gizzard constrictors. The slit-like openings through

which the arteries plunge into the lobes act as “valves” which

close upon the contraction of the muscles, much as do the ostioles.

Carmine injected into the dorsal gizzard arteries of a

“pithed” snail, by way of the dorsogastric artery, passed readily

into all the larger arteries and sinuses of the lobe, but did not

Carriker— Alimentary System of the Snail 69

extravasate. Similar injections in thoroughly relaxed snails

flowed out freely through the ostioles. The passage of the blood

in the living gizzard appears to be as follows: blood runs into

the relaxed lobes, propelled by the pumping of the heart. Upon

contraction of the gizzard the entrances of the gizzard artery

are blocked and some of the blood within the lobes is forced out

the ostioles. Upon relaxation the openings to the gizzard arteries

part and the blood in the arteries, under greater pressure than

that in the relaxed gizzard, flows into the gizzard. Also the ex¬

panding gizzard muscles tend to form a partial vacuum which

augments the flow of blood from the arteries and causes an in¬

flow of blood from the hemocoel through the ostioles. When

active, the gizzard lobes undergo a continuous contraction and

relaxation, so that they undoubtedly act also as a kind of pump

which keeps the blood flowing through the gizzard musculature.

Heidermanns (1924) in his work on the gizzard of Lymnaea

found essentially the same picture as presented above for L. s.

appressa. He does not believe that the blood passes freely

through the gizzard. However, the anatomy of the gizzard alone

indicates that in the relaxed gizzard the blood may flow freely

through it.

Pylorus and Cecum (figs. 4, 17). The ventral fold of the

pylorus is vascularized by the ventropyloric artery; the major

and minor folds by the major and minor pyloric arteries; the

interior external portion of the retrocurrent passage wall by the

propyloric artery; the corrugations of the vestibule and the

atrium by branches of the multiramous vestibular and atrial

arteries, respectively. The enlarged posterior portion of the

major pyloric fold which occludes the hepatic vestibule also re¬

ceives smaller vessels from the vestibular arborescence. The in¬

tervening thinner walls are penetrated by capillaries branching

from the principal arteries. The cecal arteries pass externally

over and vascularize principally the cecal folds.

Liver (fig. 4). The anterior lobe of the liver is penetrated

by a minor hepatic artery branching from the major pyloric

artery, and by one or more very small minor hepatic arteries

which leave the vestibular arborescence and pass up the hepatic

duct. The principal mass of the posterior lobe is vascularized

by the major hepatic artery which enters the anteroventral por¬

tion of the lobe, and by it s offshoot, the postintestinal artery.

70 Wisconsin Academy of Sciences , Arts and Letters

The anterodorsal portions of the lobe are entered by the numer¬

ous prohepatic arteries which arise in the vestibular arbores-

cence and pass into the liver lobules by way of the hepatic ducts.

In contrast to the prohepatic arteries which pass into the liver

from a common source in palmate forking fashion, the hepatic

arteries from the major hepatic branch along the main stem of

the artery as it passes to the tip of the spire.

Intestine and Rectum (fig. 4). In the prointestinal loop the

principal sites of vascularization are the pellet-compressor and

the typhlosole. To the former pass three or more compressor

arteries which form sprays of arterioles completely around the

circumference of the compressor. To the latter passes usually a

single typhlosolic artery which sends branches in opposite direc¬

tions over the typhlosole. These in turn direct branching arteri¬

oles down into the musculature of the typhlosole where the

capillaries diffuse. The remainder of the prointestinal loop is

moderately supplied with prointestinal arteries. The mid- and

postintestines have no special points of vascularization but are

densely and uniformly covered by numerous branching mid-

intestinal and postintestinal arteries which pass arterioles com¬

pletely about the entire tube. The more numerous capillaries

pass over the intestinal costae. The two lateral rectal arteries

course anad over those portions of the rectal wall which support

the raphe and the pseudoraphe, an artery to each. From these,

branch arteries spread over the rectum, though not as profusely

as do the arteries over the midintestine. The rectum lies in the

rectal sinus.

Generalizations . Certain generalizations with regard to the

vascularization of the alimentary system may be appropriately

stressed. The organs most abundantly vascularized are those

performing the most physical work. In order of their decreasing

activity these are: the buccal mass, the gizzard, pylorus, and

liver. Of these organs, with exception of the buccal mass and

the gizzard, the parts most generously supplied with blood ves¬

sels are the muscular folds and epithelial corrugations. The

arterioles pass into capillaries which may terminate in either

the rounded ostioles of the esophagus, crop, pylorus, intestine,

rectum, liver and salivary glands ; or the slit-like ostioles between

the muscle fibers of the buccal mass and the gizzard. Passage

of blood through the organs is impelled not only by the action

Carriker— Alimentary System of the Snail

71

of the heart, but is greatly facilitated, as well as probably con¬

trolled, by the muscular activity of the organs themselves. The

rich vascularization of the mid- and postintestines is suggestive

of a function other than peristaltic.

The arterial source of each vessel named and described in

the previous paragraphs is indicated in the following outline :

Summary of the Arteries Supplying the Alimentary

System of L, s. appressa

1. aorta, A

1. compressors, CP

2. cephalic, C (arborescence)

1. right cephalic, CR

2. left cephalic, CL

1. probuccal, BP

3. ventrobuccal, BV

4. pedal, P

5. penial, PE

6. cerebral, CE

7. dorsobuccal, BD

1. proesophageal, EP

2. salivary duct, SD

3. postbuccal, BT

8. salivary, S

1. midesophageal, EM

9. prostatic, CP

10. genital, G

11. spermathecal, GS

12. postesophageal, ET

13. typhlosolic, T

14. prointestinal, IP

3. dorsovisceral, VD

1. dorsogastric, GD

1. dorsoingluvial, ID

2. dorsal gizzard, ZD

3. rectal, R

1. columellar, CO

4. minor pyloric, PN

5. hepatic, H

8. genital, G

72 Wisconsin Academy of Sciences, Arts and Letters

2. cecal, CC

3. atrial, AT

4. prointestinal, IP

5. prohepatic, HP

6. vestibular, VT (arborescence)

4. ventrovisceral, VV

1. prointestinal, IP

2. ventrogastric, GV

1. ventroingluvial, IV

2. ventral gizzard, ZV

3. ventropyloric, PV

4. major pyloric, PM

5. minor hepatic, HN

6. propyloric, PP

3. major hepatic, HM

1. midintestinal, IM

2. postintestinal, IT

3. hepatic, H

4. prointestinal, IP

5. gonadal, GO

INNERVATION OF THE ALIMENTARY SYSTEM

Central nervous system and principal nerves (figs. 1, 2, 4,

5, 11, 17, 24, 25). Bargmann (1930) in a comparative re¬

investigation of the morphology of the central nervous systems

of the Pulmonata has clarified the terminology of the central

nervous ganglia. According to her classification of eight differ¬

ent types of pulmonate central nervous systems, L. stagnalis

falls in the basommatophoran group. This is the most unspe¬

cialized group, in which the minimum concentration of the

ganglia is evident, and all nine ganglia are separated by defi¬

nite connectives (fig. 24).

As far as can be ascertained, the only extensive descriptions

of the principal nerves in the Lymnaeidae is that of Lacaze-

Duthiers (1872). The topography of these nerves, as far as

that author follows them, is in agreement with that found in

L. s. appressa. F. C. Baker (1900, 1911) gives brief descrip¬

tions of the principal nerves in the Lymnaeidae. There is so

little agreement in the literature as to the terminology, and in

some cases the course, of the principal nerves of the visceral

Carriker— Alimentary System of the Snail 73

chain! that an effort is made here to adopt a logical nomen¬

clature based as much as possible on that of such investigators

as Bargmann (1930), Merker (1913), F. C. Baker (1900,

1911), Lacaze-Duthiers (1872), Simroth and Hoffmann

(1908-1928) and Schmalz (1914). The work of Schmalz on the

morphology of the nervous systems of Helix pomatia is prob¬

ably the most detailed of any on the Pulmonata, although he

does not follow it out to the minute innervation of the organs.

Of the ganglia in the visceral chain, the pleural ganglia

do not send out nerves; the two parietal and the single ab¬

dominal ganglia together extend seven principal nerves to the

mantle and to the visceral organs. The terms suggested here

for these nerves are given below (fig. 24). Throughout this

section the parts of the nervous system will be designated in

parentheses in lower case letters which correspond to similar

notations in the illustrations.

Right parietal ganglion (gpar) : (1) The anterior pallial

nerve (pla) anastomoses with the (2) median pallial nerve

(plm) from the abdominal ganglion to form a single nerve,

the internal pallial (pli) which passes in the floor of the lung

in the direction of the pneumatoporal lip and the anus. It is

questionable whether the term internal pallial of Merker and

anterior pallial of Bargmann, F. C. Baker and Lacaze-Duthiers

is synonymous to the usage here adopted of anterior pallial,

internal pallial, or to a combination of both. The median pal¬

lial is synonymous with what Merker designates as anal, and

Bargmann as pallial (3) The right pallial (plr) innervates

the mantle over the roof of the lung and sends one large nerve

to the osphradial ganglion (go) and smaller ones in the direc¬

tion of the anus. There is no disagreement over this nerve,

although Merker adds the word external to distinguish it from

his right internal pallial.

The left parietal ganglion (gpal) sends out only one nerve,

(4) the left pallial (pli), which innervates the left portion of

the mantle. There is also general agreement on this nerve.

Abdominal ganglion (ga) : There is no confusion over the

(5) aortic nerve (ao) which passes along the cephalic artery

to the left body wall. The median pallial (plm) was discussed

above. (6) The ventropallial nerve (plv) reaches posteriad to

the portion of the mantle running under the columellar muscle.

74 Wisconsin Academy of Sciences , Ar£$ cme£ Letters

Bargmann refers to it only as a pallial nerve, F. C. Baker as

the anterior mantle, and Merker as the cutaneous. (7) The

last nerve, the genital of Bargmann and Lacaze-Duthiers, intes¬

tinal of Merker and subintestinal of F. C. Baker, passes pos¬

tered between the median pallial and the aortic nerves. Beyond

the cervical septum it branches into about five large branches

which innervate the reproductive system, the columellar muscle,

the pericardium and the digestive system. Because this nerve

supplies so many of the visceral organs, the new term splanch¬

nic nerve (sp) is suggested for it, and the older terms for it

are given to the appropriate branches, as follows : that passing

over the columellar muscle, the columellar nerve (co) ; two

nerves arising from a knotty enlargement of the splanchnic

nerve in the cervical septum and spreading over the reproduc¬

tive organs, the genitals (g) ; that passing into the peri¬

cardium, the cardiac nerve (cr) ; and that passing from the

splanchnic enlargement and along the cephalic artery to the

stomach region, the intestinal nerve (in). A branch of one

of the genital nerves also passes back dorsally along the her¬

maphroditic duct, the gonadal nerve. The term splanchnic com¬

plex (cs) is suggested for the enlargement of the splanchnic

nerve in the vicinity of which arise the intestinal and genital

nerves.

The principal nerves passing forward from the cerebral

ganglia (gc) into the head region have been described for

Lymnaea by Lacaze-Duthiers (fig. 24). (1) A pair of bucco-

cerebral connectives (bcc) extend to the buccal ganglia (gb)

located at the rear of the buccal mass under the esophagus;

(2) a penial nerve (pe) to the preputium; (3) paired tentacu¬

lar nerves (tt) to the tentacles; (4) paired optic nerves (to)

to the eyes; (5) paired superior labial nerves (Is) which ex¬

tend over the dorsolateral surface of the buccal mass and

branch once, sending the dorsolabial nerves (Id) to the dorsal

portion of the lips, and the frontals (f) outward to the base

of the tentacles; (6) and paired inferior labial nerves (li)

which pass over the ventrolateral surfaces of the buccal mass

and branch into three smaller nerves: the inner nerves, the

ventrolabials (Iv), innervate the ventral portion of the lips;

the two outer branches, the velar nerves (vl), penetrate the

outer portions of the velum. Lacaze-Duthiers names only the

main stems of the nerves, calling them the inferior labials and

Carriker — Alimentary System of the Snail 75

the superior frontolabials. The names of the ramifications

of these nerves have been taken in part from this author's

terms for the principal nerves.

Specific innervation of the alimentary system (figs. 4, 5,

11, 17, 24, 25). With the exception of Heidermanns’ work

(1924) there has been no intensive work on the innervation

of these organs in the Lymnaeidae. Meager descriptions are

given by Lacaze-Duthiers (1872), Amaudrut (1898) and F. C.

Baker (1911). The detailed structure of the nervous system of

such Stylommatophora as Helix has been investigated by such

workers as Smidt (1899, 1901), Baecker (1932) and others.

Principally concerned in the innervation of the alimentary

system of L. s. appressa are the right and left buccal, the right

and left cerebral, the right parietal and the single abdominal

ganglion.

The oral aperture, the lips, the muscles about the mouth

(the suboral and the dorsomandibular dilators and the labial

retractors and sphincter) are innervated by the dor so- and

ventrolahial (Id, lv) nerves (branches of the superior and in¬

ferior labials, respectively) which pass through this region.

These nerves were not found to pass into the buccal mass

(fig. 24).

The musculature of the anal region, including the anal

sphincter and the anal dilator, receives the anal nerves (figs.

4, 25), from the right pallial nerves. These nerve endings con¬

stitute a fine system of branchings in the pneumatoporal lip,

the angle of the mantle and the anal region. No nerves, how¬

ever, could be traced over the rectum itself. Other branches

from the internal pallial nerve, and from the splanchnic nerve

extend over the columellar muscle into the ventral wall of the

rectal sinus, but again no nerve branches could be traced to

the rectum. With more refined technics, nerves could undoubt¬

edly be found on the rectum, since like the intestine it exhibits

peristaltic movements.

The paired buccal ganglia (gb; figs. 5, 25) send out the

principal nerves innervating the greater part of the alimentary

system. These ganglia are two small oblong inconspicuous

bodies connected to each other by the buccal connectives (be;

figs. 24, 25) and lie under the esophagus at its point of entrance

into the buccal mass. Union with the cerebral ganglia is ac-

76 Wisconsin Academy of Sciences, Arts and Letters

complished by means of two lateral buccocerebral connectives

(bcc). In addition to the connectives, six principal nerve

paths pass from the buccal ganglia: (1) the postbuccal, (2)

ventrobuccal, (3) laterobuccal, (4) dorsobuccal, (5) salivary

and (6) gastric nerves; all are paired with the exception of

the postbuccal nerve and this splits as soon as it passes into

the buccal mass.

(1) The postbuccal nerve (bt; figs. 5, 11, 25) arises as a

single nerve from the midventral portion of the buccal con¬

nective and passes ventrally to the posterior lower surface of

the radular sac, penetrating through the posterior jugalis mus¬

cular sac. It forks once over the radular sac sending the

right and left postbuccal nerves (btr, btl) laterad. The right

and left branches fork symmetrically in three directions: the

principal branch, the collostylar nerve (pp), passes down onto

the radular sac, innervates the hood tensor muscle, then swings

up along the radular sac, extending small nerves to the supra-

lateral radular muscle. It terminates along the junction of the

supralateral radular muscle and the subradular epithelium.

The two smaller branches of the right and left postbuccals

reach in a dorsolateral direction over the supralateral radular

muscles, innervating these muscles and the epithelium and

glandular tissue about the entrance of the proesophagus in the

buccal mass. In many specimens ganglionic cells are found

along these outer nerves. The ramifications of the postbuccal

nerve are only exposed when the posterior jugalis muscle is

removed from the buccal mass.

(2) The ventrobuccal nerves (bv; figs. 5, 25) emerge from

the lateral ends of the buccal ganglia and run in a ventrolateral

direction, sometimes in company with the buccocerebral con¬

nectives. Over the cartilage several nerves are extended into

the origin of the supralateral, supramedian and inframedian

radular and infralateral cartilage muscles and into the cartilage

itself. A subsidiary nerve continues antero vent rad, passing un¬

der the anterior jugalis muscle and branching under the infra¬

lateral cartilage muscle in the direction of the odontophoral

and buccal valves. The ventrobuccal nerves thus innervate the

cartilage and all those muscles passing about and from the

cartilage,

Carriker— Alimentary System of the Snail 77

(3) The laterobuccal nerves (bl; figs. 5, 25) extend forward

over the lateral surfaces of the buccal mass a short distance

with the ventrobuccal nerves, and then alone, as the ventro-

buccal nerve swings ventrad. The laterobuccal nerves on each

side of the buccal mass branch in three directions: the ventral

branch sometimes arises from the ventrobuccal nerve and

passes in the anterior jugalis muscle to the lateral region

immediately about the oral aperture, especially into the man¬

dibular approximator and buccal sphincter muscle. The median

branch passes directly into the first lengths of the dorsal odonto-

phoral flexor muscle as it leaves the cartilage. The dorsal branch

plunges forward beneath the anterior jugalis, the anterior ex¬

tensions of the posterior jugalis, and the buccal sphincter mus¬

cles. It splits profusely over and through the buccal glands

lying in the dorsolateral, anterior portion of the buccal mass

and directly adjacent the buccal cavity epithelium. Numerous

nerve branches continue into the region directly about the mouth

and the buccal cavity epithelium. Two or three conspicuous

ganglionic cells are found along some of the branches of this last

nerve.

(4) The dorsobuccal nerves (bd; figs. 5, 25) originate in

the dorsomedial surface of the buccal ganglia and pass over the

buccal mass to the sides of the proesophagus, innervating the

glandular and epithelial regions, as well as the portions of the

posterior jugalis, anterior jugalis, buccal sphincter and odonto-

phoral flexor muscles found over the dorsum of the mass. No

branches could be traced to the mouth. An anterior esophageal

nerve (ea) passes on the median side of each dorsobuccal nerve

over the dorsal surface of the proesophagus as it enters the

buccal mass.

(5) The salivary nerves (s; figs. 5, 25) pass forward as a

part of the dorsobuccal nerves for a short distance before they

run onto the salivary ducts. A single nerve extends over each

duct to the salivary glands, branching generously along its

course. Three or four fine threads fork from the parent nerve

stem and penetrate the principal lobules of each salivary gland

in relatively sparse ramifications. The nerve sheath described

for the nerves of the stomach region by Heidermanns (1924) is

very clearly observed in the salivary ducts and glands. Here by

the methylene-blue technic of Alexandrowics (1932) the prin-

*78 Wisconsin Academy of Sciences, Arts and Letters

cipal nerves may be seen in vivo enclosed in relatively large

sheaths, in which the branch nerves travel some distance before

passing into the surrounding tissues.

Before the last of the principal buccal nerves is described,

two minor buccal nerves will be mentioned. The first of these

are the paired posterior jugalis nerves (jp; figs. 5, 25) which

pass, one from each of the buccal ganglia, to the rear lateral

surfaces of the posterior jugalis muscles. The second, the small

paired buccal retractor nerves (br) , extend, one from each bucco-

cerebral connective, to the buccal retractor muscles, where the

latter run near the connectives.

(6) The right and left gastric nerves (gr, gl; figs. 4, 5, 17,

25) also run forward from the buccal ganglia as part of the

dorsobuccal nerves for a short distance, then jump to the lateral

sides of the proesophagus, to extend along almost the entire

length of the alimentary tract. The gastric nerves, and the

splanchnic nerve from the abdominal ganglion, are the most

extensive and the longest nerves in the snail body. The two gas¬

tric nerves pass down the esophagus in the two largest lateral

esophageal folds, branch profusely over the esophagus and form

a fine network of pro - and postesophageal nerves (ep, et) over

the surface of the tube. As the alimentary tube is followed pos¬

tered, it is evident from the course of the gastric nerves over

it, and from the complete turn of the cephalic artery about it,

that the tube undergoes a clockwise torsion (as seen from the

front) , or a turning to the animal’s left, at the postesophagus and

crop in its development. The torsion appears to have involved

one complete turn of the tube (360°) , so that the topographically

dorsal surface of the alimentary tube posteriad the site of torsion

is also the true morphologically dorsal surface. At the site of

torsion the left gastric nerve turns gradually ventrad and makes

a turn under and around the crop to the right, emerging finally

on the left dorsal surface of the gizzard between the two giz¬

zard lobes (figs. 17, 25). It continues rearward over the left

pyloric fold. The right gastric nerve swings over the post¬

esophagus and completes a full turn about it, emerging ultimately

on the right dorsal surface of the gizzard. It reaches posteriad

over the right pyloric fold. Heidermanns (1924) writes that the

left nerve makes a turn of 180° and the right one, 860°. In L. s.

appressa the left gastric nerve makes almost a 270° turn.

Carriker— Alimentary System of the Snail 79

The crop is heavily innervated by ingluvial (i) ramifications

of the gastric nerves which form an extensively ramifying net¬

work over this organ (fig. 25). The principal branches pass

down over the ingluvial and the anterior gizzard constrictor

muscles. Nerves from the crop and from the pylorus pass into

the muscular layers of the gizzard lobes. The pylorus is even

more thickly innervated than the crop by pyloric (p) nerves

branching from the gastric nerves. The denser nerve patterns

occur over and in the major, minor and ventral folds, and the

posterior gizzard constrictor muscle. Particularly heavy innerva¬

tions are found about the vestibular region and the enlargement

of the major fold there.

From the pylorus the gastric nerves pass down over the

intestine to either side of the proximal base of the cecum. The

left nerve runs on the left dorsal side of the pylorus and under

the hepatic duct of the posterior liver lobe; the right nerve

hugs the right dorsal side of the pylorus reaching beneath the

hepatic duct to the anterior liver lobe, and extends over the

atrium into the intestine. Both nerves undergo considerable

ramification as they spread out over the walls of the pro- and

midintestines, becoming, respectively, the prointestinal (ip) and

the midintestinal nerves (im). A large branch of the left

gastric nerve, the typhlosolic nerve (t), inserts itself into and

follows through the tissue of the typhlosole and the intestinal

raphe. The prointestinal pellet-compressor is fully innervated

by the prointestinal nerves as they pass down the intestine.

Branches of the gastric nerves could be traced only into the

anterior lengths of the postintestine with the technics employed;

nor could branches of the genital, columellar, or internal pallial

nerves which penetrate the rectal sinus, be found reaching over

the postintestine or rectum.

Innervation of the remaining gastric regions is brought about

by nerves leading from the dor so gastric nerve plexus (dp; fig.

25) ; this is a nerve center composed of a large number of giant

intestinal ganglionic cells and nerve fibers and is located on the

vantage position of the hepatic vestibule, slightly to the front

of the proximal base of the cecum. The plexus makes connection

with the gastric nerves by means of two stout nerves. It sends

one nerve also out over the ventral pyloric fold, others over the

vestibule, the atrium and the cecum. Most numerous and con-

80 Wisconsin Academy of Sciences , Arts and Letters

spicuous are those nerves sent posteriad into the lobules of

the posterior liver lobe. This heavy ramification spreads over

the surface of the duct and is soon lost in the glandular mass

of the liver. Smaller nerves are sent also to the anterior liver

lobe. The ganglionic cells of the plexus are most concentrated

directly over the vestibule and diminish in number out over the

hepatic ducts and pylorus. They exist in varying numbers

throughout the pylorus, liver, gizzard, crop, esophagus, buccal

mass, and intestine. In the gizzard, as found also by Heider-

manns (1924), these cells are most numerous in the subepithelial

connective tissue layer between the epithelium and the muscle

layers. The largest cells are found in the dorsogastric plexus

embedded in the vestibular vascular arborescence and measure

about 80^ in diameter. A second nerve center, the ventrogastric

nerve plexus (pv; fig. 25) is found on the ventral efferent ex¬

ternal surface of the hepatic duct leading from the posterior

liver lobe, and partly adjacent to the poster o ventral proximal

base of the cecum. This plexus is relatively inconspicuous and

is composed of noticeably smaller ganglionic cells and nerve

threads which lie in the path of the left gastric nerve. The plexus

is in communication with the dorsogastric plexus by means of

the complex network of nerves which interlace throughout this-

heavily innervated stomach region. The gastric nerves also inter¬

communicate by anastomoses which may occur anywhere between

them over the crop, gizzard or pylorus. The nervous picture

found in the stomach region of L. s. o,ppressa is in agreement

with that described by Heidermanns (1924), with one major

exception: Heidermanns states that both gastric nerves pass

directly into the liver; in L. s. appressa this was not found to

be the case.

The dorsogastric plexus anastomoses with the central nervous

system by way of ramifications of the splanchnic nerve: the

intestinal nerve (in) and the gonadal nerve (o; fig. 25). The

intestinal nerve passes to the rear along the cephalic artery,

and after sending off the cardiac nerve (cr) to the pericardium,

reaches the junction of the cephalic, dorsovisceral, ventrovisceral

and aortic arteries. At a point where this arterial branching

presses against the right lobe of the gizzard the intestinal nerve

enters a small group of ganglionic cells, comprising the intestinal

complex (ci). The complex sends minute nerves out over the

Carriker— Alimentary System of the Snail 81

arteries, to the gizzard lobe, into the connective tissue, and two

large intestinal nerves rearward to the dorsogastric plexus.

One of these intestinal nerves passes along the pylorus, and the

other by way of the cecum. The gonadal nerve arises from the

genital nerves and extends posteriad over the hermaphroditic

duct to the ovotestis, passing over the prointestinal loop and the

connective tissue of the dorsal surface of the stomach region.

Several ganglionic cells are evident along its course. At a point

over the pylorus, it sends a fine nerve ventrad which anastomoses

with the dorsogastric plexus. Several ganglionic cells are pres¬

ent at the source of the anastomosing nerve.

Discussion. Schmalz (1914) gives the names of anterior, me¬

dian, and posterior buccal to the buccal nerves in Helix pomatia.

These do not appear to fit the case in L. s. appressa, and probably

correspond as follows with the terms which have been suggested

here for L. s. appressa : dorsobuccals, laterobuccals and ventro-

buccals, respectively. The two gastric nerves he terms the poste¬

rior gastric nerves. The anterior companions to the posterior

gastrics he refers to as the anterior gastrics, which in L. s. ap¬

pressa correspond either with the anterior esophageal or the

dorsobuccal nerves. Other terms here introduced to designate

nerves described in this section are : collostylar, anterior esopha¬

geal, posterior jugalis, proesophageal, postesophageal, ingluvial,

pyloric, vestibular, dorso- and ventrogastric plexi, typhlosolic,

intestinal, intestinal and splanchnic complexes, prointestinal,

midintestinal and gonadal.

Unquestionably the densest nerve patterns of the alimentary

system in L. s. appressa are found in the stomach region, particu¬

larly about the pylorus. This is perhaps indicative of the sig¬

nificant role played by this portion of the digestive tract. The

second densest innervation is that located in the buccal mass.

This could be anticipated from the complexity of the muscula¬

ture and the intricacy of the movement of this organ in food

getting.

82 Wisconsin Academy of Sciences , Arts and Letters

Summary of the Nerves Passing to the Alimentary System

OF L. s. appressa

(arranged according to the source of the nerves)

1. Cerebral Ganglia, gc

a. buccal ganglia, gb

1. gastric, gr, gl

a. proesophageals, ep

b. postesophageals, et

c. ingluvials, i

d. pyloric, p

e. hepatic, h

f. vestibular, v

g. dorsogastric plexus, pd

h. ventrogastric plexus, pv

i. prointestinals, ip

j. typhlosolic, t

k. midintestinals, im

2. salivary, s

3. anterior esophageals, ea

4. dorsobuccals, bd

5. posterior jugalis, jp

6. laterobuccal, bl

7. ventrobuccal, bv

8. buccoeerebral connective, bcc

9. buccal connective, be

a. buccal retractor, br

10. postbuccal, bt

a. right postbuccal, btr

b. left postbuccal, btl

1. collostylar, pp

b. superior labials, Is

1. f rentals, f

2. dorsolabials, Id

c. inferior labials, li

1. ventrolabials, lv

2. velars, vl

d. tentacular s, tt

1. optics, to

e. penial, pe

Carriker — Alimentary System of the Snail

83

2. Right Parietal Ganglion, gpar

a. anterior pallial, pla

1. internal pallial, pli

a. anal, a

b. right pallial, plr

3. Left Parietal Ganglion, gpal

a. left pallial, pll

4. Abdominal Ganglion, ga

a. aortic, ao

b. splanchnic, sp

1. columellar, c

2. genitals, g

a. gonadal, o

3. cardiac, cr

4. intestinal, in

a. intestinal complex, cin

c. median pallial, plm

1. internal pallial, pli (anastomosis)

d. ventropallial, plv

DISCUSSION

Embryological evidence for the specialization and differen¬

tiation of the simple embryonic alimentary tube to give the fully

developed alimentary system of the mature snail is supplemented

by information obtained from a study of the configuration, vas¬

cularization, innervation and particularly the muscularity of

the adult system. This is in keeping with Amaudrut’s theory

(1898) that the complex buccal mass arose from a simple in¬

vagination of the cephalic integument, and Heidermanns, sug¬

gestion (1924) that the gizzard developed in front of the first

flexure of the midgut. The fact that the thin cuticular layer

covering the gizzard lobes internally is not chitinous in contrast

to the thicker chitinous layers of the buccal cavity, may suggest

the relatively more recent phylogenetic appearance of the gizzard

as compared to that of the buccal mass. The various coilings of

the intestine about the other organs attest the extent to which

this part of the alimentary system has followed the spiral turning

of the visceral hump over the foot in the embryological develop¬

ment of the individual. Torsion of the tract itself at the crop

and postesophagus is made conspicuous by the course of the

esophageal folds, the gastric nerves and the cephalic artery over

this region.

84 Wisconsin Academy of Sciences , Arts and Letters

SUMMARY

1. The alimentary system of L. s. appressa is composed of

the following morphologically (histologically and functionally—

see later papers) distinct parts: buccal mass, buccal glands, sali¬

vary glands, proesophagus, postesophagus, crop, gizzard, pylorus,

liver lobes, cecum, prointestine, midintestine, postintestine and

rectum.

2. Forty morphologically distinct muscles are described for

the alimentary tract. The majority of these are concerned

with the manipulation of the complex mouth parts of the buccal

mass. The stomach region is next in muscular complexity, but

does not compare with the buccal mass in the intricacy of mus¬

cular specialization.

3. The arterial system of L. s. appressa is composed of a

much-branching system of arteries and capillaries. About fifty

distinct arteries are described. The capillaries pass over the

walls of the organs and empty into the hemocoels surrounding

them by means of ostioles. The venous system is composed of

successive series of open sinuses, the largest being the cephalic

and the visceral hemocoels. The alimentary system receives the

greatest number of arterial vessels and consequently the largest

volume of blood of any of the body organs.

4. The alimentary system is innervated by a densely ramify¬

ing system of nerves and by variously distributed giant gangli¬

onic nerve cells. The pair of gastric nerves passing from the

buccal ganglia supply the majority of the nerves from the buccal

mass to the intestine. The large splanchnic nerve from the ab¬

dominal ganglion sends an intestinal nerve to the pylorus where

it anastomoses with the gastric nerves in the gastric plexuses.

These are conspicuous concentrations of the ganglionic nerve

cells and mark the site of greatest innervation of the alimentary

system. Some fifty separate nerves are described.

5. Morphological study indicates that torsion of the gut has

occurred at the crop and postesophagus.

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MAPLE SUGAR: A BIBLIOGRAPHY OF EARLY

RECORDS. II.*

H. A. SCHUETTE AND A. J. IlIDE

University of Wisconsin , Madison

Acknowledgment

Grateful acknowledgment is made to Yale University Press and the au¬

thors for permission to quote verbatim from “Sketches of Eighteenth Cen¬

tury America.”

A continuation of the search of early American narratives of

travel and books on natural history, inclusive of diaries, journals

and official communications, in the rich collection of the State

Historical Society of Wisconsin has uncovered sufficient material

to warrant publication of another communication on the subject

of a typically American food. It is quite apparent that a chron¬

ology of maple sugar records as observed in the voyages of ex¬

ploration, missionary activities, colonization with its economic

problems, the botanizing travels of Bartram, Nuthall, Michaux,

and Kalm, Indian lore, the Lewis and Clark expedition, the

slavery issue, technological advancement, and the rise of food

adulteration, is in a sense a segment of North American history

itself.

Observation of maple trees in Canada during the sixteenth

century is attributed to Cartier (1540), discoverer of the St.

Lawrence river. Champlain (1603) unquestionably observed

these trees and Hariot reported them in Virginia in 1590.

The fact that maple trees were tapped for their sap was

mentioned early in the seventeenth century but the production

of sugar was not recorded until much later in the century. The

question of whether the Indians made sugar or even syrup is

controversial. The writings of Thornton (1684), an anonymous

author (1685), Beverly (1722), Keating (1824), Lahontan

(1703), and Verwyst (1744) permit the assumption that sugar¬

making was probably a native art. Charlevoix (1721), on the

* For Part I see Trans. Wisconsin Acad . Sci. 29:1935, 209-236.

89

90 Wisconsin Academy of Sciences, Arts and Letters

other hand, contended that while they made constant use of the

sap, the Indians learned from the white man how to make sugar.

If this is true the knowledge certainly spread among the natives

with great rapidity as did modifications in the process of con¬

centrating sap, such as freezing or the addition of hot stones.

Interest in maple sap as a significant source of supply of

sugar was particularly marked around 1800. The production

of maple sugar was encouraged as a means of freeing the country

of the need to import cane sugar from the West Indies. As the

slavery issue became acute maple sugar production was cited

as a means of combatting slavery.

Chemical interest is first noted in the records of Boyle

(1664), but not until late in the ninteenth century do chemical

analyses appear. Scientific investigation became necessary at

this time when food adulteration became so flagrant that control

legislation became essential.

78. Cartier, Jacques 1585

Bref recit, & succincte narration, de la nauigation faicte

es yles de Canada, Hochelage & Saguenay & autres, auec

particulieres meuers, langaige, & cerimonies des habitans

d’icelles: fort delectable a veoir. Paris, 1545, (a) p. 14;

Richard Hakluyt, The Principal Navigations, Voyages,

Traffiques, and Discoveries of the English Nation. Lon¬

don, 1600. Reimpression verbatim by Edmund Goldsmid.

Edinburgh, 1889, Vol. XIII ii, (b) p. 109; J. P. Baxter,

A. Memoir of Jacques Cartier, Sieur de Limoilou. His

Voyages to the St. Lawrence. New York, 1906, (c) p.

145; H. P. Biggar, The Voyages of Jacques Cartier. Pub¬

lications Public Archives Canada, No. 11, 1924, (d) p. 123.

a. (No mention of the maple in Cartier's book)

“ . . . Nous nommasmes ledict lieu saincte Croix, par

ce que ledict iour y arrivasmes. Aupres d’iceluy lieu ya

ung peuple, dont est seigneur ledict Donnacona, & y est

sa demeurance qui se nomme Stadacone, qui est aussi

bonne terre qui’l soit possible de veoir & bien fructiferete,

pleine de fort beaulx arbres de la nature & sorte de France.

Comme chesnes, ormes, fresnes, noyers, yfz, cedres, vignes,

aubespines, qui portent le fruict aussi gros que prunes de

damas, aujtres arbres ; , . f ”

Schuette and Ihde— Maple Sugar

91

b. (The maple mentioned in Hakluyt's English version

of Cartier's second voyage )x

“ ... we named it the holy Crosse, for on that day

we came thither. Neere vnto it, there is a village, whereof

Donnacona is Lord, and there he keepeth his abode : it is

called Stadacona, as goodly a plot of ground as possibly

may be seene, and therewithal! very fruitfull, full of good¬

ly trees euen as in France, as Okes, Elmes, Ashes, Walnut

trees, Maple tres, Cydrons, Vines, and white Thornes, that

bring foorth fruit as bigge as any damsons, . . . ”

c. (Occurrence of the maple in Canada revealed

in a contemporary Cartier manuscript)

“ . . . We named the said place St. Croix because we

arrived there the said day. Near this place there is a

people of whom the said Donnacona is lord, and there is

his dwelling place, which is called Stadacone, which is as

good land as it is possible to behold, and very fruitful,

full of exceeding fair trees of the nature and kinds of

France, as oaks, elms, ashes, nuts, plum-trees, maples,

cedars, vines, white thorns, which bear fruit as big as

damson plums.”

d. (Occurrence of maple not revealed in copy

of original Cartier manuscript)

“ . . . Nous nommasmes ledict lieu saincte Croix pource

que ledict jour y arrivasmes, Aupres d’icelluy lieu y a

vng peuple dont est seigneur ledict Donnacona, et y est

sa demeurance, lequel se nomme Stadacone, qui est aussi

bonne terre qu’il soit possible de veoyr, et bien fructif-

ferante, plaine de moult beaulx arbres, de la nature et

sorte de France, comme chaisnes, hourmes, frennes, noyers,

prunyers, yfz, seddrez, vignes, aubespines, qui portent

1 The accounts of Cartier’s first voyage to Canada, in 1534, all fail to record observation

of the maple tree (erable, arable). An account of the second voyage, made a year later, was

first published in book form in 1545 under the title Bref recit. This was translated into

Italian and published, along with an account of the first voyage, in Ramusio’s Navigation*

et Viaggi (Venice, 1556). Bref recit apparently disappeared from circulation about this

time. The only known existing copy was discovered in the British Museum more than two

centuries later.

The source for Hakluyt’s version of the first two voyages appears to be an English

translation of Ramusio’s book which he induced John Florio to make in 1580.

Baxter had access to three manuscripts, believed to be Cartier originals, in the Bibli-

oth&que Nationale in Paris. He found none of them in exact agreement with Bref recit

which he believed was printed from the one numbered 5653. He translated the one numbered

5589. Biggar used these same manuscripts, publishing a verbatim copy of 5689 as well as a

translation. At the same time he collated the manuscript with 5653, 6644, and Bref recit.

His work indicates that the maple tree was not mentioned in any of these works by Cartier.

92 Wisconsin Academy of Sciences , Arts and Letters

fruict aussi groz que prunes de Damas, et aultre ar-

bres, ...”

74. Cartier, Jacques 1540

The third voyage of discouery made by Captaine Iaques

Cartier, 1540. unto the Countreys of Canada, Hochelaga,

and Saguenay. Richard Hakluyt. Ibid., p. 149; H. S. Bur-

rage, Editor. Early English and French Voyages. Chiefly

from Hakluyt. 1534-1608. New York, 1906, p. 97.

(Earliest existing version of the third voyage

records the maple in Canada)2

“ . . . Moreover there are great store of Okes the most

excellent that ever I saw in my life, which were so laden

with Mast that they cracked againe: besides this there

are fairer Arables, Cedars, Beeches, and other trees than

grow in France.”

75. Alphonse of Xanctoigne, Jean 1542

Richard Hakluyt. Ibid., p. 163. Here followeth the

course from Belle Isle, Carpont, and the Grand Bay in

Newfoundland vp the Riuer of Canada for the space of

230. leagues, obserued by Iohn Alphonse of Xanctoigne

chiefe Pilote to Monsieur Roberval, 1542.

(The chief pilot of a colonizing party

reports arables in Cartier’s Canada)

“ . . . and in all these Countreys there are okes, and

bortz, ashes elmes, arables, trees of life, prusse trees, co¬

ders, great wall nut trees, and wilde nuts, hasel-trees,

wilde peare trees, wilde grapes, and there have been found

redde plummes.”

76. Thevet, Andre 1575

La cosmographie universelle. Paris, 1575, pp. 1008-

1016.

(Maple trees noted in Canada)

“Entre autres s’en trouve un, qu’ils nomment Cotonyf

lequel est de la grosseur d’un gros noyer pardega. Cest

arbre a este long temps inutile & sans aucun profflt, ius-

ques a ce que quelcun des nostres le voulans coupper, des

2 Cartier’s third voyage of 1540 and the records of Jean Alphonse, chief pilot of

Roberval’s colonizing expedition in 1542, have come down to the present only through the

English version of Hakluyt. He was chaplain to the British Ambassador in Paris in 1583

and, according to Biggar, possibly had access to original French manuscripts which have

since been lost.

8 Rousseau from whose monograph, La Botanique canadienne & 1’epoque de Jacques

Cartier, ( Contributions du Laboratoire de Botcmique de VUniversite de Montreal, No. 28:1937,

p. 29 ) these lines are quoted, opines that Thevet refers to the sugar maple.

Schuette and Ihde — Maple Sugar 93

tite; laquelle estant goustee, fut trouvee de si bon goust,

que plusieurs 1’esgalloient a la bonte du goust de vin, de

sorte que plusieurs recueillerent de ceste liqueur en abon-

dance, & ayda a rafraischir les nostres: Et pour voir &

experimenter dont procedoit la source de ceste boisson,

ledit arbre fut scie, le tronc duquel estant par terre, fut

trouve comme chose miraculeuse au coeur de l’arbre, une

Fleur de lys bien effigiee. . . Les Canadeens n’oubliront

pas l’excellence de ceste liqueur, & se souviendront tou-

siours de ceux qui en trouverent l’usage, veu la bonte de

ce breuvage, meilleur pour vray, que celuy duquel aupara-

vant ils usoient, comme plusieurs de leurs voisins.”

77. Hariot, Thomas 1590

A briefe and true report of the new found land of

Virginia. Franckfort, 1590, p. 23.

Of commodities for building and other necessary vses

“Maple, and also Wich-hazle , whereof the inhabitants

vse to make their bowes.”

78. Champlain, Samuel de 1603

Des Savvages, ou, Voyage de Samvel Champlain, de

Brovage, fait en la France novvelle Tan mil six cens trois.

Paris, 1603, Vol. I, Chap. VIII; H. P. Biggar, Editor, The

Works of Samuel de Champlain. Toronto, 1922, Vol. I,

p. 144.

(Maple trees observed in Richelieu River region)

“ . . . I went in a canoe to the south Shore, where I

saw a number of islands, very productive of fruits, such

as grapes, walnuts, hazelnuts, and a kind of fruit like

chestnuts, cherries, oaks, aspens, poplars, hops, ash, ma¬

ple, beech, cypress, very few pines and fir trees.”

79. Smith, John 1616

A Description of New England: or the Observations,

and Discoveries, of Captain Iohn Smith (Admirall of that

Country) in the North of America, in the year of our Lord

1614 : with the successe of sixe Ships, that went the next

yeare 1615 ; and the accidents befell him among the French

men of warre. London, 1616. Coll. Mass Hist. Soc., [3]

6, 120 (1837).

94 Wisconsin Academy of Sciences, Arts and Letters

(Maple tree not noticed)

Oak is the chief wood, of which there is great differ¬

ence in regard of the soil where it groweth, fir, pine, wal¬

nut, chestnut, birch, ash, elm, cypress, cedar, mulberry,

plum-tree, hazel, sassafras, and many other sorts.

80. Lescarbot, Marc 1617

Histoire de la Nouvelle-France. Paris, 1866, new ed.,

Vol. Ill, (a) p. 750, (b) p. 815; W. L. Grant, The History

of New France. Toronto, 1914, Vol. Ill, (a) p. 194, (b)

256.

a . (Sap as thirst-quencher)

“ . . . And we in France are troubled when we have

lost our way ever so little in some great forest. If they

are tormented with thirst, they have the skill to suck

certain trees, whence tricles a sweet and very pleasant

liquor, as I myself have sometimes proved.”

b. (Forest trees)

“As for the trees of the forests, the most common in

Port Royal be oaks, elms, ashes, birch (very good for

joiner’s-work), maples, sycamores, pine-trees, fir-trees,

whitethorns, hazel-trees, willows, bay-trees, and some oth¬

ers besides which I have not yet marked.”

81. Sagard, Gabriel 1632

Le Grand voyage du pays des Hurons situe en l’Ame-

rique vers la mer douce, es derniers confins de la Nou-

velle France dite Canada. Paris, 1632, pp. 102 and 331;

W. F. Ganong. The Long Journey to the Country of the

Hurons. Toronto, 1930, (a) p. 82, (b) p. 240.

a. (Beech sap as a tonic)

“ ... In the season when sap is rising in the trees we

would sometimes make an incision into the bark of some

big beech, and holding a bowl underneath get the juice

and liquid which dropped from it; this served us as a

tonic for the digestion whenever we were indisposed in

that way. It is, however, a very crude remedy and of little

effect, which sickens rather than strengthens, and the

reason we employed it was the lack of any other sub¬

stance more suitable and better. ...”

Schuette and Ihde — Maple Sugar 95

b. (Forest trees)

“In the woods there are many cedars, called Asquata ,

very fine large oaks, beeches, maples, wild cherry trees,

and a great number of other trees of the same species as

ours, . . . ”

82. Boucher, Pierre 1664

Histoire veritable et natvrelle des moevrs et Prodvctions

dv pays de la Novvelle-France vvlgairement dite le Canada.

Paris, 1664; B. Suite, 1896. Pierre Boucher et son livre.

Proc. Trans. Roy. Soc. Canada, [2] 2 i, 184.

(Water from the maple)

“IFy a vne autre esspece d’arbre, qu’on apelle Herable,

qui vient fort gros & haut: le bois en est fort beau, non-

obftant quoy on ne s’en fert a rien qu’a bruler, ou pour

emmancher des outils, a quoy il est trespropre, a cause

qu’il est extrememet doux & fort. Quand on entaille ces

Herables au Printeps, il en dequote quantite d’eau, qui

est plus douce que de Teau detemplee dans du sucre; du

moins plus agreable a boire.”

83. Boyle, Honourable Robert 1664

Some considerations Touching the Vsefvlness of Ex¬

perimental Natural Philosophy. Oxford, 1664, 2 ed. p. 102.

(Sugar from trees in Massachusetts)

“Since the writing of these last lines, being visited by

an ancient Virtuoso, Governour to a considerable Colony in

the Northern America, and inquiring of him, among oth¬

er particularities touching his Countrey, something in

relation to the thoughts I had about the making of several

kinds of Sugar, he assured me, upon his own Experience,

that there is in some parts of New England, a kind of

Tree, so like our Walnut-trees, that it is there so called;

whose juice, that weeps out of its Incision &c. if it be

permitted slowly to exhale away the superfluous moisture,

doth congeal into a sweet and saccharine substance; and

the like was confirmed to me, upon his own knowledge,

by the Agent of the great and populous Colony of the

Masathusets,”

98 Wisconsin Academy of Sciences , Arts and Letters

84. Hubbard, William 1680

A General History of New England from the discovery

to MDCLXXX. Boston, 1815. Coll . Mass. Hist . Soc., [2]

5, 24 (1815).

(Maple trees in colonial New England)

"... the same trees, plants and ‘shrubs', roots, herbs

and fruits being found either naturally growing here that

are known to do in the northern countries of the like

climate of Europe, and upon trial have been found as

effectual in their operation, and do thrive as well when

transplanted ; as the oak, walnut, ash, elm, maple. . . . ”

85. Hennepin, Louis 1698

A Continuation of the New Discovery of a Vast Coun¬

try in America, Extending above Four Thousand Miles,

between New France and New Mexico with a Description

qu'il 1'eut touche au vif, en feit sortir une liqueur en quan-

of the Great Lakes , Cataracts , Rivers, Plants, and Ani¬

mals. London, 1698, (a) Bon ed., p. 147; (b) Tonson

ed., p. 158.

a. (Maple sap a sovereign remedy)

"... Our common Drink was fair Water, which we

took out of the Springs, Rivers or Lakes; but if any of

us were indisposed at the time when the Trees were in

Sap, more especially if he were afflicted with an Oppres¬

sion or Weakness of the Stomach, we usually made a

Cleft in the Bark of a Maple-Tree; out of which issued

forth a kind of sweet Liquor, which was receiv'd into a

Dish made of Birch-Tree Bark: This Liquor was drank

as a Sovereign Remedy, altho in reality the effects of it

were not very considerable. There are great store of

Maple-Trees growing in the Vast Forests of those Coun¬

tries, and Distill'd Waters may be drawn from them:

Insomuch that having caus'd them to boil for along time,

we made a Reddish sort of Sugar, much better than that

which is taken from the ordinary Canes in the Islands of

America.”

b. (A kind of reddish sugar from maple sap)

"... Our ordinary Drink was Water. If any of us

was indisposed, while the Sap was up in the Trees, we

Schuette and Ihde— Maple Sugar

97

made a hole in the Bark of a Maple, and there dropt out

a sweet Sugar-like Juice, which we saved in a Platter

made of the Bark of a Birch-tree; we drank it as a

Souvereign Remedy, tho it had but small effects. There

are in the Yallies of those Forests great store of Maples,

from whence may be drawn distilFd Waters. After a long

boiling, we made of it a kind of reddish Sugar, much bet¬

ter than that which is drawn from the ordinary Cane in

the Isles of America ”

86. Penicaut 1701

Relation de Penicaut. Pierre Margry, Decouvertes et

Etablissements des Francais dans Fouest et dans le sud du

L’Amerique Septentrionale (1614-1754) Memoires et Doc¬

uments Originaux. Paris, 1883, Vol. Y, p. 417; R. G.

Thwaites, Editor, The French Regime in Wisconsin.

1634-1729. Wisconsin Hist. Coll , 16, 196 (1902).

(Maples near Le Seuer’s fort on the Mississippi)

“There are also more different species of trees than

are found on the lower river— for instance, of the wild

cherry, maple, and plains , (red or swamp maple, Acer

ruhrum. ed.) and of the poplar, a tree which grows so

large that some specimens of it are five brasses in cir¬

cumference. As for the trees called “maple" and plains ,

incisions are made in them early in March, and a tube is

placed in each incision to catch the sap ; this passes

through the tube and falls into a vessel, which is placed

underneath to receive it. These trees flow abundantly

during three months, from the first of March to the end

of May; the Water which they distill is very sweet; to pre¬

serve it, this is boiled until it becomes a syrup, and if it

is boiled longer it turns to sugar."

87. Neville, Ella H., Sarah G. Martin, and Deborah

B. Martin 1703

Historic Green Bay. Green Bay, 1893, p. 70.

(Louis baron de Lahontan honored)

“A marvelous banquet was served by the Indians in

the baron's honor, at which the guests were seated in ori¬

ental fashion on the green sward, under the lofty trees.

98 Wisconsin Academy of Sciences , Arts and Letters

Successively they partook of whitefish boiled in water,

cutlets of the tongue of buck, followed by hazel hen — a

fowl fattened on nuts — a bear’s paw, and, greatest deli¬

cacy of all, the tail of a bear. Then came a bouillon pre¬

pared from a variety of meats, the whole washed down by

what the baron calls a most delicious liquor, made of

maple sugar, beaten up with water.”

88. Diereville 1708

Relation du Voyage du Port Royal de L’Acadie ou de

Lanouvelle France. Rouen, pp. 108-110; L. U. Fontaine,

Voyage du Sieur de Diereville en Acadia. Quebec, 1885,

p. 61; John C. Webster, Relation of the Voyage to Port

Royal in Acadia or New France. Toronto, 1983, pp. 117

and 270.

(Strawberries and maple sugar)

“ . . . Strawberries are no less plentiful in the fields

everywhere, one has the pleasure of eating them with a

Sugar produced in the Country.

Instead of Canes, whose Pores secrete

White Sugar, brought from afar,

Nature, for the Acadian, with kind

Forethought, has put some in the Sycamore.4

When Springtime comes, this tree gives forth

A sweetish liquor from its bark,

And this, in each vicinity,

The Settlers all collect with care.

This seemed a pleasant brew to me

In copious draughts I drank it down;

And Lemons only did we need

To make it into Lemonade.

To obtain this sweet Liquor, which is as clear as

Spring water, a hole, fairly deep, shaped like a trough,

is made in the tree with an axe, a frame of bark is joined

to this reservoir, so that the sap, as it flows, may drop

into it. When it is full, which occurs quickly enough, for

the sap at this time is in its greatest vigour, the fluid

runs by means of a little scupper attached to the trough,

4 This is an incorrect designation as the true sycamore did not grow in Acadia. Diereville

means the sugar maple. Europeans used the term sycamore maple, to designate a native

species, Acer psuedo-platanus.

Schuette and Ihde — Maple Sugar 99

into a vessel at the base of the tree. Several trees are

treated in this way at the same time, so that a quantity

of fluid is obtained, which is carefully collected each day

as long as it is forthcoming. It is boiled in a large caul¬

dron until the drying point is reached; as it condenses

little by little, it becomes a Syrup, & then a reddish Sugar

which is very good/’

89. Byrd Family, Member of 1729

Letters of the Byrd Family. Virginia Mag . Hist., 36,

117 (1928).

(The sugar tree in Virginia)

“To Mr. Warner in England July, 1729 (Virginia)

"As it was our Fortune to pass over all sorts of

ground, we saw many new plants, the most remarkable

of which was, the Sugar Tree, which grows as tall, as an

Elm, and big in proportion. By Tapping this Tree in the

Spring, a Liquor issues out of it, which may be boiled

into good sugar. I shall endeavor to get you one of these,

against the return of the Ships, and every thing else, that

I believe may be agreeable.”

90. Anon. 1730

Observations Botaniques III.

Histoire de P Academic Royale des sciences, Paris,

1730, 65; l’Abbe Laflamme, Michael Sarrazin: Materiaux

pour servir a Phistoire de la science en Canada. Proc.

Trans. Roy. Soc. Canada, 5, iv, 21 (1887).

(A report on the maple tree)

“M. Sarrazin, Medecin de Quebec, Correspondant de

l’Academie, a trouve dans PAmerique Septentrionale qua-

tre especes d’Erable qu’il a envoyees au Jardin Royal,

apres leur avoir impose des noms. La 4me qu’il appelle

Acer Canadense Sacchariferum, fructu minori. D. Sarra¬

zin, est un arbre qui s’eleve de 60 a 80 pieds, dont la

seve qui monte depuis les premiers jours d’Avril jusqu’a

la moitie de Mai est assez souvent sucree, ainsi que Pont

ai semen t reconnu les Sauvages & les Frangois. On fait a

l’arbre une ouverture, d’ou elle sort dans un vase qui la

regoit, & en la laissant evaporer, on a environ la 20me

partie de son poids, qui est de veritable sucre propre a

100 Wisconsin Academy of Sciences, Arts and Letters

etre employe en confitures, en sirops, &c. Un de ces

arbres qui aura 3 ou 4 pieds de circonference, donnera

dans un printemps, sans rien perdre de sa vigueur, 60 a

80 livres de seve. Si on en voulait tirer davantage, comme

on le pourrait, il est bien clair qu’on affoibliroit 1’arbre,

& quo’on avanceroit sa vieillesse.

“Cette seve pour etre sucree demande des circonstances

singulieres, qu’on ne devineroit pas, & que M. Sarrazin a

remarquees par ses experiences. 1°. It faut que dans le

temps qu’on la tire, le pied de l’arbre soit couvert de

neige, & il y en faudroit apporter, s’il n’y en avait pas.

2°. Il faut qu’ensuite cette neige soit fondue par le soleil,

& non par un air doux. 3°. Il faut qu’il ait gele la nuit

precedente. Cette espece de manipulation, dont la nature

se sert pour faire le sucre d’Erable, ressemble assez a

quelques operations delicates de Chymie, ou l’on fait des

choses qui paroissent opposees, ou celles qui paroissent le

plus semblables ne sont pas equivalentes pour 1’effet.

“Encore une remarque curieuse de M. Sarrazin, c’est

que le seve de tel Erable qui ne sera point bonne a faire

du sucre, le deviendra une demi-heure, ou tout au plus

une heure apres que de la neige, dont on aura couvert

le pied de l’arbre, aura commence a fondre. Cette neige

s’est done portee dans les tuyaux de 1’Erable, & y a

opere avec une grande vitesse.”

91. Bartram, John 1743

Observations on the Inhabitants, Climate, Soil, Rivers,

Productions, Animals, and other Matter Worthy of Notice

in his Travels from Pensilvania to Onondago, Oswego and

the Lake Ontario. London, 1751, (a) p. 28, (b) p. 39,

(c) p. 71.

a. (Sugar maple in Pennsylvania)

“The 15th, we set out a N.E. course, and passed by

very thick and tall timber of beach, chestnut, linden ash,

great magnolia, sugar-birch, sugar-maple, poplar, ...”

b. (Sugar maples along the Susquehanah river)

“ ... we descended easily for several miles, over good

land producing sugar-maples many of which the Indians

had tapped to make sugar of the sap, ...”

Schuette and Ihde — Maple Sugar

101

c. (Parched corn and maple sugar)

“ . . . They take the corn and parch it in hot ashes,

till it becomes brown, then clean it, pound it in a mortar

and sift it, this powder is mixt with sugar.”

92. Verwyst, Chrysoston 1744

Historic sites on Chequamegon Bay. Wisconsin Hist.

Coll., 13, 429 (1895).

(Maple sugar made by Indians along the

Swamp River)

“Mashki-Sibi (Swamp River, misnamed Bad River) :

About two miles up this river are pictured rocks, now

mostly covered with earth, on which in former times

Indians engraved in the soft stone the images of their

dreams, or the likenesses of their tutelary manitous. Along

this river are many maple groves, where from time im¬

memorial they have made maple-sugar.”

93. Kalm, Peter 1749

Travels into North America. J. R. Forster, trans.

London, 1772, Vol. II, 2 ed., p. 411.

(Sugar from the juice of the maples and the

sugar-birch)

They boil a good deal of sugar in Canada of the juice

running out of the incisions in the sugar-maple, the red

maple, and the sugar-birch; but that of the first tree

is most commonly made use of. The way of preparing

it has been more minutely described by me in the Memoirs

of the Royal Swedish Academy of Sciences.

94. B., J. C. 1751-1761

Voyage au Canada, dans le nord de TAmerique Sep-

tentrionale, fait depuis Tan 1751 a 1761; H. R. Casgrain,

Editor. Quebec, 1887; Travels in New France, S. K.

Stevens, D. H. Kent, Emma F. Woods, Editors. Penn¬

sylvania Historical Survey, Harrisburg, 1941, p. 93.

(A most remarkable tree in the Ohio Country)

“The maple is the most remarkable of all these trees,

because every year, in February and March, there exudes

from this tree an abundant flow of a delicious, sweet and

clear liquid which is fragrant and very wholesome. The

tree will die, however, if it is used too often.

102 Wisconsin Academy of Sciences , Arts and Letters

“Maple sap can be drawn from the same tree for five

or six consecutive days, if care is taken to make new

grooves every day always on the side toward the noon¬

day sun. This must be, too, when it has been cold the

night before, and when there is bright sunshine without

a cold, fierce wind. It can be determined that the tree

contains no more sap when the sap appears whitish and

runs slowly.

“When it is at that stage, vinegar or a drink like cider

can be made from it, if you go on extracting, but it still

must be boiled down into sugar.

“Maple sap must be cooked for two full hours to

make syrup, and two more hours to make sugar, which

is always brown. It is very good for the lungs and never

causes heartburn. Maple sugar is made into small cakes

like chocolate so that it can more easily be carried on

trips. It keeps a long time if dry, otherwise becoming

moldy, spoiling because of dampness. Maple trees usual¬

ly have large growths on them, which are cut and dried

in the sun, making a sort of touchwood which Canadians

call tondre .

“The plane-tree, the wild cherry, the ash, and the

walnut, also produce sap which yields sugar. But as the

flow is much less and the sugar not so good, it is almost

never made. Maple sugar, therefore, is most used in

Canada, as much as white sugar is used. . . . Maple

sugar is very easily digested.”

95. Barney, George 1755

The history of the Town of Swanton in Franklin

County. Vermont Historical Gazetteer, Vol. IV, p. 957

(1882).

(Indians made maple sugar at Missisqoi)

“ . . . From Dr. Belknap's History of New Hampshire

we learn that in 1755, the Indians in the vicinity of Mis¬

sisqoi were in the habit of tapping the maples in spring,

and making sugar. This perhaps was no more than they

had been accustomed to do for many years. ...”

96. Dodge, J. R. 1755

Red Men of the Ohio Valley. Springfield, 0., 1860,

(a) p. 80, (b) p. 85.

103

Schuette and llnde — Maple Sugar

a. Their Character and Customs

“Impelled by hunger, a supply of wild fowl, fat rac¬

coons, deer and bears, was secured; at the proper season,

a quantity of maple sugar made ; then, with hominy, which

became a luxury with a dressing of bear’s fat and sugar,

the feast would be prolonged, day after day, till all was

gone; ...”

b. Narrative of Colonel James Smith

“This Wyandot encampment consisted of eight hunt¬

ers, and thirteen squaws, boys and children. ... In

February, they commenced sugar making. First, the

squaws, finding an elm that would strip at this season,

cut it down, and with a crooked stick, broad and sharp

at the end, took the bark from the tree, and of this bark

curiously wrought vessels holding about two gallons each,

making more than a hundred in number. In the sugar-

tree, they cut a sloping notch, at the lower extremity of

which they struck a tomahawk, and, in the cut, drove a

long chip to convey the sap to the vessels. They notched

only the trees of two feet or more in diameter, so plenty

and large were they. They had bark vessels of four gal¬

lons each, two brass kettles of fifteen gallons each, and,

as they could not boil the sap as fast as collected, they

had large vessels of bark, holding about one hundred

gallons each.

“Their mode of using sugar was by putting it on

bear’s fat, until the fat was almost as sweet as the sugar

itself, and in this was dipped the roasted venison — by no

means an unpalatable morsel.”

97. Gautier, Jean Francois 1755

Histoire du sucre d’erable. Acad. Roy . Sciences, Mem.

math . phys., 2, 378-392 (1755).

(Adulterated maple sugar)

Described is not only the preparation of syrup and

sugar in Canada from the sap of the maple ( Acer coma-

dense , folio tridenato) but also experiments in support of

the observations that the optimum meteorological condi¬

tions for the flow of sap are warm, sunny days with the

wind blowing from the southwest, on which side of its

trunk the tree should be tapped.

104 Wisconsin Academy of Sciences, Arts and Letters

The addition of wheat flour to the syrup or sugar is a

frequently practiced form of adulteration. Because the

syrup prepared from the sap obtained at the end of the

season is of poor quality, it is often flavored with an

extract of maiden hair fern.

Maple sugar is used in the preparation of foods in

the home; as a medicine it is deemed to be pectoral and

emollient.

98. Alden, Timothy 1756

An Account of the Captivity of Hugh Gibson among

the Delaware Indians of the Big Beaver and the Muskin¬

gum, from the latter part of July 1756, to the begin¬

ning of April, 1759. Boston, 1887. Coll. Mass. Hist . Soc.,

[3] 6, 147 (1837).

(Indian Sugar Camp)

“Gibson and a little boy, of twelve years of age, went

on a hunting expedition, were absent three days, killed

two turkeys, and returned; but Bisquittam, whether sus¬

pecting the plan or not is unknown, was still at the place.

He, with the little boy, again took a tour into the woods.

They reached an Indian sugar camp the first evening, stole a

horse and a bag of corn, rode several miles to a cranberry

swamp, tarried there seven days, parched and ate their

corn, threw away their bag, killed one turkey, and returned

to the sugar camp.,,

99. Henry, Alexander 17604-

Travel and Adventures in Canada and the Indian Ter¬

ritories between the Years 1760-1765. New York, 1809,

(a) p. 68, (b) p. 216, (c) p. 217.

a. (Sugar-making near the Sault de Sainte-Marie)

“The lands, between the Bay of Boutchitaouy and the

Sault, are generally swampy, excepting so much of them

as compose a ridge, or mountain, running east and west,

and which is rocky, and covered with the rock or sugar

maple, or sugar-wood ( Acer saccharinum) . The season

for making maple-sugar was now at hand; and, shortly

after my arrival at the Sault, I removed, with some other

inhabitants, to the place at which we were to perform

the manufacture. . . .

Schuette and Ihde~Maple Sugar

105

“The next day was employed in gathering the bark of

white birch-trees, with which to make vessels to catch

the wine or sap. The trees were now cut or tapped,

and spouts or ducts introduced into the wound. The bark

vessels were placed under the ducts; and, as they filled,

the liquor was taken out in buckets and conveyed into

reservoirs or vats of moose-skin, each vat containing a

hundred gallons. From these, we supplied the boilers, of

which we had twelve, of from twelve to twenty gallons

each, with fires constantly under them, day and night.

While the women collected the sap, boiled it, and com¬

pleted the sugar, the men were not less busy in cutting

wood, making fires, and in hunting and fishing, in part of

our supply of food.

“The earlier part of the spring is that best adapted to

making maple-sugar. The sap runs only in the day; and

it will not run unless there has been a frost the night

before. 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.

“ ... we hunted and fished, yet sugar was our prin¬

cipal food, during the whole month of April. I have

known Indians to live wholly upon the same, and become

fat.”

b. (Sugar maples at Michipicoten)

“The country, immediately contiguous to my wintering-

ground, was mountainous in every direction; ... On

the summits of some of the mountains there were sugar-

maple trees; but, with these exceptions, the uplands had

no other growth than spruce-firs and pines, nor the low¬

lands than birch and poplar.”

c. Maple-sugar making

“In the beginning of April, I prepared to make maple-

sugar, building for this purpose a house, in a hollow dug

out of the snow, the house was seven feet high, but yet

was lower than the snow.

“On the twenty-fourth, I began my manufacture. On

the twenty-eighth, the lands below were covered with a

thick fog. All was calm, and from the top of the mountain

106 Wisconsin Academy of Sciences , Arts and Letters

not a cloud was to be discovered in the horizon. Descend¬

ing the next day, I found half a foot of new-fallen snow,

and learned that it had blown hard in the valleys the day

before; so that I perceived I had been making sugar in

a region above the clouds.

“Sugar-making continued till the twelfth of May. On

the mountain, we eat nothing but sugar, during the whole

period. Each man consumed a pound a day, desired no

other food, and was visibly nourished by it.”

100. Anon. 1765

The Annual Register, or a View of the History, Pol¬

itics, and Literature for the Year 1765. London, 1778, 2

ed., Yol. VIII, Chronicles, p. 141.

(New method of obtaining sugar and melasses)

“A very singular method of obtaining sugar and me¬

lasses has been lately introduced in New England, espe¬

cially at a place called Bernardston, almost twenty miles

from Athol; and as the vegetable, from which that valu¬

able article may be obtained by this new method, grows

in the coldest climates, it promises great advantages to

mankind, especially in those countries, which, like New Eng¬

land, are already plentifully stocked with it by the hand of

nature. This vegetable is no other than the maple-tree.

The process in Bernardston is as follows. Having chosen

out a large tree, suitable for the purpose, they with an

axe box it, much after the same manner, that they box

that kind of fir, which produces turpentine. This being

done, they prepare a kind of trough, extending from the

trunk of the tree on each side, in order to retain the sap

as it runs down. By this means they have obtained up¬

wards of thirty gallons from one tree in a day; which,

being treated like the syrup proceeding from the sugar

cane, produces a sugar equal in fineness of grain to the

Jamaica sugar, and as pleasant to the taste; and the mak¬

ers insist that it is a medicinal, and very popular to give

children for the chin-cough, at this time very prevalent in

New England. This sugar produces melasses, or treacle,

very little, if anything, inferior to West Indian melasses.

Of this sugar, above 600 lb. was made by one man during

the last season, that is, from February last to April last

Schuette and Ihde — Maple Sugar

107

inclusive; and several hundred weight of it were in July

last brought for sale to Boston in New England, from

various towns situated on the eastern and western parts

of that province.”

101. Pond, Mrs. Nathan G. ca. 1765

Journal of “Sir” Peter Pond. The Conn. Magazine,

10, (a) 244, (b) 245 (1906).

a. (Maple sugar traded at Mackinac)

“ . . . Hear I Met with a Grate meney Hundred Peo¬

ple of all Denominations . . . Sum trading with the tribes

that Came a Grate Distans with thare furs, Skins & Mapel

Suga &c to Market.”

b. (Maple sugar made near Mackinac)

“ . . . Most of the frenchmens wives are white women.

In the spring they made a Grate Quantity of Maple Suga

for the youse of thare families & for sale som of them.”

102. Rogers, Robert 1765

A Concise Account of North America. London, 1765,

p. 251.

(Loaf sugar from maple sap)

“The Indians, in the months of February and March,

extract the juice from the maple-tree; which is wholesome

and delicious to the palate. The way they extract it is by

cutting a notch in the body of the tree, and by means of

a piece of wood or quill, convey the juice from the tree

to a vessel placed to receive it. The same tree may be

tapped for several years successively. The liquor is as

clear as spring-water, and is very refreshing. It is ac¬

counted a very good pectoral; and was never known to

hurt any one, tho' he drank ever so freely of it. The liquor

will not freeze; but, when kept any time, becomes excel¬

lent vinegar. The Indians, by boiling it, make from it a

kind of sugar, but is milder, and answers all the ends of

sugar for sweetening; and, no doubt, was it properly

manufactured, might be rendered equal to that extracted

from sugar-cane. A manufactory is begun in the Prov¬

ince of New York, near South Bay; which, I am told,

answers very well; and produces considerable quantities

of powder and loaf sugar.”

108 Wisconsin Academy of Sciences, Arts and Letters

103. Grignon, Augustin 1766-1816

Seventy-two years’ recollections of Wisconsin. Wis¬

consin Hist. Coll., 3, 255 (1857).

(Maple sugar used in fur trade at Green Bay)

‘The early commerce of the country deserves a pass¬

ing notice. . . . There was some considerable quantity of

deer’s tallow, saved by the Indians and sold to the traders,

taken to Mackinaw, and some maple sugar ; . . . But as

there was much sugar manufactured around Mackinaw,

not much was sent there to market ; the Indians made

large quantities as far back as I can remember. To the

traders passing into the Indian country, cattle for beef

were sold, sugar and tallow, potatoes and other vegeta¬

bles. ...”

104 Anon 1767

A New Collection of Voyages, Discoveries and Trav¬

els, etc. London, 1767, Vol. II, (a) p. 138, (b) p. 179.

a. (Beverage of the Souties or Attawawas)

“They, as yet, make very little use of spiritous liquors,

nor do they manufacture any kind of drink, except the

juice of the maple-tree, of which they likewise make su¬

gar.”

b. (Properties of maple sap)

“The Indians, in the months of February and March,

extract the juice from the maple-tree, which is whole¬

some and delicious to the palate. The way they extract it

is by cutting a notch in the body of the tree, and, by

means of a piece of wood or quill, convey the juice from

the tree to a vessel placed to receive it. The liquor is as

clear as spring-water, and is very refreshing. It is ac¬

counted a very good pectoral, and was never known to

hurt any one, though he drank ever so freely of it. The

liquor will not freeze, but, when kept any time, becomes

excellent vinegar. The Indians, by boiling it, make from it

a kind of sugar, which has a taste very much like honey,

but is milder : A manufactory of this kind of sugar is

begun in the province of New York, near South Bay,

which is said to answer very well; and produces consid¬

erable quantities of powder and loaf-sugar.

Schuette and Ihde— Maple Sugar

109

105. Crevecoeur, St. John de 1770-1774

Henri L. Bourdin, Ralph H. Gabriel and Stanley T.

Williams, Editors. Sketches of Eighteenth Century Amer¬

ica. More “Letters from an American Farmer,” by St.

John de Crevecoeur. New Haven, 1925, p. 98.

( Careful “ bleeding ” prolongs life of the tree.)

“In clearing his farm my father very prudently saved

all the maple trees he found, which fortunately are all

placed together in the middle of our woodland; and by

his particular caution in bleeding them, they yield sap as

plentifully as ever. The common method is to notch them

with an axe. This operation, after a few years, destroys

the tree entirely. That which my father followed is much

easier, and gives these trees wounds which are almost

imperceptible. The best time to make this sugar is be¬

tween the months of March and April, according to the

season. There must be snow on the ground, and it must

freeze at night and thaw in the day. These three circum¬

stances are absolutely requisite to make the sap run in

abundance. But as my trees are but a little way from my

house, I now and then go out and try them, and, as soon

as the time is come, then I bring all my hands, and we go

to work. Nothing can be simpler than this operation. I

previously provide myself with as many trays as I have

trees. These I bore with a large gimlet. I then fix a spile

made of elder through which the sap runs into the trays.

From them it is carried into the boiler which is already fixed

on the fire. If the evaporation is slow, we are provided with

barrels to receive it. In a little time it becomes of the con¬

sistency of syrup. Then it is put into another vessel and

made to granulate. When in that state we cast it into little

moulds made according to the fancy of the farmer. Some

persons know how to purify it, and I am told that there

are some people at Montreal who excel in this branch. For

my part, I am perfectly well satisfied with the colour and

taste which Nature has given it. When the trees have

ceased to run we stop the holes with pegs made of the

same wood. We cut them close to the bark, and in a little

time the cicatrice becomes imperceptible. By these simple

means our trees will afford sugar for a long time, nor

110 Wisconsin Academy of Sciences , Arts and Letters

have I ever observed that it impaired their growth in the

least degree. They will run every year, according to the

seasons, from six to fifteen days until their buds fill. They

do not yield every year the same quantity, but as I regu¬

larly bleed two hundred trees, which are all I have, I have

commonly received six barrels of sap in twenty-four hours

which have yielded me from twelve to eighteen (pounds

of sugar).

“Thus without the assistance of the West Indes, by the

help of my trees and of my bees, we yearly procure the

sweetening we want, and it is not a small quantity, you

know, that satisfies the wants of a tolerable American

family.”

106. Le Page du Pratz 1774

The History of Louisiana or of the Western Parts of

Virginia and Carolina. T. Becket, translator. London,

1774, new ed., p. 240.

(An excellent stomachic)

“The maple grows upon declevities in cold climates,

and is much more plentiful in the northern than southern

parts of the colony. By boring it they draw from it a

sweet syrup which I have drunk of, and which they allege

is an excellent stomachic.”

107. Buchanan, James ca. 1775

Sketches of the History, Manners, and Customs of the

North American Indians. New York, 1824, Vol. I, p. 163.

(An episode at an Indian sugar-camp)

“ . . . One hope, however, still remained. The sugar¬

making season was at hand, and they were shortly to re¬

move to their sugar camps, where he flattered himself

his wife would not be followed by the disturber of his

peace, whose residence was about ten miles from thence.

But this hope was of short duration. They had hardly

been a fortnight in their new habitation, when, as he

returned one day from a morning's hunt, he found the un¬

welcome visiter at his home, in close conversation with

his faithless wife. This last stroke was more than he

could bear; without saying a single word, he took off a

large cake of his sugar, and with it came to my house,

Schuette and Ihde — Maple Sugar

111

which was at the distance of eight miles from his tempo¬

rary residence.”

108. Ansbury, Thomas 1776

Travels through the Interior Parts of America; in a

Series of Letters. London, 1791, new ed., Vol. I, p. 80.

(Maple sugar used as a pectoral)

Quebec, November 5th 1776

“My Dear Friend,

• • •

“The maple tree yields in great quantities a liquor

which is cool and refreshing, with an agreeable flavor.

The Canadians make a sugar of it, a very good pectoral,

and used for coughs. There are many trees that yield a

liquor they can convert into sugar, but none in such

abundance as the maple. You will, no doubt, be surprised

to find, in Canada, what Virgil predicted of the Golden

Age, Et durae quercus subabunt roscida mella.”

109. Askin, John 1778

Fur-trade on the upper lakes - 1778-1815. Wiscon¬

sin Hist. Coll, 19, p. 248 (1910) .

(Letter to Benjamin Frobisher mentions sugar)

“St. Cir arrived last night. I have delivered him the

Canoes, all your Corn, Sugar, Gum, Bark & Watap now

remaining here shall be delivered him to Day, all the rum

coming up in the Canoes he shall also have. ...”

110. Broadhead, Daniel 1780

Letters from Col. Daniel Broadhead. Pennsylvania

Archives, Vol. XII, p. 212 (1856).

(Murders at sugar camp)

Fort Pitt, March 8th, 1780

“Dear General,

The savages have already begun their hostilities — last

Sunday they killed five men at a Sugar Camp on Racoon

Creek, Youghaghany County, and took prisoners three

girls and three lads. It is generally conjectured that the

Delawares perpetrated this Murder, but it is possible that

it may have been Done by other Indians. . . .

Most Obed’t Serv’t.

Daniel Broadhead

112 Wisconsin Academy of Sciences y Arts and Letters

Directed

To His Excellency General Washington

By Mr. D. Duncan”

111. Alexander, M.T.C. 1784

Vermont Historical Gazetteer. Burlington, 1876, Vol.

I, (a) p. 313, col. 1; (b) col. 2

a. Danville - to 1860

In recalling the activities of Capt. Charles Sias, a pio¬

neer settler, the author states:

“ ... In three days more the effects were all removed,

and the lone family began their hard labors upon the

wilderness. They commenced by tapping the maples,

which stood thick around them in the most beautiful

groves, affording them sugar in abundance, and supplied,

in a great degree, the lack of other food.”

b. (Maple sugar an emergency food)

A too rapid increase in population caused a scarcity

of provisions in 1789. In describing the conditions exist¬

ing at that time, the author states:

“The sufferings of that time were very severe. Maple'

sugar formed the chief article of food. Like the manna

of the ancient Hebrews, it was really a providence in the

time of hunger and famine. No doubt, those stern old

fathers blessed the forest trees that gave them food and

life.”

112. Brissot de Warville, J. P. 1788

Nouveau Voyage dans les Etats-Unis de L’Amerique

Septentrionale, fait en 1788. Paris, 1791, Vol. II, p. 41 ; New

Travels in the United States of America, including the

Commerce of America. London, 1792, p. 301.

On replacing the Sugar of the Cane by the

Sugar of the Maple

“On this continent, my friend, so polluted and tormented

with slavery, Providence has placed two powerful and in¬

fallible means of destroying this evil. The means are, the

societies of which we have been speaking, and the sugar-

maple.

“Of all vegetables containing sugar, this maple, after

the sugar-cane, contains the greatest quantity. It grows

Schuette and Ihde— Maple Sugar 113

naturally in the United States, and may be propagated

with great facility. All America seems covered with it,

from Canada to Virginia; it becomes more rare at the

southward, on the east of the mountains; but it is found

in abundance in the back country.

“Such is the beneficent tree which has, for a long

time, recompensed the happy colonists, whose position de¬

prived them of the delicate sugar of our islands.

“They have till lately contented themselves with be¬

stowing very little labour on the manufacture, only bring¬

ing it to a state of common coarse sugar; but since the

Quakers have discerned in this production, the means of

destroying slavery, they have felt the necessity of carry¬

ing it to perfection; and success has crowned their en¬

deavors.

“You know, my friend, all the difficulties attending

the cultivation of the cane. It is a tender plant, it has

many enemies, and requires constant care and labour to

defend it from numerous accidents: add to these, the

painful efforts that the preparation and manufacture

costs to the wretched Africans; and, on comparing these

to the advantages of the maple, you will be convinced,

by a new argument, that much pains are often taken to

commit unprofitable crimes. The maple is produced by

nature ; the sap to be extracted, requires no prepara¬

tory labour; it runs in February and March, a season

unsuitable for other rural operations. Each tree, with¬

out injury to itself, gives twelve or fifteen gallons, which

will produce at least five pounds of sugar. A man aided

by four children, may easily, during four weeks running

of the sap, make fifteen hundred pounds of sugar.

“Advantages, like these, have not failed to excite the

attention of the friends of humanity; so that, besides the

societies formed for the abolition of slavery, another is

formed, whose express object is to perfect this valuable

production.

“Whenever these shall form from North to South a

firm coalition, an ardent emulation to multiply the prod¬

uce of this divine tree, and especially when it shall be

deemed an impiety to destroy it, not only America may

supply herself, but she may fill the markets of Europe

114 Wisconsin Academy of Sciences, Arts and Letters

with a sugar, the low price of which will ruin the sale

of that of the islands — a produce washed with the tears

and blood of slaves.

“What an astonishing effect it would produce, to nat¬

uralize this tree through all Europe! In France we might

plant them at twenty feet distance, in a kind of orchard,

which would at the same time produce pasture, fruits,

and other vegetables. In this manner an acre would

contain 140 trees, which, even when young, would pro¬

duce three pounds of sugar a-year. This would give 420

pounds the acre, which, at threepence sterling the pound,

and deducting one half for the labour, would yield annually

52 £. 6s. sterling, clear profit; besides other productions,

which these trees would not impede. . . .

“Thus we should obtain a profitable production in

Europe, and diminish so many strokes of the whip, which

our luxury draws upon the blacks. Why is it, that, in our

capital where the delicacy of sentiment is sometimes equal

to that of sensation, no societies are formed, whose ob¬

ject should be to sweeten their coffee with a sugar not

embittered by the idea of the excessive tears, cruelties

and crimes, without which these productions have not been

hitherto procured ? — an idea which cannot fail to present

itself to the imagination of every humane and enlightened

man. Our devotees, our ignorant and inhuman priests,

who never fail to be great lovers of coffee and sugar,

would, by these means, be saved from the horrible part

which they take in the most enormous crime on which

the sun ever shown. In consuming these articles, do they

not encourage those whose guilt is more direct in the

operation of producing them? and yet, with what cold¬

ness, with what culpable indifference, do these pious men

look upon our Society of the Friends of the Blacks.”

113. Anon. 1792

A plan for moderating the price of sugar. The Bee ,

or Literary Weekly Intelligencer , 7, 330.

.

(Sugar from the maple for Britain ?)

“The present extravagent price of sugar has attracted

the attention of every class of persons in this island, and

has brought forward many plans for remedying that

Schuette and Hide— Maple Sugar

115

evil, some of which will no doubt take effect at some

future period; but there is reason to suspect, that the

nation must submit to the hardship for a good while,

before things can be brought to bear.

“Among the first plans that was suggested for this

purpose, was that of manufacturing sugar from the maple

tree, in America. It has been long known, that the juice

of one kind of maple, common in most of the American

states, can afford a grained sugar, without any other

process than that of evaporating the watery parts by

boiling; but the quantity of water that requires to be

dissipated, renders that process so tedious and expensive,

in a country where labour is very high, as gives reasons

to fear the assistance that can be derived from thence

will be but very inconsiderable.”

114. Biggs, Benj. 1793

Calendar of Virginia State Papers. Sherwin McRae,

Editor. Richmond, 1886, Vol. VI, p. 307.

(Sugar making near Marietta, Ohio)

“I just Received authentick Intelligence of the capture

of Major Goodall by the Savage, which happened near

Bellspray on the Ohio, a small distance below Muskingum.

About seven days previous to the capture a number of

Kittles were taken from the Sugar Camp, within a small

distance of Mariatta.”

115. Allen, William 1794

The History of Norridgewock. Norridgewock, 1849,

p. 99.

(Rum and maple sugar)

“In the summer of 1794, the meeting house was erect¬

ed, and among other things preparatory to the raising,

it was ‘Voted to get one barrel of good W. I. Rum, and

one-hundred pounds of maple sugar, to be used at the

raising of the meeting house’. ”

116. Coxe, Tench 1794

A View of the United States of America, in a Series

of Papers, Written at Various Times, between the Years

1787 and 1794. Philadelphia, 1794, (a) p. 65, (b) p. 77,

(c) p. 453, (d) p. 455; Dublin, 1795, (a) p. 54, (b) p.

65, (c) p. 386, (d) p. 388.

116 Wisconsin Academy of Sciences , Arts and Letters

a. ( Sugar from the maple trees of Pennsylvania)

“ A new article is added to the list of our pro¬

ductions, which is a well-tested and wholesome sugar,

made of the maple tree. It has been proved, by many fair

and careful experiments, that it is in the power of a

substantial farmer, who has a family about him, easily

to make twelve hundred weight of this sugar every season,

without hiring any additional hands, or utensils, but those

which are necessary for his family, and farm use. The

time, in which it can be made, is from the middle of Feb¬

ruary to the end of March, when farmers in this country

have very little to do, as it is too early to plough or

dig. The price of sugar being lower here than in Europe,

this article may be reckoned at one hundred and fifty

dollars per annum, to every careful and skilful farmer,

who owns land bearing the sugar maple. Of these there

are some millions of acres in Pennsylvania and the ad¬

jacent states. It seems also highly probable, that this

valuable tree may be transplanted, and thus be obtained

by almost any farmer in the state ; and that men of prop¬

erty, who will purchase kettles, and hire hands for the

above short period, may make larger quantities.”

b. (The possible value of maple sugar in

certain parts of the United States)

“ . . . The easy and profitable practice of making su¬

gar from the sap or juice of the maple tree, had pre¬

vailed for many years in the northern and eastern states.

The facility and advantages of this pleasing branch of

husbandry, had attracted little attention in Pennsylvania,

though a few of its inhabitants were in the habit of manu¬

facturing small quantities of this kind of sugar. In the

year 1790, it became more generally known to the Penn¬

sylvanians that their brethern in the eastern and northern

parts of the union, had long made considerable quantities,

with their family utensils, and without the expense of

hiring assistance, that the same tree might be carefully

tapped without injury for many successive years; that the

process was simple and very easy, and only required to

be carried on between the middle of February and the

end of March when the farmer has little to do, and that

Schuette and Ihde — Maple Sugar

117

a very large proportion of the unsettled lands of the

state abound with this valuable tree. The great and in¬

creasing dislike to negro slavery, and to the African trade

among the people of that state, occasioned this new

prospect of obtaining a sugar, not made by the unhappy

blacks, to be particularly interesting to them. The fol¬

lowing estimate, which was founded on the best materials

obtainable at that time, was published among other things

to elucidate the subject.”

c. A method of clearing a farm lot of new wood¬

land, easily practicable by persons having no more

money or provisions than are sufficient to provide

the food and clothing of their families, during the

first year of their settlement.

“ ... If he has sugar maple trees on his land, he

may also obtain money, by making sugar in February and

March, and felling or bartering, it for cash, or goods to

be laid out in like manner, in hiring hands the next sea¬

son. If money is scarce in a new settlement, and he bar¬

ters pot ash or maple sugar, for strong trowsers, shirts,

hats or jackets, he will find it easy to procure laborers

for such necessaries.,,

d. (The valuable maple)

“The United States have been brought, by slow de¬

grees, to their present knowledge of the value of their

wood and timber. The value of the maple sugar tree is

not yet universally known. . . . ”

117. Wansey, Henry 1794

An Excursion to the United States of America. Salis¬

bury, 1798, 2 ed., (a) p. 48, (b) p. 47, (c) p. 47, (d)

p. 262.

a. (Maple sugar served in Hartford, Conn.)

“At Frederick Bull's tavern, where I lodged, we had

excellent provisions: beef, mutton, and veal, as good as in

England; tea and coffee of the best kind; three sorts of

sugar brought always to the table; — the muscovado, the

fine lump sugar, and the maple; from the novelty of it,

I preferred the last, though I could not find much dif¬

ference in the taste of it."

118 Wisconsin Academy of Sciences , Arts and Letters

b. (England forbids importation of maple sugar)

“At breakfast I was offered by one of the passengers

five hundred weight of it, for fourpence halfpenny sterling

per pound, but it is contrary to the laws of England to

import it.”

c. (Maple groves in Dunham toivnship, Conn.)

“Under many of the maple trees, I observed many of

the wooden troughs remaining, and the taps still in the

trunks, although the sap season had been over about six

weeks, being only while the sap is rising. A frosty night

always makes a plentiful distillation next morning. A

passenger told me that a barrel of juice made six quarts

of molasses, which produces ten or eleven pounds of

sugar.”

d. (Granulated maple sugar)

“The sugar maple, is a tree that I should suppose

would grow in this country as well as in many parts of

Europe. In Connecticut, it is exposed to as severe win¬

ters as any in this island. I have a tree in my garden,

seven feet high, that has stood the severe winter of 1794.

The chief thing to attend to, is to see it planted in good

rich soil. Those settlers in America, who clear the lands,

always begin with cutting down the sugar maples, because

they are generally found on the richest and best land. — -

This is one reason why America will not be sufficient to

supply its own sugar.

“I was given the following as the method in which

they make it: Draw off the sap into wooden vessels, by

wooden taps fixed in the bark, seven feet from the ground.

Boil it away next day;— provide three kettles of different

sizes — say, of fifty, sixty, and seventy gallons; boil it first

in the large kettle, adding as much lime as will make the

liquor granulate; as it boils, take off the scum, encreasing

the heat, till it evaporates to sixty gallons; then strain

it through a woolen cloth into the sixty gallon kettle.

This must be boiled and skimmed in the same manner,

till it is reduced to fifty, and then be strained into the

fifty gallon kettle. And each kettle must be continued in

succession, till you have boiled your whole quantity — say

two hundred gallons.

Schuette and Ihde — Maple Sugar

119

“When it is boiled enough, which is known by its be¬

coming ropy between the finger and thumb, it is turned

out into a wooden cooler, and stirred with a kind of wood¬

en paddle, till it granulates ; and then it is put into earthen

moulds, in the same manner as the West-Indian planters

practice.”

118. La Rochefoucault-Liancourt, Duke de 1795

Travels through the United States of North America,

the Country of the Iroquois, and Upper Canada in the

Years 1795, 1796, and 1797. London, 1799, Vol. I, (a) p.

79, (b) p. 96, (c) p. 107, (d) p. 125, (e) p. 153, (f) p.

283.

a . (Maple sugar at Fishing Creek, Penn.)

“ . . . This was the first place, where we used maple

sugar, which we found excellent. Abraham Miller sells

yearly about five or six barrels of this sugar. He buys it at

thirteen pence a pound, and sells it at fifteen; the brown

moist sugar of the colonies he sells at fourteen pence.”

b. (Maple Sugar made at Asylum, Penn.)

“ . . . Maple-sugar is made here in great abundance.

Each tree is computed to yield, upon an average, from

two pounds and half to three a year. Melasses and vinegar

are also prepared here. I have seen Messrs. De Vilaine

and Dandelot make sugar in this place, which much sur¬

passes any of the same kind, which has hitherto come

under my observation.”

c. (Price of maple sugar at Painted Post, N. Y.)

“ . . . There are however few sugar-maple trees. The

price of this sugar at the beginning of last spring was

one shilling per pound.”

d. (Maple sugar production at Genessee)

“The whole country abounds in sugar-maple trees (Acer

saccharinum, Lin. called by the Indians Ozeketa .-Trans.),

and very considerable quantities of this sugar are made

here. The following is the substance of the information,

which we were able to procure on this head:

“1. The medium produce of a tree, standing in the

midst of a wood, is three pounds of sugar.

120 Wisconsin Academy of Sciences, Arts and Letters

“2. The average produce of trees, standing on ground

which has been cleared of all other wood, is from five to

seven pounds per tree.

“3. A barrel of the first juice, which comes from the

maple-tree, will yield seven pounds of sugar, if the tree

stand single, and four, if it stand in the midst of other

wood. This sugar is sold at one shilling per pound.

“4. A barrel of the second juice will yield three gal¬

lons and a half of treacle.

“5. Four or five barrels of the third juice will yield

one barrel of a good and pleasant vinegar.

“6. The vinegar is found to be better, in proportion

as it is more concentrated. This is the case with Robin¬

son'S vinegar, who, from ten barrels of the third juice,

brews but one barrel of vinegar.

“7. To clarify the vinegar, it must be boiled with

leaven.

“8. The third juice, which is not used for vinegar,

yields cyder of an excellent flavour, when mixed with

an equal quantity of water.

“9. The longer the first juice is boiled, the better and

finer the sugar will become.

“10. In order that the trees may continue productive,

they require to be tapped with extraordinary care; i.e. the

fissures must be neither too deep, nor too wide, so that no

water may settle in them, after the juice is extracted, and

that the wood may close again in the space of a twelve-

month.

“11. During the time the juice is flowing out, which

lasts about six weeks, and generally begins on the 1st of

February, all the days on which it freezes or rains are

lost, so that the number of days on which the business

can be pursued to advantage is frequently, from these

circumstances, much diminished.

“12. Maple sugar, however, is already obtained in

sufficient quantities, to form a respectable article of trade,

as during the above time two persons can frequently make

from five to six hundred pounds of it, and this quantity

will be increased in proportion to the number of workmen

Schuette and Ihde — Maple Sugar

121

employed. As the maple-tree, wherever it goes, multiplies

with astonishing rapidity, we found, almost everywhere

on our journey, no want of excellent sugar. At Robin¬

son’s it was better and finer than we had met with any

where else; although in general it was not so white here

as at Asylum, where Messrs, de Villaine and D’Andlau

refine it with the yolks of eggs. ...”

e. (Maple sugar production at Genessee Flats)

“ . . . Maple-sugar, of which great quantities are usu¬

ally obtained in this neighbourhood, has not answered this

year, from the uncommon wetness of the season. It is

sold for one schilling a pound.”

/. (Production neglected at Kingston , Canada)

“ . . . The farmers make but little maple-sugar, though

the woods abound with the trees, from which it is pro¬

duced. The Indians import about two or three thousand

pounds, and sell it to the retail traders for one shilling

a pound. Maple-sugar is prepared in much larger quan¬

tities in Lower Canada. The Canadians eat it here on

bread, or make cakes of it, mixed up with flour of wheat,

or Indian corn. ...”

119. Weld, Isaac, Jr. 1795

Travels through the States of North America, and the

Provinces of Upper and Lower Canada, during the years

1795, 1796 and 1797. London, 1799, pp. 219-224.

Observations on the Manufacture of

Sugar from the Maple-tree

“The variety of trees found in the forests of Canada

is prodigious, . . . ; the sugar maple tree is also found in

almost every part of the country, a tree never seen but

upon good ground. There are two kinds of this very valu¬

able tree in Canada ; the one called the swamp maple, from

its being generally found upon low lands; the other, the

mountain or curled maple, from growing upon high dry

ground, and from the grain of the wood being very

beautifully variegated with little stripes and curls. The

former yields a much greater quantity of sap, in propor¬

tion to its size, than the other, but this sap does not

afford so much sugar as that of the curled maple. A

122 Wisconsin Academy of Sciences , Arts and Letters

pound of sugar is frequently procured from two or three

gallons of the sap of the curled maple, whereas no more

than the same quantity can be had from six or seven

gallons of that of the swamp. . . .

“The maple is the only sort of raw sugar made use of

in the country parts of Canada; it is very generally used

also by the inhabitants of towns, whither it is brought

for sale by the country people who attend the markets,

just the same as any other kind of country produce. The

most common form in which it is seen is in loaves or

thick round cakes, precisely as it comes from the vessel

where it is boiled down from the sap. These cakes are of

a very dark colour in general, and very hard; as they are

wanted they are scraped down with a knife, and when

thus reduced into powder, the sugar appears of a much

lighter cast and not unlike West Indian muscovada or

grained sugar. If the maple sugar be carefully boiled

with lime, whites of eggs, blood, or any of the other

articles usually employed for clarifying sugar, and prop¬

erly granulated, by the draining off of the melasses, it is

by no means inferior, either in the point of strength,

flavour, or appearance to the eye, to any West Indian

sugar whatsoever: simply boiled down into cakes with

milk or whites of eggs it is very pleasing to the taste. . . .”

120. Winterbotham, W. 1795

An Historical, Geographical, Commercial, and Philo¬

sophical view of the American United States, and of the

European Settlements in America and the West Indies.

London, 1795, Vol. Ill, (a) p. 498, (b) p. 500.

An extended report on maple sugar in America. The

first part deals with the occurrence of the maple tree in

New York and Pennsylvania. The tree is described. Fac¬

tors which influence the flow of sap are discussed along

with methods of concentrating the sap by freezing, spon¬

taneous evaporation and boiling. The possibility of large

scale manufacture is considered.

a. (Maple sugar is clean and not made by slaves.)

“The quality of this sugar is necessarily better than

that which is made in the West-Indies. It is prepared

in a season when not a single insect exists to feed upon

Schuette and Ihde — -Maple Sugar

123

it, or to mix its excrements with it, and before a particle

of dust or of the pollen of plants can float in the air. The

same observation can not be applied to the West-India

sugar. The insects and worms which prey upon it, and

of course mix with it, compose a page in the nomencla¬

ture of natural history. We shall say nothing of the hands

that are employed in making sugar in the West-Indies,

but that men who work for the exclusive benefit of others

are not under the same obligations to keep their persons

clean while they are employed in this work, that men,

women and children are, who work exclusively for the

benefit of themselves, and who have been educated in the

habits of cleanliness. The superior purity of the maple

sugar is farther proved by its leaving a less sediment

when dissolved in water than the West-India sugar.

“It has been supposed that the maple sugar is inferior

to the West-India sugar in strength. The experiments

which led to this opinion we suspect to have been inaccu¬

rate, or have been made with maple sugar prepared in

a slovenly manner. Dr. Rush examined equal quantities

by weight of both the grained and loaf sugar, in hyson

tea, and in coffee, made in every respect equal by the

minutest circumstances that could affect the quality of

taste of either of them, and could perceive no inferiority

in the strength of the maple sugar. The liquors which

decided this question were examined at the same time

by Alexander Hamilton, Esq. secretary of the treasury

of the United States, Mr. Henry Drinker, and several

ladies, who all concurred in the above opinion.”

h. (Maple sugar in relation to alcoholic liquor)

“ . . . The sap of the maple is moreover capable of

affording a spirit, but we hope this precious juice will

never be prostituted by American citizens to this ignoble

purpose. Should the use of sugar diet become more gen¬

eral in America, it may tend to lessen the inclination or

supposed necessity for spirits, for a relish for sugar in

diet is seldom accompanied by a love of strong drink. It

is the sugar which is mixed with tea which makes it so

generally disagreeable to drunkards.”

124 Wisconsin Academy of Sciences, Arts and Letters

121. Thwaites, R. G. 1797-98

Narrative of Andrew J. Vieau, Sr. Wisconsin Hist.

Coll., 11, (a) 228, (b) 231 (1898).

a. (Maple sugar at Milwaukee, Wisconsin)

“ . . . Each spring, after packing up the winter’s

peltries and buying all the maple sugar available from

the Indians, father would start out with his family and

goods on his return to Mackinaw. . . . Upon his return

down the lake, father would stop at his various jack-knife

posts and collect their furs and maple sugar. ...”

b. (A venture in maple sugar)

“In the spring of 1839, I closed up my post, bought a

lot of sugar from the Indians, loaded up a boat with the

sugar and furs that I had collected and went up to Mil¬

waukee, where I disposed of my venture, having had an

excellent winter’s trade.”

122. Adams, George ca. 1798

History of Enosburgh, Franklin County. Vermont

Historical Gazetteer, Vol. II, p. 135 (1871).

(Wooden faucets for drawing off sap)

“Sugar is something of an article of revenue. Since

the high prices occasioned by the war, great improvements

have been made in the process of manufacture. Two con¬

siderable sugar orchards are in use: one by Virgil Bogue,

and one by James Kidder, where grain was once grown.

Among all the improvements in this business, whether in

theory or practice, the most novel is in that of tapping,

proposed by one of our first settlers, Isaac B. Farrar, who

settled on the lot on which V. Bogue lives. Mr. Farrar

was a son of Priest Farrar, of New Ipswich, N. H. — had

a liberal education, and doubtless thought it best to bring

his knowledge to bear on his business, and pursue a kind

of 'scientific farming.’ He brought with him a large quan¬

tity of wooden faucets. When inquired of what he de¬

signed these for, said 'he had formed a favorable opinion

of the manufacture of maple sugar; and, upon inquiry,

thought the method then pursued of tapping with an axe,

gouge and split spouts, must occasion great waste, as well

as hurry in gathering and boiling, when it run rapidly.’

Schuette and Ihde— Maple Sugar

125

Said his 'plan was to obviate both difficulties, by tapping

with an auger, and putting in a faucet; and when he

wanted sap, to draw a pailful, and take it leisurely/ He

afterwards moved to Fairfax, and established himself in

the pottery business. Whether his improved manner of

tapping was generally introduced, I have never learned.”

123. Stuart, John 1798

Narrative of Col. John Stuart, of Greenbrier. William

and Mary College Quart. Hist. Mag., 22, 234 (1914).

(Sugar trees in Virginia)

"I here hazard a conjecture that has often occurred

to me since I inhabited this place, that nature has designed

this part of the world a peaceable retreat for some of her

favorite children, . . . mines pragnant with saltpeter, &

forrests of sugar trees so amply provided and so easily

acquired . . . ”

124. French, Samuel 1799

History of the Town of Hardwick, Caledonia County.

Vermont Historical Gazetteer, Vol. I, p. 325 (1876).

(Sugaring at Hardivick)

"The last of March the snow lay 4 feet deep on a

level, but the weather was mild, and we prepared for

sugaring; but there came two feet more of snow, and

not a tree was tapped until the 15th of April. We gath¬

ered our buckets the 15th of May. Snow-banks were

visible the 9th of June.”

125. Smith, James 1799

An Account of the Remarkable . Occurrences in the

Life and Travels of Col. James Smith, during his Captiv¬

ity with the Indians, in the Years 1755, ’57, ’58 and '59.

Lexington, 1799, (a) p. 15, (b) p. 16, (c) p. 22, (d) p.

23, (e) p. 27, (f) p. 39, (g) p. 41, (h) p. 45.

a. (Sugar maples in Pennsylvania)

"On the head waters of this branch, and from thence

to the waters of Canesadooharie, there is a large body of

rich, well lying land — the timber is ash, walnut, sugar-

tree, buckeye, honey-locust and cherry, intermixed with

some oak, hickory, . . . ”

126 Wisconsin Academy of Sciences, Arts and Letters

b. (Results of the winter hunt)

“As the Indians on their return from their winter

hunt, bring in with them large quantities of bear's oil,

sugar, dried venison, &c. at this time they have plenty,

and do not spare eating or giving . . . thus they make

way with their provision as quick as possible. ..."

c. (Sugar tubs of elm bark)

See Drake. (Pt. I, ref. 42a of this bibliography)

d. (A load of maple sugar)

“When all things were ready we moved back to the

falls of Canesadooharie. In this route the land is chiefly

first and second rate, but too much meadow ground, in

proportion to the up land. The timber is white ash, elm,

black-oak, cherry, buckeye, sugar-tree, lynn, mulberry,

beech, white-oak, hickory, wild apple-tree, red-haw, black-

haw, and spicewood bushes. There is in some places, spots

of beech timber, which spots may be called third rate land.

Buckeye, sugar-tree and spice wood, are common in the

woods here. . . .

“On our arrival at the falls, (as we had brought with

us on horse back about two hundred weight of sugar, a

large quantity of bears oil, skins, . . . ) the canoe we

had buried was not sufficient to carry all, therefore we

were obliged to make another one of elm bark. ..."

e. (Homony, bear's oil and maple sugar)

“At this time homony plentifully mixed with bears'

oil and sugar; or dried venison, bears oil and sugar, is

what they offer to every one who comes in any time of the

day; and so they go on until their sugar, bears oil and

venison, is all gone. ..."

/. (Concentration of maple sap by freezing)

See Drake. {Ibid., 42b)

g. (Sugar camp on Big Beaver Creek)

“From our sugar camp on the head waters of Big

Beaver creek, to this place is not hilly, and some places

the woods are tolerably clear: but in most places exceed¬

ing brushy. ..."

Schuette and Hide — Maple Sugar

127

h. (Forest trees near Fort Detroit)

“There is plenty of good meadow ground here, and a

great many marshes that are overspread with water. . . .

The timber is elm, sugar-tree, black-ash, white-ash, abun¬

dance of water-ash, oak, hickory, and some walnut.”

126. Draper, Lyman C. 1800-1809

Antoine Le Clair's statement. Wisconsin Hist, Coll,, 11,

241 (1888).

(Maple sugar made in Milwaukee)

“The Indians at Milwaukee had no fruit trees, except

wild plums, which were plenty; there were blackberries,

grapes and strawberries, but no raspberries. There were

no nuts,— no pecans, no persimmons. The Indians manu¬

factured large quantities of maple sugar for their own

use, and for sale; they would live on it fast, and sell to

the traders,— the rule in this, as in other things, being

first a feast, then a famine. ...”

127. Sumner, Samuel ca. 1800

History of the Town of Troy, Orleans County. Ver¬

mont Historical Gazetteer, Vol. Ill, p. 819 (1877).

(Sugar maples yield a luxury)

“The sugar maple was a blessing to the early settlers

of Vermont. Those beautiful groves yielded an abundant

supply of sugar, affording to the indigent settler a neces¬

sary and luxury of life which the wealthy in older coun¬

tries could scarce afford, whilst the cheerful fires of this

wood, which, in our infancy, we saw blazing in the old

stone-backed chimneys, call up recollections of an enjoy¬

ment we cannot now find in the dull invisible warmth of

an air-tight stove, and the ashes of this generous tree,

when manufactured into potash or pearlash, furnished an

article for exportation, and almost the only one which

would warrant the expense in transporting it to the then

distant markets.”

128. Michaux, F. A. 1802

Travels to the Westward of the Allegany Mountains,

London, 1805, Vol, II, p. 81,

128 Wisconsin Academy of Sciences , Arts and Letters

(The sugar maple in southwestern Pennsylvania)

“The sugar maple is very common in all that part of

Pennsylvania, which is watered by the Monogahela and

the Allegany. This tree thrives best in cold, humid, and

mountainous countries, and its sap is more abundant, as

the winter has been more severe. The sugar obtained

from it has as dark a colour as that of the clayed sugar

of the first boiling, it is sold in loaves of 6, 8, or 10

pounds, at seven pence per pound. The inhabitants only

make it for their own use, most of them have tea or coffee

every day, but they only use it in the state in which it is

obtained from the first evaporation of the sap: they do

not refine it because of the great loss which it sustains

in this operation."

129. Curot, Michel 1804

A Wisconsin fur-trader's journal, 1803-04. Wisconsin

Hist . Coll. , 20, (a) 449, (b) 457 (1911).

a. ( Sugar made by Indians near Jaune, or

Yellow , River)

“Friday 9. (March) Mr. Sayer sent his wife this morn¬

ing to the Savages' lodges to make sugar. 4 Men went

with her to carry her Baggage and provisions."

b. “Tuesday 27th. (March) Mr. Sayer sent yesterday

with Kitchinimiscoutte, & Payechgigue who brought him

a Mocock of about 30 lbs. of sugar, two of his men and a

3 Gallon Keg of Pure H. W. . . . His wife (Savoiard's)

went to the Lodges to make sugar with her sister. Atao-

wabe came to the fort with them bringing nothing but

about Two Livres of sugar that he gave me."

130. Lewis, Merriweather, and George Rogers Clark 1804

Original Journals of the Lewis and Clark Expedition,

1804-1806. R. A. Thwaites, Editor. New York, 1904, (a)

Vol. I, p. 8; (b) p. 291; (c) Vol. VI, p. 169.

a. (Extra ration of whiskey for sugar makers)

“Detachment Orders

Camp River Dubois, Febr. 20th, 1804

“Lewis :

“The four men who are engaged in making sugar will

continue in that employment untill further orders, and

Schuette and Ihde — Maple Sugar

129

will receive each a half a gill of extra whiskey pr. day

and be exempt from guard duty.

Merriweather Lewis Cap1.

1st. U. S. Reg1. Infty.”

b. (Maple in bud-Mandan to Yellowstone)

“9th of April Tuesday 1805.

“Set out this morning verry early under a gentle breeze

from the S.E. at Brackfast the Indian deturmined to re¬

turn to his nation. I saw a Musquetor to day great num¬

bers of Brant flying up the river, the Maple, & Elm has

buded & cotton and arrow wood beginning to bud. ...”

c. (Meteorological observation for February , 180 4-)

“11th. The Sugar Maple runs freely, Swans pass from

the North.”

131. Malhiot, Frangois Victor 1804

A Wisconsin fur-trader’s journal - 1804-05. Wiscon¬

sin Hist. Coll, 19, (a) 174, (b) 232 (1910).

a. (Maple sugar exchanged for a gum)

“He did me the favor of giving me a keg of sugar for

a keg of gum, which had been given me at Kamanaiti-

quoya instead of a keg of sugar.”

b. (Mococks of maple sugar sold)

“1 A Mocock of Sugar. . . .”

132. Pike, Zebulon Montgomery 1805-1806

An Account of a Voyage up the Mississippi River,

from St. Louis to its Source. Compiled from Mr. Pike’s

journal. (Washington, 1807 ?), (a) p. 49. Elliott Coues,

Editor. The Expeditions of Zebulon Montgomery Pike, To

Headwaters of the Mississippi River, through Louisiana

Territory, and in New Spain, During the years 1805-6-7.

New York, 1895, new ed., (b) Vol. I, pp. 184-186.

a. (Visit to camp of Thomas, the Fols Avoine chief)

“ . . . The camp was situated in one of the finest sugar

groves imaginable. They were received in a truly patri¬

archal style: the chief pulled off Mr. Pike’s moccasins,

assigned him the best place in the lodge, and offered

dry cloths. After being presented with the syrup of the

130 Wisconsin Academy of Sciences , Arts and Letters

maple to drink, the chief asked his guest which he pre¬

ferred, beaver, swan, elk or deer, to eat. . . . they were

presented with something to eat ; at one a bowl of sugar , —

at another the tail of a beaver : generally, with what was

esteemed a delicacy by their Indian friends.

“The next morning Mr. Pike purchased two baskets of

sugar ; . . . ”

b. (Visit to Thomas sugar camp)

“Mar. 18th. We marched (up Spunk river), deter¬

mined to find the (Menomonee) lodges. Met an Indian

whose track we pursued through almost impenetrable

woods for about 2 1/2 miles to the camp. Here there was

one of the finest sugar-camps I almost ever saw, the whole

of the timber being sugar-tree. ... He then presented us

with syrup of the maple to drink, and asked whether I

preferred eating beaver, swan, elk, or deer; . . . We

were presented with something to eat; by some, with a

bowl of sugar : . . . ”

“March 19th. This morning purchased two baskets

of sugar. . . . ”

133. Dubuque, J. 1807

Fur-trade on upper lakes. 1778-1815. Operations of

Dubuque. Wisconsin Hist. Coll., 19, 319 (1910).

(Sugar at Prairie du Chien)

“As for the Accounting that you ask me for, I make it

the same as to what I owe as you and every one does. But

there are some small differences in regard to the price made

on sugar, rum, and powder; and after these are settled I

will adjust the Balance whenever you wish.”

134. Grignon, Pierre 1807

Fur-trade on the upper lakes - 1778-1815. Grignon

accounts 1806-1807. Wisconsin Hist. Coll., 19, 321 (1910).

A typical invoice

“1807 Rentree de L’envoice Oliva

Par diverses agrets

1000 lvs de Sucre .10 500”

135. Askin, Jno., Jr. 1808

Fur-trade on the upper lakes - 1778-1815. Wisconsin

Hist Coll, 19, 325 (1910).

Schuette and Ihde — Maple Sugar

131

(Son sends father maple sugar from

St. Joseph's Island)

“1 send Mr. J. & Mrs. Barthe Senr a Mocouts of sugar

addressed to your Care. One for Mr. Badishon wh Madelain

& my comp8, a Mocouts marked I P for Mr Peltier & a

Bundle. You'll receive One Mocouts Sugar, a Bundle of

Mats, & a mocout of dryed Huckleberrys which you'll

please accept of. The sugar is very clean I believe having

received it from a clean woman."

136. Smith, J. P. ca. 1810

History of the Town of Newark, Caledonia County.

Vermont Historical Gazetteer, Vol. I, p. 357.

(Maple sugar production in Newark)

“This town is also celebrated for its large productions

of maple sugar. The original growth of timber upon two-

thirds of its area, consisted of maple, beech and birch,

maple being in the excess; many beautiful groves of this

usefull tree have been cut down, but many yet remain.

The eastern slope of a mountain which extends from East

Haven to the centre of the town (a distance of three

miles), is covered for two miles or more with a continu¬

ous forest of sugar-maple. Many tons of sugar are made

here annually."

137. Tyrrel, J. B. ca. 1810

David Thompson's Narrative of his Explorations in

Western America. 1784-1812. Toronto, 1916, (a) p. 274,

(b) p. 275, (c) p. 283.

a. (Diet of wild rice and maple sugar)

“ . . . Proceeding five miles over the Lake we came to

the trading house of Mr. John Sayer, a Partner of the

North West Company, and in charge of this Department,

. . . Mr. Sayer and his Men has passed the whole winter

on wild rice and maple sugar, which keeps them alive,

but poor in flesh : . . . "

b. (Indian family property rights to maple groves)

“In the Spring the Natives employ themselves in mak¬

ing Sugar from Maple Trees, the process of doing which

is well known. The old trees give a stronger sap than the

young trees; The Canadians also make a great quantity,

132 Wisconsin Academy of Sciences , Arts and Letters

which when the sap is boiled to a proper consistence, they

run into moulds where it hardens. But the Indians prefer

making it like Muscovado sugar, this is done simply by

stirring it quickly about with a small paddle. The Plane

Tree also makes a good sugar, the sap is abundant, and

the sugar whiter, but not so strong. Both sugars have a

taste, which soon becomes agreeable, and as fine white

loaf sugar can be made from it as from that of the West

Indies. The natives would make far more than they do

if they could find a Market. . . . The Natives here call

themselves ‘Oochepoys’ and for some years have begun to

give something like a right of property to each family on

the maple sugar groves, and which right continues in the

family to the exclusion of others. But as this appropri¬

ated space is small in comparison of the whole extent ; any,

and every person is free to make sugar on the vacant

grounds. The appropriation was made by them in a coun¬

cil, in order to give to each family a full extent of ground

for making sugar, and to prevent disputes that would

arise where all claim an equal right to the soil and its

productions. And as in the making of sugar, several ket¬

tles and many small vessels of wood and birch rind for

collecting and boiling the sap are required, which are not

wanted for any other purpose, (they) are thus left in

safety on their own grounds for future use.”

c. (Sugar from the juice of the ash-leaved maple)

“He had traded 16 Cwt. of Maple Sugar from the

Natives; this was packed in baskets of birch rind of 28

to 68 lbs. each. The Sugar appeared clean and well made ;

that of the Plane Trees, looked like the East India Su¬

gars, and (was) much the same in taste: In this article

I have always noticed the supply is greater than the

demand.”

138. Wakefield, Priscilla. 1810

Excursions in North America, described in Letters

from a Gentleman and his Young Companion, to their

Friends in England. London, 1806, 2 ed., (a) p. 56,

(b) p. 191, (c) p. 295.

Schuette and Ihde — Maple Sugar 133

a. (Maple trees in Virginia)

“ . . . Virginia is intersected by numerous rivers and

creeks, and in many parts covered with forests of maples,

pines, cedars, the climbing trumpet-flower tree, the Caro¬

linian allspice, cornel trees, walnuts, laurels, bay-trees,

tulip trees, poplars, oaks, sumachs, acacias, and many

others : . . . ”

b. (The red maple in Maine)

“ . . . The black fir, the Weymouth pine, the red cedar,

the common fir, the red maple, the Pennsylvania ash, the

black birch, and the dwarf birch, are also common. . . . ”

c. (The sugar maple in Canada)

“ . . . The sugar maple grows in all parts of the coun¬

try, and is a very useful tree; as not only sugar may be

made from it, but vinegar, table beer, and an excellent

spirit. The country people pierce these trees with an

augur, and put a vessel beneath, to catch the sap as it

falls, which they refine by boiling till it is converted into

sugar, and a sufficient quantity is procured to nearly supply

the inhabitants, who seldom use any other.”

139. Lambert, John 1814

Travels through Canada and the United States. Lon¬

don, 1814, p. 83.

( Quality of Canadian maple sugar)

“Large quantities of maple sugar are sold at about

half the price of the West India sugar. The manufactur¬

ing of this article takes place early in the spring, when

the sap or juice rises in the maple trees. It is very labori¬

ous work, as at that time the snow is just melting, and

the Canadians suffer great hardships in procuring the

liquor from an immense number of trees dispersed over

many hundred acres of land. The liquor is boiled down

and often adulterated with flour, which thickens, and

renders it heavy: after it is boiled a sufficient time, it is

poured into tureens, and, when cold, forms a thick hard

cake of the shape of the vessel. These cakes are of a

dark brown colour, for the Canadians do not trouble them¬

selves about refining it. The people in Upper Canada

134 Wisconsin Academy of Sciences , Arts and Letters

make it very white; and it may be easily clarified equal to

the finest loaf sugar made in England.

“It is very hard, and requires to be scraped with a

knife when used for tea, otherwise the lumps would be

a considerable time dissolving. Its flavour strongly resem¬

bles the candied horehound sold by the druggists in Eng¬

land, and the Canadians say that it possesses medicinal

qualities, for which they eat it in large lumps. It very

possibly acts as a corrective to the vast quantity of fat

pork which they consume, as it possesses a greater degree

of acidity than the West India sugar. Before salt was in

use, sugar was eaten with meat in order to correct its

putrescency.”

140. Baird, Eliz. T. ca . 1815

Reminiscences of early days on Mackinac Island. Green

Bay Gazette, Dec. 4, 1886 ; Nov. 19, 1887 ; Wisconsin Hist.

Coll ., 14, 29 (1898).

( An American forest scene - maple sugar-making)

“A visit to the sugar camp was a treat to the young

folks as well as the old. In the days I write of, sugar was

a scarce article, save in the Northwest, where maple sugar

was largely manufactured. All who were able, possessed

a sugar camp. My grandmother had one on Bois Blanc

Island, about five miles east of Mackinac. About the first

of March, nearly half of the inhabitants of our town, as

well as many from the garrison, would move to Bois Blanc

to prepare for the work. Our camp was delightfully situ¬

ated in the midst of a forest of maple, or a maple grove.

A thousand or more trees claimed our care, and three

men and two women were employed to do the work.

“The ‘camp’,— as we specifically styled the building in

which the sugar was made, and the sugar-makers housed,

— was made of poles or small trees, enclosed with sheets of

cedar bark, and was about thirty feet long by eighteen

feet wide. On each side was a platform, about eighteen

inches high and four feet wide. One side was intended

for beds, and each bed when not in use was rolled up

nicely, wrapped in an Indian mat, then placed against

the wall; the bedroom then became a sitting room. The

walls on the inside were covered with tarpaulin, also the

Schuette and Ihde — Maple Sugar

185

floor. The women’s bedding was placed at one end of the

platform. The platform at the other side served as a

dining floor, one end of which was enclosed in cedar bark,

forming a chest for the dishes and cooking utensils. The

dishes consisted of some crockery, tin plates and cups,

and wooden dishes and ladles. The wing was added at

one end, for the men’s bedroom.

“At either end of the camp were doors, made large

to admit heavy logs for the fire. The fire-place was mid¬

way between the two platforms, and extended to within

six feet of the doors. At each corner of the fire-place

were large posts, firmly planted in the ground and ex¬

tending upward five feet or more. Large timbers were

placed lengthwise on top of these posts, and across the

timbers extended bars from which, by chains and hoops,

were suspended large brass kettles, two on each bar. On

the dining-room side, half way up the wall, ran a pole,

horizontally. This was to hold in place hemlock branches,

which were brought in fresh every evening. The place

between the fire and the platforms was kept very heat

by a thick, heavy broom, made of cedar branches, cut off

evenly on the bottom, and with a long handle. These

brooms are still used by semi-civilized Indians.

“The hanging of the kettle was quite a test of skill,

requiring three persons to perform the task. The fire had

to be burning when the hanging began. It was the duty

of one person to hang the kettle properly; of the second,

to pour in immediately a small quantity of sap to keep

the kettle from burning; of the third, to fill it with sap.

The peak of the roof was left open to allow the smoke

to escape, — and at night to let in the stars, as was my

childish fancy. . . .

“Now for the work; All of the utensils used in the

making of sugar were of that daintiest of material, birch-

bark. The casseau, to set at the tree, to catch the sap,

was a birch-bark dish, holding from one to two gallons.

The pails for carrying the sap were of the same material,

and held from three to four gallons. The men placed a

gauje or yoke on their shoulders, then a bucket would be

suspended on each side. The women seldom used this

yoke, but assisted the men in carrying the buckets, doing

136 Wisconsin Academy of Sciences , Arts and Letters

so in the usual manner. The mocock, in which the sugar

was packed, was also of birch-bark and held from thirty

to eighty pounds. The bark was gathered in the summer

at Bark Point. The name was afterward done in French

as ‘Point aux Ecorces,’ meaning ‘bark point’. The sailors

now miscall it, ‘Point au Barques.’

“The gouttiere or spout, which was made of basswood,

had to be cleaned each spring, before it was placed in the

tree; the birch-bark of the casseau was cleaned by taking

off a layer of the inner bark and then washing it. The

buckets were made by sowing the seams with hast (which

is taken from the inner bark of bass-wood), then gummed

over with pine pitch. They also were carefully washed

and dried before use. As a matter of course, the large

vessels to receive the sap were barrels made of oak. No

pine was ever used about the camp, as that would impart

a disagreeable taste. The strainers were made of a par¬

ticular kind of flannel, of very coarse thread and not wooly,

brought especially for this purpose by the merchants. I

remember well, the cleaning of these. After they had been

used, they were put in a tub of hot water and washed

(without soap) ; or pounded, rather, with a hattoir or

beetle, then rinsed in many waters.

“By this time the sap must be boiling. It takes over

twenty-four hours to make the sap into syrup, and the

boiling is usually begun in the morning. The fire is kept

bright all day and night. Two women are detailed to

watch the kettles closely, for when the sap boils down

nearly to syrup, it is liable to bubble over at any moment.

The women therefore stand by with a branch of hemlock

in hand; as soon as the liquid threatens to boil over they

dip the branch in quickly, and, it being cool, the syrup is

settled for a while. When at this stage, it requires closest

watching. When the sap has boiled down about one-half,

the women have to transfer the contents of one kettle,

which would spoil all. As fast as a kettle is emptied it

will be filled with water and set aside, awaiting the general

cleaning. The kettles require the utmost care, being

scoured each time emptied, keeping one woman employed

nearly all the time. Sand and water are the cleansing

agents used.

Schuette and Ihde — Maple Sugar 137

"All this time, if the weather favors the running of

the sap, it is brought as fast as possible, and the boiling

goes on. At this period, my grandmother would send me

my little barrel full of syrup. The miniature barrel I

still have in my possession. The barrel bears the date

1815, and is now dark and polished with age, and is a

rare memento of those halcyon days. It holds less than a

pint, and was made by an Ottowa Indian, out of a solid

piece of wood, sides and ends all one, the interior being

ingeniously burned out through the bung hole. The re¬

ceipt of this was the signal that the time had come when

I too might visit the camp.

"When made, the syrup is put in barrels, awaiting

the time when it can be made into sugars of various

kinds, the modus operandi thus : A very bright brass ket¬

tle is placed over a slow fire (it can not be done at boiling

time, as then a brisk fire is required),— this kettle contain¬

ing about three gallons of syrup, if it is to be made into

cakes; if into cassonade, or granulated sugar, two gallons

of syrup are used. For the sugar cakes, a board of bass¬

wood is prepared, about five or six inches wide, with

moulds gouged in, in forms of bears, diamonds, crosses,

rabbits, turtles, spheres, etc. When the sugar is cooked

to a certain degree, it is poured into these moulds. For

the granulated sugar, the stirring is continued for a longer

time, this being done with a long paddle which looks like

a mushstick. This sugar has to be put into the mocock

while still warm, as it will not pack well if cold. The

work is especially difficult; only a little can be made at

a time, and it was always done under my grandmother's

immediate supervision.

"The sugar-gum, or wax, is also made separately. Large

wooden bowls, or birch-bark casseaus, are filled with snow,

and when the syrup is of the right consistence it is poured

on the snow in thin sheets. When cooled, it is put into

the birch-bark, made into a neat package, and tied with

bast. The syrup made for table use is boiled very thick,

which prevents its souring. For summer use it is put into

jugs and buried in the ground two or three feet deep,

where it will keep for a year, more or less.

138 Wisconsin Academy of Sciences, Arts and Letters

“One time, a party of five ladies and five gentlemen

were invited to the camp. Each lady brought a frying-pan

in which to turn les crepes or pancakes, which was to be

the special feature and fun of the occasion. All due prepa¬

ration was made for using the frying-pans. We were noti¬

fied that no girl was fitted to be married until she could

turn a crepe . Naturally, all were desirous to try their

skill in that direction, whether matrimonially inclined or

not. The gentlemen of the party tried their hand at it,

as well as the ladies. It may not be amiss here to explain

what turning a crepe meant ; when the cake was cooked on

one side, it was dexterously tossed in the air, and ex¬

pected to land, the other side up, in the pan. Never did I

see objects miss so widely the mark aimed at. It seemed

that the crepes were influenced indeed by the glee of the

party; they turned and flew everywhere, but where want¬

ed. Many fell into the fire, as if the turner had so in¬

tended. Some went to the ground, and one even found its

way to the platform, over the head of the turner. One

gentleman (Henry S. Baird) came up to Mrs. John Dous-

man, and holding out his nice fur cap, said, 'Now turn

your cake, and I will catch it.' Mrs. Dousman was an

adept at turning, and before the challenger had time to

withdraw his cap, with a toss she deftly turned the cake

and landed it fairly into the cap. You may imagine the

sport all this afforded. In due time, a nice dinner was

prepared. We had partridges roasted on sticks before the

fire; rabbits and stuffed squirrel, cooked French fashion;

and finally had as many crepes, with syrup, as we desired.

Every one departed with a bark of wax, and sugar cakes.”

141. Heckewelder, John 1817

History, Manners and Customs of the Indian Nations

who once Inhabited Pennsylvania and the Neighboring

States; Memoirs Historical Society Penn., 12, 194-195

(1876).

Food and Cookery

"... In the dough of this kind of bread they fre¬

quently mix boiled pumpkins, green or dried, dry beans,

or well pared chestnuts, boiled in the same manner, dried

Schuette and Ihde— Maple Sugar

139

venison well pounded, whortle berries, green or dry, but

not boiled, sugar and other palatable ingredients.

“The Indians have a number of manners of prepar¬

ing their corn. They make an excellent pottage of it,

by boiling with fresh or dried meat (the latter pounded),

dried pumpkins, dry beans, and chestnuts. They some¬

times sweeten it with sugar or molasses from the sugar-

maple tree. . . .

“ . . . They make an excellent preserve from the cran¬

berry and crab-apple, to which, after it has been well

stewed, they add a proper quantity of sugar or molas¬

ses. ...”

142. Evans, Estwick 1818

A Pedestrious Tour of Four Thousand Miles through

the Western States and Territories. Concord, 1819, p. 171 ;

R. G. Thwaites, Editor, Early Western Travels 1748-1846.

Cleveland, 1904, Vol. 8, p. 275.

(Maple sugar production)

“ . . . Several millions of pounds of mapel sugar are

made here annually ; . . . ”

143. Michaux, F. A. 1819

The North American Sylva, or a Description of the

Forest Trees of the United States, Canada and Nova

Scotia. Paris, 1819, Vol. I, (a) p. 216, (b) p. 218, (c) p.

221, (d) p. 223, (e) p. 228, (f) p. 237.

a. White Maple (Acer eriocarpum)

“In the Atlantic parts of the United States, this spe¬

cies is often confounded with the Red Maple which it

nearly resembles ; west of the Mountains, they are con¬

stantly distinguished, and the Acer eriocarpum is known

by no other name than White Maple.

“Some of the inhabitants on the Ohio make sugar of

its sap, by the same process which is employed with the

Sugar Maple. Like the Red Maple, it yields about half the

product from a given measure of sap; but the unrefined

sugar is whiter and more agreeable to the taste than that

of the Sugar Maple. The sap is in motion earlier in this

species than in the Sugar Maple, beginning to ascend about

the 15th of January; so that the work of extracting the su-

140 Wisconsin Academy of Sciences, Arts and Letters

gar is sooner completed. The cellular integument rapidly

produces a black precipitate with sulphate of iron.”

b. Red Flowering Maple (Acer rubrum)

“Different names are given to this tree in different

parts of the United States; east of the Alleghany moun¬

tains it is called Red flowering Maple, Swamp Maple and

Soft Maple; in the Western Country, simply Maple. The

first denominations, which is most generally in use, is also

most appropriate, as the young shoots, the flowers, and

the fruit are red. ...”

c. (Sugar from the red maple, or plaine)

“The French Canadians make sugar from the sap of

this Maple, which they call Plaine, but, as in preceding

species, the product of a given measure is only half as

great as is obtained from the Sugar Maple.”

d. Sugar Maple (Acer saccharinum)

“This species, the most interesting of the American

Maples, is called Rock Maple, Hard Maple, and Sugar

Maple. The first of these names is most generally in use,

but I have preserved the last, because it indicates one of

the most valuable properties of the tree.

“The Sugar Maple covers a greater extent of the

American soil than any other species of this genus. It

flourishes most in mountainous places, where the soil

though fertile is cold and humid. Besides the parts which

I have particularly mentioned, where the face of the

country is generally of this nature, it is found along the

whole chain of the Alleghanies to their termination in

Georgia, and on the steep and shaly banks of the rivers

which rise in these mountains.”

e. (Manufacture of maple sugar)

Descriptive.

/. Black Sugar Tree (Acer nigrum)

“In the Western States, and in the parts of Pennsyl¬

vania and Virginia, which lie between the mountains and

the Ohio, this species of Maple is designated by the name

of Sugar Tree, and frequently, by the more characteristic

denomination of Black Sugar Tree; probably, on account

Schuette and Ihde — Maple Sugar 141

of the dark colour of its leaves, in comparison with those

of the true Sugar Maple, which sometimes grows with it.

In the extensive country of Genesee both species are in¬

discriminately called Rock Maple and Sugar Maple. This

confusion seems to have arisen from the country's being

settled principally by emigrants from the Eastern States,

who, finding the Black Sugar Tree applicable to the same

uses with the other, and equally productive of sugar, have

given it the same name. The two species have also been

confounded by* Botanists, in describing the vegetable pro¬

ductions of America."

144. Childs, Ebenezer 1820

Recollections of Wisconsin since 1820. Wisconsin Hist.

Coll , 4, (a) 161, (b) 175 (1858).

a . (Maple-sugar making at Green Bay)

“There were quite a number of very respectable French

families residing at the Bay when I arrived there. . . .

They caught large quantities of sturgeon and trout, and

they made immense quantities of maple sugar. At the

proper season in the spring, the entire settlement would

remove to their sugar-campe, often remain two months,

each family making eight to ten hundred pounds of the

finest sugar I ever saw."

6. (Barter with maple sugar at Green Bay, Wis.)

“I furnished the Indians with provision that fall and

winter; they paid me in furs and maple sugar. I pur¬

chased some six tons of sugar of them."

145. Doty, James Duane 1820

Northern Wisconsin in 1820. Wisconsin Hist. Coll. , 7,

199 (1876).

(Indians make maple sugar at Rice Lake)

“ . . . Their families being left at home in this hunt,

repair to the sugar camps, and are engaged in manufac¬

turing sugar during the absence of the men, of which

they make large quantities."

146. Doty, James Duane 1820

Official Journal, 1820. Expedition with Cass and School¬

craft. Wisconsin Hist. Coll, 13, 207 (1897).

142 Wisconsin Academy of Sciences, Arts and Letters

(Indian sugar camp near Sandy Lake)

“ . . . Thus far the land has been low,-— timber birch,

maple, bass wood, and elm. Next we steered S. 20 west

6 miles over a ridge one half (the first covered chiefly

with pine the residue the sugar maple). The Indians

had a large establishment in this wood, this last spring

where from every appearance they must have made great

quantities of sugar.”

147. Irwin, M. 1820

Fur trade and factory system at Green Bay. 1816-21.

Letter of M. Irwin, U. S. Factor, to Doctor J. Morse. Wis¬

consin Hist . Coll, 7, 286 (1876).

(Maple sugar traded for whiskey)

“ ... He adds, T will venture to say, that out of two

hundred barks of sugar taken, not five have been pur¬

chased with any other commodity than whiskey. I have

not been able to procure a pound (of sugar) from the

Indians, but can get a supply from the traders at ten

cents a pound.’

“Independent of the known veracity of Mr. Varnum,

the fact that private traders can sell sugar at ten cents

a pound, is pretty conclusive evidence of the manner in

which they obtain it.”

148. Locke, John 1820

On the manufacture of sugar from the river maple

(Acer eriocarpum of Linnaeus) . Am . /. Sciences, Arts, 2,

(a) 258, (b) 261.

a. (Sugar from the river maple)

“It seems not to be generally known, that sugar is

afforded in any considerable quantity, by any other species

than the sugar maple, (Acer saccharinum) ; but I have

found that in some parts of New-England, more sugar is

made from the river, than from the sugar maple.5

“A peculiar method of tapping is practiced in Frye-

burg. The incision from which the sap issues is made

by driving a gouge a little obliquely upward, an inch or

more into the wood. A spout or tap about a foot long,

to conduct off the sap, is inserted about two inches below

this incision with the same gouge. . . . One principal

6 The author bases his statements upon observations made in Fryeburg, Maine, on the

Saco river, where large quantities of maple sugar were annually made. The sap of the river

maple was generally deemed there to be sweeter than that of the sugar maple; its sugar

whiter and of a better quality.

Schuette and Ihde— Maple Sugar

148

advantage of this method is, that the wound in the tree

is so small that it is perfectly healed or ‘grown over* in

two years, the tree sustaining little or no injury. The

other common methods of tapping are two. 1. With an

axe. An oblique incision three or four inches long, is

made in such a manner that all the sap will be conducted

to the lower corner, where it passes into a spout inserted

with a gouge as above. Disadvantages of this method.

The surface being much exposed to the air and sun, is

presently dried, so as to diminish very much the quantity

of sap. The wound in the tree is extensive and a ruinous

decay is often the consequence, the tree becoming rotten-

hearted. 2. With an auger. The tree is perforated an

inch or more with an auger three fourths of an inch di¬

ameter, and a tube made of elder or sumach is inserted

to conduct off the sap. The end of the tube is made taper¬

ing so as to bear only at the outer edge of the tube. Dis¬

advantage. The tap presses upon the external grains so

as to obstruct the flow of sap from them; and it is from

these external grains that most of the sap is obtained. The

method tapping with the gouge is undoubtedly superior

to either of the others, but in a sugar maple there might

be difficulty in inserting the gouge to a sufficient depth on

account of its superior hardness.

“It seems that the superiority of the river over the

sugar maple is not universal ; for Micheaux says, that on

the Ohio only one half the quantity is obtained from the

river, that is afforded by the sugar maple.”

b . (Lin9* to the maple tree)

“In a poem written in Fryeburg called ‘The Village' the

following lines are bestowed upon it.

‘More sacred than the thunder chosen oak,

‘Let not the maple feel the woodman's stroke.

‘Fair maple! honours purer far are thine

‘Than Venus's myrtle yields, or Bacchus's vine;

‘Minerva's olive consecrated tree,

‘Deserves not half the homage due to thee.

‘The queen of trees, thou proudly towerist on high,

‘Yet wave thy limbs in graceful pliancy.' "

144 Wisconsin Academy of Sciences , Arts and Letters

149. Morse, Jedidiah 1820

Report to the Secretary of War of the United States.

New Haven, 1822, Appendix p. 50.

(Maple products made at Green Bay , Wis.)

In commenting upon the activities of the half-breed

population at Green Bay, the author states, “These peo¬

ple, and the Menominees with whom, by the ties of re¬

lationship, they are connected, make from the maple tree

about one hundred pounds of sugar annually; and from

three to four hundred gallons of molasses. These, with

their skins, etc. are nearly all sold for whiskey, at an

immense sacrifice.”

150. Finley, J. B. 1821

History of the Wyandott Mission. Cincinnati, 1840,

(a) p. 124, (b) p. 125, (c) p. 126, (d) p. 128.

a . (Sugar-making season)

“In February, nearly all of the Indians went to the

woods, to trap and make sugar. They seldom return from

these expeditions until the first of April.”

b. (Sugar- flavored boiled raccoon meat)

“Soon we had placed before us a kettle filled with fat

raccoons, boiled whole, after the Indian style, and a pan

of good sugar molasses. These we asked our heavenly

Father to bless, and then each carved for himself, with

a large butcher knife. I took the hind quarter of a rac¬

coon, and holding it by the foot, dipped the other end

in the molasses, and eat it off with my teeth. Thus I con¬

tinued dipping and eating until I had pretty well finished

the fourth part of a large coon.”

c. (Troughs as sap containers)

“The troughs in which they catch their sugar water,

are made of bark, and hold about two gallons. They have a

large trough, made like a bark canoe, into which they

gather from the small ones. The women make the sugar,

and stretch all the skins. The men trap and hunt.”

d. (War-time rations of the Indian)

“Deer meat is sliced thin; and dried over the fire,

until it can be easily pounded in a mortar. This, mixed

Schuette and Ihde — Maple Sugar

145

with sugar and dipped in bear’s oil, is the greatest lux¬

ury of an Indian table. This, with corn parched in a

kettle, and pounded to meal, then sifted through a bark

sieve, and mixed with sugar, makes the traveling provi¬

sion of an Indian in time of war.”

151. Grignon, L. 1821

Fur-trade in Wisconsin. Letter to Robt. Stuart, Michi-

milimackinac from La Bay verte 7 June 1821. Wisconsin

Hist Coll , 20, 200 (1911).

(Sugar supplies plentiful in Green Bay)

‘There is much sugar here but few Peltries, ...”

152. Nuttall, Thomas 1821

A Journal of Travels into the Arkansas Territory.

Philadelphia, 1821, p. 113; R. G. Thwaites, Editor, Early

Western Travels 1748-1846. Cleveland, 1905, Vol. XIII,

p. 161.

(Low-cost sugar)

“ . . . Sugar and Coffee are also high priced articles,

more particularly this year. There is a maple in this

country, or rather, I believe, on the banks of the White

river, which has not come under my notice, called the

sugar-tree (though not, as they say, the Acer Sacchari-

num), that would, no doubt, by a little attention afford

sugar at a low rate ; . . . ”

153. Deane, Samuel 1822

The New England Farmer; or Georgical Dictionary.

Boston, 1822, 3 ed., p. 262.

( Observations on making sugar from the

sap of the rock-maple)

This is an edited quotation from the American Mu¬

seum, vol. 6, pp. 98-101 for which see reference 28, part

1, of this bibliography.

154. Ellis, Albert G. 1822

Fifty-four Years’ Recollections of Men and Events in

Wisconsin. Wisconsin Hist Coll, 7, 1876, (a) 220, (b)

222.

a. (Sugar-making at Green Bay)

“The product of the sucreries of the better class of

the French, was a fair article of sugar, of ready sale,

146 Wisconsin Academy of Sciences , Arts and Letters

and in some respects preferable to the best muscovado.

They had learned to use the utmost neatness and caution

to keep out all impurities, and had attained to great per¬

fection in the purifying process. All the sap was strained

through a fine sieve into the kettles — the syrup was

strained twice before granulating; and here came in the

product of the chickens, to wit, the eggs, the whites of

where were broken in the boiling syrup, when all impuri¬

ties immediately came to the surface and were removed.

The sugar, when strained off and cooled, was quite fair

and pure. Some of the more enterprising and forehanded,

bought syrup and coarse sugar of their Indian retainers,

and their less able neighbors, and went into the purify¬

ing process on a large scale, and thus largely increased

their product for the season. A few families of this

class had a preference in the sugar market at the frontier

trading posts, their mococks, branded with their names,

always being sought, at advanced prices.

“As before stated, the Easter festival was generally

celebrated at those sucreries; for this reason those who

had the chickens, and could do it, took them into the

woods, made houses for them, and saved a store of eggs

for this festival. Then it was that their friends at the

settlement, the Americans and army officers, were in¬

vited to visit them, and the invitations were rarely de¬

clined. The American citizens, the gentlemen and ladies

of the army, found no greater enjoyment than one of

these spring festivals in the depth of the great maple

woods, in their commodious sugar-houses. . , . These frol¬

ics were often enlivened by an old-fashioned ‘candy-pulT,

when the French girls presented their sweet-hearts, on

parting, with a cake of candy, folded in a strip birch-

bark, which they called their ‘billet doux\”

b. (Maple sugar not accepted in trade)

“ . . . the natives, as well as the French inhabitants,

made quantities of maple sugar; this was not current at

New York, for payment of goods, as peltries were; and so

not much cared for by the old traders. The Indians

resorted with it to the United States factor, Major Irwin,

who bought large quantities of it; and had many thou-

Schuette and Ihde— Maple Sugar

147

sand pounds in store at the time of our arrival in

1822. ...”

155. Moore, Jacob B. 1822

Topographical and historical sketch of Andover, N. H.

Farmer and Moore's Historical Collections, Vol. I, p. 12.

( A neglected branch of domestic economy)

“ . . . The growth of wood, in the other parts of the

town, is principally oak, beach and sugar-maple. . . .

From its first settlement until within a few years of the

inhabitants have anually supplied themselves with sugar

from their own farms; but the trees now beginning to

decay, and little pains being taken in their preservation,

this branch of domestic economy is almost wholly neg¬

lected.”

156. Woods, John 1822

Two Years' Residence in the Settlement on the English

Prairie in the Illinois Country, United States. London,

1822, (a) p. 235, (b) p. 306, R. G. Thwaites, Editor, Early

Western Travels 1748-1846. Cleveland, 1904, Yol. X, (c)

p. 312, (d) p. 354.

a . (Maple trees tapped near Harmonie, Indiana)

“In passing some woods, we saw some sugar-maple

trees that were tapped, with the liquor then running into;

we dismounted, and had a good draught or two of the

liquor; it was pleasant-tasted.”

b. (Sugar made in Illinois)

“ . . . This year some sugar has been made near us,

from the white maple, and it appears to answer nearly

as well as the sugar-maple. I suppose, another season,

it will be made in considerable quantities.”

157. Hunter, John D. 1823

Manners and Customs of Several Indian Tribes Lo¬

cated West of the Mississippi. Philadelphia, 1823, p. 269.

(Indian beverage)

“Their usual drink is pure cold water; though some¬

times they mix maple sugar with it, or honey, which they

procure in considerable quantities from the stores of

honey bees, deposited in hollow trees; . . . ”

148 Wisconsin Academy of Sciences, Arts and Letters

158. Schoolcraft, Henry R. 1823

Personal Memoirs of a Residence of Thirty Years with

the Indian Tribes on the American Frontiers. Philadel¬

phia, 1851, p. 162.

(Sugar-making season at hand)

“It is now the season for making sugar from the rock

maple by the Indians and Canadians in this quarter. And

it seems to be a business in which everyone is more or

less interested. Winter has shown some signs of relax¬

ing its iron grasp, although the quantity of snow upon

the ground is still very great, and the streams appear

to be as fast locked in the embraces of frost as if it were

the slumber of ages. Sleighs and dog trains have been

departing for the maple forests, in our neighborhood,

since about the tenth instant, until but few, comparatively,

of the resident inhabitants are left. Many buildings are

entirely deserted and closed, and all are more or less

thinned of their inhabitants. It is also the general sea¬

son for sugar making.

“I joined a party in visiting one of the camps. . . .

We found a large temporary building, surrounded with

piles of ready split wood for keeping a fire under the ket¬

tles, and large ox hides arranged in such a manner as to

serve as vats for collecting the sap. About twenty ket¬

tles were boiling over a central elongated fire.”

159. Anon. 1824

Topographical sketch of Salisbury, New-Hampshire.

Farmer and Moore’s Historical Collections, Vol. Ill, p.

297.

(Forest trees at Salisbury)

“ . . . The hilly lands in their natural state, were

covered with a heavy growth of the sugar maple, white

maple, beech, birch, elm, ash and red oaks; the valleys

were interspersed with ever-greens. ...”

160. Lawe, John 1824

Fur-trade in Wisconsin. Letter to Jacques Porlier,

Portage, Green Bay 25th April 1824. Wisconsin Hist . Coll.,

20, 338 (1911).

Schuette and Ihde— Maple Sugar

149

(Sugar abundant at Green Bay)

. . Sugar will be somewhat abundant but there is

so many purchasers we stand but a small chance of geting

our share & they estimate it so high that it will cost us

more than the first cost of our Goods to pay for it.”

161. Evelyn, John 1825

Silva: or, A Discourse of Forest-Trees. London, 1825,

Vol. I, 5 ed., p. 199.

(Maple sugar— refined in Normandy)

“The Savages in Canada, when the sap rises in the

Maple, by an incision in the Tree, extract the liquor;

and having evaporated a reasonable quantity thereof, (as

suppose seven or eight pounds) there will remain one

pound as sweet and perfect sugar as that which is gotten

out of the cane; part of which sugar has been for many

years constantly sent to Rouen in Normandy, to be re¬

fined: There is also made of this sugar an excellent syrup

of Maiden-hair and other capillary plants, prevalent

against the scurvy; though Mr. Ray thinks otherwise by

reason of the saccharine substance remaining in the de¬

coction.”

162. McKenney, Thomas L. 1826

Sketches of a Tour to the Lakes, of the Character

and Customs of the Chippeway Indians, and of Incidents

connected with the Treaty of Fond Du Lac. Baltimore,

1827, (a) p. 192, (b) p. 193.

a. (Trade in maple sugar at Sault de St. Marie)

“The staples of the place, are the white fish and maple

sugar, and some few, but not many furs.”

b. (Mococks of maple sugar)

“Sugar is the next great staple. It is made from the

maple, and principally by the Indian women. You know

the manner of tapping the tree, and boiling the sap, and

fining the sugar, and therefore it is not necessary that

I should trouble you with an account of it. Henry tells us

the earlier part of the spring is that best adapted to make

maple sugar. The sap runs only in the day, and it will

not run unless there has been a frost the night before.

When, in the morning, there is a clear sun, and the night

150 Wisconsin Academy of Sciences , Arts and Letters

has left ice the thickness of a dollar, the greatest quantity

is produced. Three families in this neighborhood, of

which my old friend Mr. J . . ,’s is one, make generally

four tons of sugar in a season. Some of it is very beau¬

tiful. I have some mococks of it given to me by Mrs.

Johnson, of her own make. It is as white as the Havanna

sugar, and richer. A mocock is a little receptacle of a

basket form, and oval, though without a handle, made of

birch bark, with a top sewed on with wattap , (the fine

roots of the red cedar, split) the smaller ones are orna¬

mented with porcupines’ quills, died red, yellow, and

green. These ornamented mococks hold from two to a dozen

table spoons full of sugar, and are made for presents,

or for sale, to the curious. The larger ones, also of birch

bark, are not ornamented, and contain from ten to thirty

pounds of sugar. This is an article of exchange with

those who make it. They give for labour, for goods, &c.

and generally at about ten cents per pound. Indians often

live wholly upon it ; and Henry tells us he has known them

to grow fat upon this sugar alone.”

163. Doty, James Duane 1827

Letter to The Honorable James Strong, Chn. of the

Comt. on Territories, Dec. 25, 1827. Wisconsin Hist . Coll. ,

13, 245 (1897).

(Sugar exported from i(that part of the Michigan

Territory which lies to the north and west

of Lakes Huron and Michigan ”)

“The exports from this Territory have been usually

estimated to consist annually of Furs and Peltries, valued

at 300,000$ — White fish 800 to 1000 barrels. — Sugar

200,000 lbs. and Lead 10,000,000 lbs.

164. Lockwood, James H. 1827

Early times and events in Wisconsin. Wisconsin Hist .

Coll., 2, 156 (1856).

(Sugar-making in Iowa)

“ ... he went with his family up the Yellow or Paint¬

ed Rock Creek, above twelve miles above the Prairie, on

the Iowa side of the Mississippi River, to make su¬

gar _ ” (cf. Snelling, 1827)

Schuette and Ihde— Maple Sugar

151

165. (Snelling, Wm. J.) ? 1827

Early days at Prairie do Chien. Wisconsin Hist Coll.,

5, 126 (1868).

(Sugar-making by white men in Iowa)

“ . . . Every one knows that, in the Western country,

French people make maple sugar in the spring. M. Me-

thode chose to set up his sugar camp at the mouth of the

Yellow River, two miles from Prairie du Chien. ...”

166. Smith, Thomas and John 0. Charles 1828

The Origin and History of Missions. Boston, 1887,

Yol. II, p. 389.

(A penitent interpreter of dreams among

the Ojibway Indians)

The Rev. Wm. M. Ferry, the founder of the mission

at Mackinaw, wrote of an “excessively intemperate” Indian

woman of Ojibway blood who, because of influential con¬

nections, had been selected to become an interpreter of

dreams. In about 1825 “her serious attention to religion

commenced, the amount of which for some length of time

was very fluctuating.” In one of her drunken orgies she

“lost her sack.” Repentance followed. Of her it was

stated: “During the spring, 'while at the sugar camp, she

says she was greatly distressed during the whole time.

When gathering sap, she often had feelings like these -

Here I am going the same round daily from tree to

tree, and can find no relief - 1 must always carry this

wicket heart, and when I die, be miserable forever.”

167. Macauley, James 1829

The Natural, Statistical, and Civil History of the

State of New York. New York, 1829, p. 536.

Remarks on some of the trees, etc .

“The Indians in this State and Canada are said to have

made sugar anterior to the colonization. The quantity

made lessens as the woods are cut down. There are many

districts, in which large quantities were formerly made,

which, at present, do not afford a pound. In general, no

attention has been paid to the preservations of the trees.

In addition to this, the snow, where the lands are cleared,

go off much earlier, and the seasons have become shorter

152 Wisconsin Academy of Sciences , Arts and Letters

and more irregular: hence, the trees do not afford so

much sap as formerly. ...”

168. Sheppard, W. 1829

Observations on the American Plants described by

Charlevoix. Trans. Literary Hist. Soe. Quebec , 1, 228,

(1829).

(Erable a fleurs rouges)

“Acer rubrum. Red maple. Called by the Canadians,

Plane. A large tree growing commonly about Quebec, and

in common with the other maples yields a saccharine sap ;

but the sugar made from it, is inferior in quality to that

obtained from Acer saccharinum and nigrum. It is sur¬

prizing that Charlevoix should mention this species only,

out of the nine found in America.”

169. Stambaugh, Samuel 1881

Report (to Secretary of War) on the quality and con¬

dition of Wisconsin Territory, 1831. Wisconsin Hist. Coll.,

15, (a) 408, (b) 413, (c) 415 (1900).

a. (Brothertown Indioms settle on Menominie

land east of Fox River)

“ . . . I assured the Menominies that the removal of

the stranger Indians upon this land would not impair

their treaty stipulations with the United States, nor dimin¬

ish the kind feeling entertained toward them by the

government, but that, should the Treaty be ratified by

the Senate, all its provisions would be carried into effect.

With this assurance the Chiefs left me apparently satis¬

fied, although some of their finest sugar Camps are on

the land occupied by the Brothertown Indians, which will

be much injured if not entirely destroyed by their settle¬

ment.”

b. (Maple groves near Oconto Falls)

“ . . . There are exuberant groves of Maple and Beech

in the neighborhood of these falls, which is the best evi¬

dence of the fertility of the soil ; and a short distance be¬

low, on the south side of the mill, the Indians have Sugar

Camps, at which they manufacture large quantities of

sugar. . . . The land is covered with a thriving growth

of Oak, Beech and Maple. ...”

Schuette and Ihde~Maple Sugar

153

c. (Sugar camps near Little Kaccalin)

“There are several sugar camps in the neighborhood,

where large quantities of sugar are manufactured from

large and beautiful Maple groves.”

170. Anon. 1832

History of the Delaware and Iroquois Indians former¬

ly Inhabiting the Middle States. Philadelphia, 1832, p.

89.

(The great value of the sugar maple tree)

“Of all the productions of the earth, however, with

which the Indians were familiar, none was better esteemed,

or more interesting than the Sugar maple tree , so called

because of the sap which runs from it at a particular

season of the year, from which they make a quantity of

delightful sugar. This sap is found in greatest plenty in

the spring of the year. At this time they make an incision

into the tree; and through this hole the sap is received,

by means of a funnel, into wooden troughs or vessels.

It is then boiled over a slow fire in kettles, and becomes

as good as any sugar in the world. The flowing season

lasts generally one or two months.

“Sugar boiling is chiefly the employment of women.

A kettle holding between sixty or seventy quarts, with

two of a smaller size, for ladles, will boil, with ease, two

hundred pounds of sugar in one season, besides furnish¬

ing a large quantity of molasses. Instances have been

known of one tree producing above three hundred quarts

of good sap for sugar, and as much for molasses. About

thirty-five or forty quarts of sap produce one pound of

sugar. Thus about eight pounds of sugar, and as many

of molasses, may be collected from one tree, and the

trees last eight or nine years. A large quantity of maple

sugar is made every year in the United States. Dr. Rush,

who wrote on this subject of sugar maple, which he re¬

garded as a peculiar gift of a benevolent providence, cal¬

culates that the cultivation of these trees would furnish

support to many thousand families, and even become an

important branch of revenue to the government.”

154

Wisconsin Academy of Sciences , Arts and Letters

171. Ferrall, S. A. 1832

A Ramble of Six Thousand Miles through the United

States of America. London, 1832, p. 173.

(Sugar Maple in Indiana)

‘The farmers use, almost exclusively, the sugar of the

maple ( acer sacckarinum) which they manufacture them¬

selves. The space in which a number of these trees are

found, they call a ‘sugar camp.’ The process of manufac¬

turing is as follows : - After the first frost, the trees are

tapped, by perforating the trunk in an ascending direc¬

tion. A spout of alder is inserted in the perforation,

and the sap drips through this conduit into a trough of

wood. The sap is then boiled with a spoonful of slaked

lime, the white of an egg or two, and about a pint of

milk, to every fifteen gallons. An ordinary tree commonly

gives four pounds of good course brown sugar, which

when refined can be made equal to superior lump sugar.”

172. Hudson, Charles 1832

History of the Town of Westminster, Mendon, Mass.,

1832, p. 6.

(Sugar-making in Massachusetts)

“ It is a good grazing township. The growth of

wood is beech, maple, birch, oak, chestnut, ash, hemloc,

and pine. Beech and rock maple, are the most abundant.

From the latter sugar for family consumption, is made

in considerable quantities.”

173. Radcliff, T. 1832

Authentic Letters from Upper Canada. Dublin, 1833,

p. 229.

(Shuggar from the “maypole” )

“But what flogged all that I had ever seen, was making

sugar out of a tree, Mary -not a word of a lie do I tell

you; you take a big gimlet and make a hole in the tree,

(the maypole I think they call it,) and out comes the

shuggar, like sweet water thick like, and you boil it, and

you — but where’s the use of my telling you any thing

about it, as you have no sugar trees at home.

“I remember when you and I thought a shuggar stick,

a mighty good sort of thing, never thinking I’d lay my

eyes on a sugar tree”

Schuette and Ihde — Maple Sugar

155

174. Flint, Timothy 1833

History and Geography of the Mississippi Valley. Cin¬

cinnati, 1833, Vol. I, 3 ed., p. 43.

(“Country sugar ” and cane sugar)

“The sugar maple is very abundant in the northern

and middle regions of this valley. ... In different parts

of Ohio, Kentucky, Tennessee, Illinois, and Missouri, it is

made, not only for consumption, but for sale. . . . The

season of making it is generally one of festivity and high

holiday. We have tasted sugar loaf made from it, which

could in no way be distinguished from that made from

the cane. The cheapness of the latter kind, the abundance

and excellence of its growth in the lower country, and

the diminished expense of transporting it to the upper

states, in consequence of the multiplication of steam boats,

has diminished the demand for what is called 'country

sugar/ and the manufacture of it has decreased, since

the use of steam boats.”

175. Porter, Jacob 1834

Topographical Description and Historical Sketch of

Plainfield, in Hampshire County, Massachusetts. Green¬

field, 1834, (a) p. 11, (b) p. 12.

a. (Native timber)

“The native timber of our forests consists principally

of maple, (of which we have four species,) beech, birch,

hemloc, spruce, fir, and cherry. From the sugar maple

large quantities of sugar are manufactured. ...”

6. (Plantings of sugar maple)

“Several years since the practice of setting out that

very beautiful and useful tree, the sugar maple, by our

road sides, was introduced by the writer of this article.

Several of our streets are now ornamented in this way;

and it is highly desirable that the practice should become

general. ...”

176. Gatlin, George 1835

Letters and Notes on the Manners, Customs, and Con¬

dition of the North American Indians. Philadelphia, 1859,

p. 604; Thos. Donaldson, The George Gatlin Indian Gal-

156 Wisconsin Academy of Sciences , Arts and Letters

lery. Report of the U. S. National Museum. 1885, Pt. V,

p. 240.

(A Chippeway gift of maple sugar)

“Through this curious scene I was strolling a few days

since with my wife, and I observed the Indian women

gathering around her, anxious to shake hands with her

and shew her their children, of which she took especial

notice; and they literally filled her hands and her arms

with muk-kuks of maple sugar, which they manufacture

and had brought in, in great quantities for sale.”

177. Featherstonhaugh, Geo. Wm. 1885

A Canoe Voyage up the Minnay Sotor. London, 1847,

Vol. I, (a) p. 327, (b) 338.

a . (Maple sap restores Indians strength)

“The banks of the river were generally low, but occa¬

sionally immense bluffs of granite came jutting in, maple,

oak, poplar, and willow abounding. Milor informed me

that the sugar-maple was a great blessing to the Indians ;

for that often in the spring, before the snow has melted,

and they are almost reduced to starvation, they watch the

maple-tree, and as soon as the sap begins to run in March,

drink it and soon recover their strength.”

b. (Maple sugar camp near Lac qui Parle)

“ ... At 9 A.M. we stopped in a clump of sugar maple

trees to breakfast, where we found a great number of lit¬

tle wooden troughs, which the Indians, after making an

incision in the trees, place beneath them to collect the

sap. ...”

178. Fitch, Martha E. 1839

A little girl of old Milwaukee. Wisconsin Mag. Hist.,

9, 84 (1925).

(A “ sugaring off ” along the Menominee River)

“Whenever they came to a maple tree they stopped

and chopped a small piece of wood from it, pressed a

shingle into the cavity to carry the sweet maple sap that

was ready to flow into the bucket that they had left under

each tree. They found a nice open space for the great

kettle and put it on a chain, supported by rods, and later

Schuette and Ihde— Maple Sugar

157

built from the fallen trees a great fire under it. In a day

or two they commenced gathering the sugar sap, as it

was called. The men drove to all the trees and emptied

the buckets into a barrel, and then drove to the fire, empty¬

ing the barrels into the big kettle. We could drive around

with them or sit on a log and watch the boiling kettle, just

as we pleased. There was a long table with tin cups and

plates, and a long dipper to stir the syrup, and the nice

men would give us a cupful of the syrup, which we cooled

in the snow and it was just like candy. When the syrup

was boiled down sufficiently we had a ‘sugaring off.’ The

neighbors were invited. It was in the evening and the

moon was bright as day, and the big bonfire blazed and

glowed, and made us all warm and ‘comfy.’ The neigh¬

bors came, some in sleighs and some on horseback, and

boys and girls, too. They could have all the sugar they

wanted to eat. . . . And that is the way we made our

maple sugar.”

179. Trego, Charles B. 1843

A Geography of Pennsylvania. Philadelphia, 1843, p.

60.

Maple

“Of the Sugar Maple we have two kinds; the true

Sugar maple (Acer sacchannum) and the Black Sugar

tree, or Black maple ( Acer nigrum) . The former is most

abundant in the northern parts of the State, and along

the elevated range of the Allegheny table land, where the

soil, though fertile, is cold and moist. . . . The Black

maple is more common in the low rich soils along the

western rivers. . . . Both of these species of maple yield

the sap from which sugar is made. ... In February, or

the beginning of March, when the sap begins to ascend,

holes are bored in the tree from one to two feet from the

ground, and tubes of elder or sumach inserted to conduct

the sap into a trough or vessel placed to receive it. The

sap is collected and boiled to a syrup, after which it is

cooled and is strained through a cloth to separate impuri¬

ties. It is then boiled again, until the syrup is reduced

to the proper consistency for graining or pouring into the

moulds. The colour and quality of the sugar depend much

upon the care and judgment with which the process is

158 Wisconsin Academy of Sciences , Arts and Letters

conducted. The sap continues to flow for several weeks,

but gradually becomes less abundant and less rich in

saccharine matter. About four gallons of sap are estimated

to yield a pound of sugar, and a single tree, having twen¬

ty tubes inserted has been known to yield twenty-three

gallons of sap in a day. Large quantities of maple sugar

are still made in the northern and western counties by

the farmers, who sell that which they do not require for

their own use to the shopkeepers of the neighbouring

towns.”

180. Marshall, Josiah T. 1845

The Farmer’s and Emigrant’s Hand-Book. New York,

1845, 2 ed., p. 359.

(To make maple sugar)

In a letter to the Committee on Maple Sugar of the

New- York State Agricultural Society, Joel Woodworth de¬

scribes his method of making and clarifying the sugar for

which he received the Society’s first premium. As clarify¬

ing agents he uses for one hundred pounds of sugar “the

whites of four or five eggs well-beaten, about one quart of

new milk, and a spoonful of saleratus, all mixed with the

syrup before it is scalding hot.”

181. Beckiey, Hosea 1846

The History of Vermont; with Descriptions, Physical

and Topographical. Brattleboro, 1846, p. 311.

(A salute to the sugar-maple)

“The sugar maple is the glory of the Vermont forests,

so rich and beautiful in their great variety of trees and

shrubbery, and to the different heights to which they

grow, and shapes which they assume. The color of their

bark and lines and tinges of their foliage are almost end¬

less in their diversities. The form of the maple and the

intenseness of its foliage, the first to bud and leave out in

the spring, and the first to fade in autumn, renders it a

pleasing object of contemplation in itself. But the increas¬

ing use made of it for sugar and molasses, must greatly

enhance its value and comeliness in the eyes of the Ver¬

monters, on whose soil it stands pre-eminent and most

frequent.

Schuette and Ihde — Maple Sugar

159

“Pre-eminent and most frequent, this is true as a

state; although in some parts of New York, .particularly

the high-lands of Schoharie county, this noble tree is found

in magnitude and height and frequency equal to any part

of this state. Such significant names of neighborhoods

and villages are found as sap-bush-hill , and sap-hollow,

where and on dutch-hill, the writer has seen as noble

specimens of this tree as those given by Dr. Williams

in early periods of green mountain history; five feet in

diameter and from one to two hundred feet high.”

182. Butterfield, Consul W. 1848

History of Seneca County. Sandusky, 1848, p. 69.

(A Seneca festival)

“Large kettles of soup ready prepared, in which ma¬

ple sugar, profusely added, made a prominent ingredient,

thus forming a very agreeable saccharine coalescence. All

were invited, and all were made welcome; indeed, a re¬

fusal to partake of their bounty was deemed disrepect-

ful, if not unfriendly.”

183. Goodrich, S. G. 1848

Manners and Customs of the American Indians. Bos¬

ton, 1848, p. 204.

(Food of the North American Indian)

“They extracted sugar from the maple tree, and used

it to sweeten their cakes which were made of ground

corn mixed with chestnuts, beans and berries.”

184. Allen, William 1849

The History of Norridgewock. Norridgewock, 1849,

(a) p. 57, (b) p. 78.

a. (Forest trees at Norridgeivock , Maine)

“There was formerly a considerable quantity of pine

timber in the town which was distinguished for its size.

The hard wood growth originally consisted of beech, su¬

gar maple, yellow and white birch, white and brown

ash, intermixed with evergreens, of which hemlock pre¬

dominated ; spruce and cedar were also found, and in some

swampy places, hackmatack. . . . The margin of “the river

was lined with trees of various kinds, and the intervales

were covered with the white and sugar maple, the elm,

160 Wisconsin Academy of Sciences, Arts and Letters

the birch, the butternut, and the basswood ; balm of Gilead

and poplars were found in some places.”

b. (Source of sugar)

“The surrounding maples furnished them with su¬

gar; . .

185. Clark, Joshua V. H. 1849

Onondaga ; or Reminiscences of Earlier and Later

Times. Syracuse, 1849, Vol. I, p. 54.

(Rites and ceremonies)

“The first of these festivals is held in spring, directly

after the season for making sugar is past. They give

thanks for the abundance of sap, and for the quantity of

sugar they have been permitted to make.”

186. Eastman, Mary 1849

Dahcotah; or Life and Legends of the Sioux around

Fort Snelling. New York, 1849, p. 159.

(Sugar-feast is part of the Indians religious rites)

“After the scalp-dance had been performed long enough,

the Dahcotahs of the villages turned their attention to

making sugar. Many groves of sugar trees were in sight

of their village, and on this occasion the generous sap

rewarded their labors.

“Nor were they ungrateful; for when the medicine

men announced that they must keep the sugar-feast, all

left their occupations, anxious to celebrate it. Neither

need it be concluded that this occasioned them no loss of

time; for they were all occupied with the construction

of their summer wigwams, which are made of bark trees,

which must be peeled off in the spring.

“But every villager assembled to keep the feast. A

certain quantity of sugar was dealt out to each individual,

and any one of them who could not eat all that was given

him was obliged to pay leggins, or a blanket, or something

valuable, to the medicine man. On this occasion, indeed on

most occasions, the Dahcotahs have no difficulty in dis¬

posing of any quantity of food.”

187. (Cooper, Susan Fennimore) 1850

Rural Hours. New York, 1850, (a) p. 23, (b) p. 27,

(c) p. 28.

S dinette and Ihde— Maple Sugar

101

a. (The maple sugar scene in New York)

“Saturday, April 1st.— Fresh maple sugar offered for

sale today; it is seldom brought to the market as early

as this. A large amount of this sugar is still made in our

neighborhood, chiefly for home consumption on the farms.

In the villages, where foreign groceries are easily pro¬

cured, it is eaten more as a dainty than in any other

way; the children are very fond of it, and most grown

persons like a bit now and then, its peculiar flavor making

it pleasant when taken by itself, though it becomes a de¬

fect when used for sweetening food. In the spring, a lit¬

tle of it is not thought unhealthy, from a fancy that it

purifies the blood ; probably it is neither better nor worse

in this respect than any other sugar. With our farmers,

however, it is a matter of regular household consumption,

many families depending on it altogether, keeping only a

little white sugar for sickness ; and it is said that children

have often grown up in this country without tasting any

but maple sugar. Maple molasses is also very much used,

some persons preferring it to that of the cane, as it has

a peculiar flavor which is liked with puddings, or buck¬

wheat cakes.”

5. (“Year-round” production)

“A story is told in the village of a Scotch stocking-

weaver, who some years since bought a farm near the

lake, and the first spring after his arrival in the country

was so successful with his maple trees, that in the midst

of his labors he came into the village and gave large orders

for sap-buckets, pans, furnaces, &. The good folk were

rather surprised at the extent of these preparations, and

inquiries were made about this grand sugar-bush. They

were told by their new neighbor that as yet he had

tapped only a small number of trees, but he intended

soon to go to work in earnest among the maples, and, in¬

deed, had quite made up his mind, 'canny Scot/ as he was,

to 'give up farming altogether, and keep to sugar-making

all the year round' ; . . . ”

c. (Other trees yield saccharine sap)

“Many other trees are tapped for their juices . . .

they prepare from the sap of the Palm of Chili, a syrup

162 Wisconsin Academy of Sciences, Arts and Letters

of the consistency of honey. . . In Crimea, the Tartars

regularly make sugar from the fine walnut-trees on the

shores of the Black Sea. So says Dr. Clarke in his Travels.

The lime or basswood also yields a saccharine fluid. Our

own hickory is thought to have the sweetest and richest

sap of any tree in the woods, and we have heard of su¬

perior sugar being made in small quantities from it by

certain New England housewives. It would not be gen¬

erally available for the purpose, however, as the amount

of sap yielded is very small.”

The entry concludes with some production figures for

the various states.

188. Thatcher, B. B. 1854

Indian Traits: being Sketches of the Manners, Cus¬

toms, and Character of the North American Natives. New

York, 1854, Vol. I, p. 66.

(Maple sugar in Indian cookery)

“If ripe and dry, it was pounded as fine as possible in

the mortar, kneaded into dough, and made up into flat

cakes, which they were careful to bake on hot and clean

ashes. With this dough they frequently mixed boiled

pumpkins, green or dried, beans, chestnuts, dried venison

pounded to a powder, berries, and other things. Sugar,

made from the juice of the maple-tree, was in many sec¬

tions used to sweeten the rest.”

189. Myrtle, Minnie 1855

The Iroquois; the Bright Side of Indian Character.

New York, 1855, p. 49.

(Maple festival)

“The first festival was held in the spring when the

sap began to flow, to return thanks to the maple for its

sweet juices, and also to God for having given it to his

red children. ...”

190. Thoreau, Henry David 1856

Early Spring in Massachusetts. From the Journal of

Henry David Thoreau. Edited by H. G. 0. Blake. Boston,

1898, p. 199.

(Sugar from the red maple)

uMarch 21, 1856. 10 A.M. To my red maple sugar

camp. Found that after a pint and a half had run from

Schuette and Ihde— Maple Sugar

168

a single tube after 3 P.M. yesterday afternoon, it had

frozen about half an inch thick, and this morning a quar¬

ter of a pint more had run. Between 10 V2 and 11 V2 A.M,

this forenoon I caught two and three quarters pints more

from six tubes at the same tree, though it is completely

overcast, and threatening rain,— four and one half pints

in all. The sap is an agreeable drink like iced water, by

chance, with a pleasant but slightly sweetish taste. I

boiled it down in the afternoon, and it made one and one „

half ounces of sugar, without any molasses. This appears

to be the average amount yielded by the sugar maple

in similar circumstances, viz., on the south edge of a wood,

and on a tree partly decayed, two feet in diameter. It is

worth while to know that there is all this sugar in our

woods, much of which might be obtained by using the

refuse wood lying about, without damage to the propri¬

etors, who use neither the sugar nor the wood. I put in

saleratus and a little milk while boiling, the former to

neutralize the acid, and the latter to collect the impurities

in a scum. After boiling it till I burned it a little, and my

small quantity would not flow when cool, but was as

hard as half-done candy, I put it on again, and in a min¬

ute it was softened and turned to sugar. Had a dispute

with father about the use of my making this sugar when

I knew it could be done, and might have bought sugar

cheaper at Holden’s. He said it took me from my studies.

I said I made it my study and felt as if I had been to a

university. The sap dropped from each tube about as

fast as my pulse beat, and as there were three tubes di¬

rected to each vessel it flowed at the rate of about one

hundred and eighty drops a minute into it. One maple,

standing immediately north of a thick white pine, scarce¬

ly flowed at all, while a smaller one, farther in the wood,

ran pretty well. The south side of a tree bleeds first in

the spring. Had a three-quarter inch auger. Made a

dozen spouts five or six inches long, hole as large as a

pencil, and smoothed with one.”

191. Lapham, I. A. 1357

The forest trees of Wisconsin. Trans. Wisconsin State

Agr . Soc., 4, 207 (1854-7).

164 Wisconsin Academy of Sciences , Arts and Letters

(Acer Saccharinum, of Waugenheim . Sugar

Maple)

“This well known and highly valuable tree forms dense

groves in many places, but more especially in the eastern

and northern parts of the State. Some of these groves,

called ‘maple openings/ are among the most beautiful and

interesting of our forest scenery. These groves often oc¬

cupy the sites of deserted Indian villages,— thousands of

the trees are annually ‘tapped’ to draw sap for the manu¬

facture of ‘maple sugar.’ Over six hundred thousand

pounds of this sugar are annually made in Wisconsin.”

192. Hardy, Campbell 1869

Forest Life in Acadie. London, 1869, p. 41.

(Sugaries, maple honey , maple molasses)

“Before leaving the woods, however, we may not omit

to notice those characteristic trees of the American forest,

the maples, particularly that most important member of

the family, the rock or sugar maple -Acer saccharinum.

Found generally interspersed with other hardwood trees,

this tree is seen of largest and most frequent growth in

the Acadian forests on the slopes of the Cobequid hills,

and other similar ranges in Nova Scotia, often growing

together in large clumps. Such groves are termed ‘Sugar¬

ies’, and are yearly visited by the settlers for the plentiful

supply of sap which, in the early spring, courses between

the bark and the wood, and from which the maple sugar

is extracted. Towards the end of March, when winter is

relaxing its hold, and the hitherto frozen trees begin to

feel the influence of the sun, the settlers, old and young,

turn into the woods with their axes, sap-troughs, and

boilers, and commence the operation of sugar-making. A

fine young maple is selected; an oblique incision made

by two strokes of the axe at a few feet from the ground,

and the pent-up sap immediately begins to trickle and

drop from the wound. A wooden spout is driven in, and

the trough placed underneath; next morning a bucketfull

of clear sv/eet sap is removed and taken to the boiling

house. Sometimes two or three hundred trees are tapped

at a time, and require the attention of a large party of

men. At the camp, the sap is carefully boiled and evap-

Sehuette and Hide— Maple Sugar

165

orated until it attains the consistency of syrup. At this

stage much of it is used by the settlers under the name of

‘maple honey, or molasses’. Further boiling; and on pour¬

ing small quantities on to pieces of ice, it suddenly cools

and contracts, and in this stage is called ‘maple-wax’

which is much prized as a sweetmeat. Just beyond this

point the remaining sap is poured into moulds, in which

as it cools it forms the solid saccharine mass termed

‘maple sugar’. Sugar may also be obtained, though in¬

ferior in quality, from the various birches, but the sap

of these trees is slightly acidulous, and is more often

converted into vinegar.”

193. McAfee, H. H. 1870

The maple family of trees for cultivation. Trans . Wis¬

consin State Agr. Soc., 9, 288 (1870).

(Economic value of the maple)

An appraisal of the maple tree as a source of sugar

and of wood.

194. Fisk, E. A. 1874

Maple Sugar. Vermont State Board Agriculture, Man¬

ufacturing and Mining, Second Biennial Report , 1873-74,

(a) p. 713, (b) p. 717.

a . (Maple sugar production poorly understood)

“ . . . Perhaps there is no branch of farming in which

more improvement has been made within the last thirty

or forty years than in the manufacture of maple sugar;

and there is probably none in regard to which there is

more general ignorance among people who do not live

where it is made. I will not vouch for the truth of the

story of the man who on immigrating to Vermont, thought

he would follow sugaring the year round as he had heard

that it was a profitable business. But only a few days ago

I heard of two intelligent men within the limits of New

England who had a warm dispute whether maple sugar

was made in the fall or spring, and referred it to a Ver¬

monter for decision. How many pounds of sugar can

you make from a cord of maple wood? is also an authentic

question.”

166 Wisconsin Academy of Sciences , Arts and Letters

b. (Factors which influence quality)

“ . . . The location of the sugar place may have some

influence upon the quality of sugar produced, and if there

are many spruces or hemlocks to cast their leaves into

the sap, it cannot improve it any, and it is probable that

impurities in the soil may in some cases affect the su¬

gar. ... In one case a pile of spent tan bark was placed

near the roots of the maple, and the sap of that tree be¬

came the color of weak lye, but afterwards it was removed,

and the sap has since appeared to be pure. In another

case which came under my own observation, the sap was

evidently affected by impure substances in or on the

soil ; but making allowances for all this, I think that much

of the difference in sugar is in making.”

195. Foster, A. M. 1874

Sugar Making. Vermont State Board Agriculture, Man¬

ufacturing and Mining, Second Biennial Report , 1873-74,

p. 724.

(Production and marketing of maple sugar)

A report dealing with contemporary apparatus, manu¬

facturing processes, marketing, and care of sugar tools.

196. Willard, J. E. 1874

History of the Town of Sutton, Caledonia County.

Vermont Historical Gazetteer, Vol. V, iii, p. 159 (1891).

(Sutton, the banner maple sugar town)

“Sutton, it is understood, is the largest maple-sugar

producing town in the State, and perhaps, the largest in

the United States. In the spring of 1874, more than 140,-

000 pounds was made, and one year since, the produce

was larger than in 1874.

“In School District No. 6 of 12 families more than

28,000 pounds have been made in a single season.”

(Maple sugar in spirituous liquor)

“Our sugar is made nearly all of it in this County, dry

or what is known as stirred so that it is put into flour

barrels and headed up, and is now shipped to Chicago.

What they do with so much maple sugar is somewhat of

a mystery. Some say it is used in spirituous liquor.

Schuette and Ihde — Maple Sugar

167

especially for brandy which it gives the look of age, others

that it is used in glucose.”

197. Daily, Josiah 1882

Flavoring-Extract for Sirup and Sugar. United States

Patent 261,815. July 18, 1882.

Described as an “improved” sirup or sugar, claimed

to have a flavor which “cannot be distinguished from gen¬

uine maple-sirup.” To prepare it, a decoction of the out¬

side bark of the shell-bark hickory or wood — the sap of

this tree may be used if available — is added to a sirup

made from any kind of sugar, or the sirups “ordinarily

found on the market.”

198. Wiley, H. W. 1885

Composition of maple sugars and syrups. Chem. News ,

51, 88 (1885).

(Various forms of adulteration found)

The paucity of recorded analyses of maple sugars and

syrups led to a study of market samples of these prod¬

ucts with the discovery, as had long been suspected, that

the commercial products were largely adulterated. Ad¬

mixture with starch sugar, or glucose, substitution of

colored cane sugar syrup for the genuine article, and the

use of the sap of the butternut tree as the source of a

syrup, are examples of the forms of adulteration revealed

by this survey.

199. Wiley, H. W. 1885

The Sugar Industry of the United States. Maple Sugar.

U. S. Dept. Agric., Chem. Div., Bull . 5, iv, (a) p. 209, (b)

p. 218.

This publication summarizes the then known chemi¬

cal information on maple saps, sugars, and sirups. Ex¬

tensive new analytical data are recorded. Of special in¬

terest are the following items.

a. (A “ Jersey ” among maples?)

“The highest percentages of sucrose are found in tree

No. 3, April 24, viz., 9.88 per cent., and in tree No. 14,

April 20, viz., 10.20 per cent. In both of these cases the

flow of sap was small, being 128 and 227 grams, respec¬

tively.

168 Wisconsin Academy of Sciences , Arts and Letters

“The study of the sap from such a tree as No. 8 offers

also the interesting suggestion that it may^ be quite pos¬

sible to increase the percentage of the sugar in the sap of

future maples by planting the seed of such trees as show

the largest percentage of sucrose. . . . There is every rea¬

son to believe that a race of maples, yielding a large per¬

centage of sugar, could be developed as easily as a race of

cows, yielding large quantities of butter.

“Among the maples there may yet be a race of Jer¬

seys.”

b. (Manufacturing notes)

Descriptive.

200. Schultz, J. C. 1887

The Great Mackenzie Basin: A Summary of the Re¬

ports of the “Schultz Committees” of the Senate of Can¬

ada. Edited by F. J. Chambers. Ottawa, 1908, p. 16.

( Sugar from the ash-leaved or Red river maple)

“Prof. Bell explained that although the ordinary sugar

maple does not grow in the Northwest, there is a tree

there which yields sugar — the ash-leaved maple, some¬

times called the Red river maple. It is a very pretty tree,

grows rapidly and yields a rich sap. This tree grows na¬

tive in all the more southern parts of the northwest coun¬

try along the rivers, and Prof. Bell had seen it cultivated

by the missionaries where it does not grow naturally.

It is cultivated at Lac la Biche, some three hundred miles

northwest of its natural northern limit. The missionaries

at Lac la Biche cultivate it for the purpose of getting

sugar from it. This sugar is capable of being refined.

The sap contains two and a half per cent of sugar to its

weight. The Indians boil down the sap of this tree to make

sugar. It is the maple sugar of the Northwest.”

201. Sargent, C. S. 1891

Silva of North America. Boston and New York, 1891,

Yol. II, (a) p. 99, (b) p. 101.

a. (Sugar-making an Indian art)

“ . . . the making of maple-sugar was an established

industry of the Indians during the last half of the seven¬

teenth century, and before the discovery of the upper

Schuette and Hide— Maple Sugar

169

Mississippi River by Europeans (1673). Bossu, a French

officer of much intelligence who traveled in America be¬

tween 1756 and 1771, states explicitly that the French

learned the method of sugar-making from the Indians;

and the testimony of earlier travelers point to the same

conclusion.”

b. (Early botanists overlook the sugar maple)

“The Sugar Maple, strangely enough, escaped the at¬

tention of the early botanists who examined the forests

of North America, and it was not known to Linnaeus. . . .”

202. Wiley, H. W. 1892

Foods and Adulterants. Sugar, Molasses and Sirup,

Confections, Honey and Beeswax. U. S. Dept. Agric. Div.

Chem., Bull., 13, vi, (a) p. 645, (b) p. 675, (c) p. 710.

a. ( Adulterated maple sirup from Ohio)

“ . . . Among the 17 samples of maple molasses 6 were

found to be adulterated with commercial glucose. This

fact was a surprise to the writer, since two years ago the

dairy and food commission of Ohio had succeeded in driv¬

ing all of these spurious brands of maple sirup from the

State. . . . some of the (remaining) samples have a con¬

siderable proportion of reducing sugars, and at the same

time a low content of ash. In the manufacture of maple

sirup and sugar, the salts contained in the sap are not

separated from the finished product. ... It would seem

. . . that some of the samples not adulterated with glu¬

cose were contaminated with cane sugar or sirup. ...”

b. (Maple sugar valued for its flavor)

“ . . . The price of maple sugar, as is well known,

is out of all proportion to the saccharine matter which it

contains, and is due to its peculiar and pleasant taste, de¬

rived presumably from some ethereal matter exuded with

the sap. The natute of this substance has not . . . been

definitely determined. It is not wholly volatile, since it re¬

mains in the sugar and molasses after they have been

kept for a long time at a high temperature during the

process of concentration. ...”

170 Wisconsin Academy of Sciences , Arts and Letters

c. (Maple sirup widely adulterated)

“It has long been known that a large part of the maple

sirup sold in the market is made from glucose, understand¬

ing by this term the liquid product of the conversion of

starch into sugar. It is also well known that large quanti¬

ties of maple sirups are sold on the market which are

fabrications made up of other sweets, to which a little

maple molasses is added for the purpose of giving it

flavor, or, as is often the case, being entirely free from

any addition of maple product whatever. The maple flavor

is imparted to sirups by mixing with them an extract of

hickory bark, and this product has been made and sold

under the term of 'mapleine.' ” It is safe to say that per¬

haps the greater quantity of maple molasses or sirup sold

on the market is an adulteration in the true sense of the

word. . . . ”

203. Hoffman, W. J. 1893

The Menomini Indians. Smithsonian Institution. Bur.

Ethnology, 14 Ann. Report , 14, (a) p. 173, (b) p. 288, (c)

p. 315.

a . (Menomini tale about the origin of maple sugar)

“When Manabush returned empty-handed from his

hunting trip ... he and his grandmother, Nokomis, gath¬

ered together all their effects, moved away from the place

where they had dwelt, and built a new wigwam among

the trees in the new locality.

“These trees were maples, and the grandmother of

Manabush said to him, 'Now, my grandson, you go into

the woods and gather for me some pieces of birchbark ; I

am going to make sugar.' So Manabush went into the

woods and gathered some strips of birchbark to make

vessels to contain the sugar.

“The grandmother of Manabush then went from tree

to tree, cutting a small piece of wood over which the sap

ran into the vessels placed beneath. Manabush followed

his grandmother from tree to tre§, watching her and look¬

ing for the sap to drop into the vessels, but none was to

be seen. When she had gone around among the trees, and

cut holes for as many vessels as she had made, Manabush

went back and looking into the vessels saw that all of

them had suddenly become half full of thick syrup.

Schuette and Ihde— Maple Sugar

171

“Manabush dipped his finger into the syrup and tasted

it. Finding it sweet, he said, 'My grandmother this is all

very good, but it will not do to have these trees produce

sirup in this manner. The people will not have any work

if they make sugar so easily; they must cut wood to boil

the sirup for several nights, and to keep them occupied

that they may not get into bad habits; I will change all

this/

"So Manabush climbed to the very top of one of the

trees, when he took his hand and scattered water all over

the maples, like rain, so that the sugar should dissolve and

flow from the trees in the form of sap. This is why the

uncles of Manabush and their descendants always have

to work hard when they want to make sugar. Wood must

be cut, vessels must be made, and the sap that is collected

must be boiled for a long time, otherwise the people would

spend too much time in idleness.”

b. (The sugar-making season)

The season for sugar-making came when the first crow

appeared. This happened about the beginning or middle

of March, while there was yet snow on the ground. This

period of the season was looked forward to with great

interest, and, as among the Minnesota Ojibwa today, be¬

came a holiday for everybody. Each female head of a

household had her own sugar hut, built in a locality

abounding in maple trees — the Acer saccharinum— which

might or might not have been convenient to her camp,

but which was the place always resorted to by her, and

claimed by right of descent through her mother's family

and totem.

c. ( English-Menomini vocabulary)

Cake sugar, Bakwatenekan ; maple sugar molded in

the shape of small cakes; served to visitors and friends,

and also deposited in grave boxes of friends and relations

as an offering. Hard maple, Sheshikima — Acer sacchari¬

num; the species used for sugar-making ; sap of maple,

Shopomakwopo.

172 Wisconsin Academy of Sciences , Arts and Letters

204. Burroughs, John 1^95

Winter Sunshine. Boston, 1895, (a) p. 89, (b) p. 93.

a. A March chronicle

“ . . . The moment the contest between the sun and

frost fairly begins, sugar weather begins; and the more

even the contest, the more the sweet. I do not know what

the philosophy of it is, but it seems a kind of see-saw, as

if the sun drew the sap up and the frost drew it down;

and an excess of either stops the flow. Before the sun has

got power to unlock the frost, there is no sap; and after

the frost lost its power to lock up again the work of the

sun, there is no sap. But when it freezes soundly at night,

with a bright, warm sun next day, wind in the west, and

no signs of a storm, the veins of the maples fairly thrill.

Pierce the bark anywhere, and out gushes the clear-, sweet

liquid. But let the wind change to the south and blow

moist and warm, destroying the crispness of the air, and

the flow slackens at once, unless there be a deep snow in

the woods to counteract or neutralize the warmth, in which

case the run may continue till the rain sets in. . . . ”

b. (The charm of sugar -making)

“I think any person who has tried it will agree with

me about the charm of sugar-making, though he have no

tooth for the sweet itself. It is enough that it is the first

spring work, and takes one to the woods. The robins are

just arriving, and their merry calls ring through the

glades. The squirrels are now venturing out, and the wood¬

peckers and nuthatches run briskly up the trees. The

crow begins to caw, with his accustomed heartiness

and assurance; and one sees the white rump and golden

shafts of the high-hole as he flits about the open woods.

... I sympathize wtih that verdant Hibernian who liked

sugar-making so well that he thought he should follow it

the whole year. I should at least be tempted to follow

the season up the mountains, camping this week on one

terrace, next week on one farther up, keeping just on

the hem of Winter’s garment, and just in advance of the

swelling buds, until my smoke went up through the last

growth of maple that surrounds the summit.”

Schuette and Hide— Maple Sugar

173

205. Jenks, Albert Ernest 1899

The Bear Maiden. An Ojibwa Folk-Tale from Lac

Courte Oreille Reservation, Wisconsin. /. American Folk¬

lore , 15, 34 (1904).

(Maple sugar in Indian folk-tale)

“He asked the little Bear whether she could bring

back the sun. She said: 'Yes, give me two handsful of

maple-sugar and your oldest son/ With the maple-sugar

she went to the wigwam of the old woman, and, climbing

up to the top, threw the sugar into a kettle of wild rice

which the old woman was cooking. When the old woman

tasted the rice she found it too sweet, so she went away

to get some water to put in the kettle, and the little Bear

jumped down, ran into the wigwam, grabbed up the hid¬

den sun, and threw it into the sky. ...”

“Again the old chief got sick and he asked the little

Bear whether she could get him his lost horse which

was all covered with bells. She answered: 'Yes, give me

two handsful of maple sugar and your youngest son/ 99

206. Pokagon, Chief Simon 1899

O-gi-maw-kwe-mit-i-gwa-ki (Queen of the Woods).

Hartford, Mich., 1899, (a) p. 124, (b) p. 143.

a . (Corn cake and maple syrup)

“After eating our simple morning meal of 'manda-min>

(corn cakes) in 'gi-wa-ga-mis-i-gan’ (maple syrup)

dipped. ...”

b. (God's kettle)

“That kettle is still kept among us, and is now called

Man-i-to au-kick (God's kettle), and is used for boiling

on-si-ban sho-po-maw (maple sap into sugar). ...”

207. Parker, Arthur C. 1910

Iroquois Uses of Maize and Other Food Plants. N. Y.

State Museum, Museum Bull., 144, (a) p. 102, (b) p. 104.

a . (Iroquois' veneration of the maple)

“The maple tree was one of the trees venerated by the

Iroquois, It was in fact the goddess of trees and the only

one to which a stated ceremony was dedicated and to which

offerings were made. Pine, hemlock, elm and basswood

were esteemed, but the maple was a special gift of the

174 Wisconsin Academy of Sciences, Arts and Letters

Creator and every spring at the foot of the largest maple

tree in each village a ceremonial fire was built and a

prayer chanted by the Keeper of the Maple Thanksgiving

ceremony as he threw upon the embers pinches of sacred

incense tobacco. The maple tree started the year. Its re¬

turning and rising sap to the Indian was the sign of the

Creator's renewed covenant.

“The Iroquois will ever remember the maple tree, but

few now even remember the tradition of how it was

during the maple sap season, that the Laurentian Iroquois

struck their blow for freedom from Adirondack domina¬

tion and fled into northern and central New York. (One

Mohawk tradition relates that the women flung hot maple

sap into the faces of the Algonquin Chiefs and thus helped

their people in the fight for independence)."

6. (Iroquois vocabulary)

“Maple, wat da; sap, syrup, owa no gi ; sugar, owa no;

boiling sap, goste do; sap runs, sap time, o ga not; he

taps, ha ge o ta ; sap spout, nio geoda Java ”

208. Barbeau, C. M. 1912

Huron and Wyandot Mythology. Canada Geol. Surv.,

Memoir 80, No. 11, Anthropological Series , 1915, p. 110.

The maple and the woman

“The Sugar-tree-top, transfigured into a human form,

once appeared to a woman who was engaged in making

maple sugar.6

“The sweet sap from a maple-tree was changed at once,

as it still lay by the tree, into a sugar lump, as big as a

large round pebble. When the woman found it on the

wooden chip that she had driven into the tree for con¬

veying the sap into a bark tray, she picked it up and

started to eat it. A person whom she did not know sud¬

denly [appeared and] stood beside her, saying, T wish to

bring you good-luck. You must not eat the sugar-lump

but keep it in a box, so that it may not be spoilt. And

whenever you are making maple sugar, you may use it

for gathering as much syrup as you will desire. The only

. ® The Iroquoian tribes knew how to make maple syrup before the coming of the whites.

It is not certain whether the Wyandots belonging to the western band made any maple sugar

since they left the neighbourhood of Detroit, Ohio, in the course of the eighteenth century/*

Schuette and Ihde- — Maple Sugar

175

thing for you to do, when the sap is boiling, is to make

a mark in the big kettle with the [treasured] sugar-lump ;

and the syrup will fill the kettle up to that spot. Keep

this charm forever, and I will give you good fortune/ ”

209. Skinner, A. B. 1913

Social Life and Ceremonial Bundles of the Menomini

Indians. New York, 1915. Am. Museum Nat. Hist., An¬

thropological Papers , 13, vi, (a) p. 6, (b) p. 21, (c) p. 51,

(d) p. 62, (e) p. 66, (f) p. 148.

a. Home life of the Menomini

“ ... In early spring, too, there was the annual sugar¬

making festival at the camps when the toil of reducing

maple sap was lightened by merriment, dances, and buf¬

foonery.”

6. Social organization

“If a man met his totem animal he would often give

it tobacco or some of its favorite food. For instance, if

a bear, he would give it a piece of maple sugar.”

c. (A Menomini dream)

“Shanapow, when a young boy commenced fasting for

his fortune. He lived with his parents on the side hill

opposite Keshena Falls or Kakapakato. He fasted eight

days without eating, till he got very weak. On the eighth

night he dreamed that one of the sacred monsters who

lived in the falls appeared and told him, ‘Look yonder and

you will see something laced there as your reward for

fasting/ indicating a rock in the center of the falls. The

whole earth looked transparent and he went to the rock

island, going over ice. When he got there, he discovered

a sacred kettle which was bright as fire. It was a bear

kettle from the underneath god to fetch from when a

sacrifice feast was given. ‘Now/ said the god, ‘go a short

distance and you will find there what is granted you. You

will then break your fast and eat/ So Shanapow went

and found a large bear which he killed and made a sacri¬

fice of, and then ate with others whom he invited.

“The sacred kettle was to be hidden at first, for it

was too great and sacred to be seen. When maple sugar

is made it is the first thing to be placed in the sacred

176 Wisconsin Academy of Sciences , Arts and Letters

kettle, and it should be in it till a feast is made in its

honor. Then the feasters eat it in honor of the monster

below the falls. A song is then sung which is: ‘All of

the chiefs have given me to know this song/ This kettle

is called a bear god kettle and is sacred. Every spring,

maple sugar is put in it because all bears like sweet su¬

gar, especially the king bear beneath this great falls. The

dreamer Shanapow was told that he must keep a tiny bear

to fulfil his dream. He always kept a bear cubskin to set

up on a stick during the sacrifices.”

d. Months and seasons

April Sopomakwin keso Sugar-making moon

e. Burial customs

“ . . . Should the relatives of the deceased be so for¬

tunate as to have an unusually luxurious meal, or, in the

sugar season, when there is an abundance of sweets, some

tid-bits are placed in a tiny wooden bowl which is hung

up in the memory of the dead relative who is supposed

to come and eat it. . . . ”

/. Hunting customs

“When it drew near spring the parents of the lost

girl were making maple sugar at their sugar bush. Only a

little snow remained here and there, and in the evenings

the owls began to whoop and sing to show that they are

at last awake, for the Indians know that winter is but

a short night to all the Sacred Powers.

“In the meantime the parents of the little girl had

given her up as lost, but the owl said to her grandchild,

‘Now I will take you home, and land you at the limits

of your parent’s work on the trees they have tapped, sur¬

rounding their sugar camp. Stand there silently until

your mother comes and finds you. Don’t allow her to

touch you at all for four days. Then you must tell her to

go and prepare a tiny wigwam for you to remain in for

four days. This must be away from the sugar camp in a

clean place where no one has done any trampling on the

ground, and you shall remain there, silent.’

“The mother ran back to the sugar camp to tell her

husband and they both went back and met the girl.”

Schuette and Ihde — Maple Sugar

177

210. Barbeau, C. M. 1915

Huron and Wyandot Mythology. Canada Geol. Surv.,

Memoir 80, No. 11, Anthropological Series , p. 45.

(Maple in Wyandot myth regarding the

origin of the ivorld)

“The Good One . . . made all kinds of trees covered

with savory fruits, just within one’s hand’s reach. . . .

The maple was made so that syrup would just drip out

when the tree was tapped. Then came the Evil One. Find¬

ing the bushes too luxuriant and the fruits too sweet and

juicy, he spoiled them. . . . Into the maple tree he poured

some water and in that way Thinned’ the syrup into

sap, which could not be reduced into syrup without exact¬

ing labour and trouble.”

211. Skinner, A. B., and J. V. Satterlee 1915

Folklore of the Menomini Indians. New York, 1915.

Am. Museum Nat. Hist., Anthropological Papers, 13, iii,

298.

(Tales of the culture hero)

“Once in one of Manabus’ walks he followed up a

stream along the bank until he came to where a pair of

mated partridges were making maple sugar. They had two

children, little partridges, who were seated near their

nest and beside the sugar kettle. . . . Then said Manabus,

‘How do you eat your maple sugar? Let me see you, do.’

So both little birds flew to the rim of the kettle and sat

there while they commenced to eat maple sugar. Then

Manabus pushed them in and killed them. . . .

“In the meantime the old partridges returned and

when they saw their young ones in the kettle of syrup

they said : ‘Manabus must have come here.’ ”

212. Barrus, Clara 1916

The Life and Letters of John Burroughs. Boston, 1925,

Vol. II, p. 230.

C'Lock jaw”, a maple sugar confection)

The author uses the colloquial name of “lockjaw” in

referring to the sugar-gum, or wax, whose preparation

from the hot syrup before it had reached the crystalliza¬

tion stage was usually part of every maple sugar picnic.

178 Wisconsin Academy of Sciences , Arts and Letters

The art of making this confection by pouring thin sheets

of syrup of the right consistency upon clean snow had

been known for at least one hundred years.

213. Waugh, F. W. . 1916

Iroquois foods and food preparation. Canada Geol.

Surv. Memoir 86, No. 12. Anthropological Series, p. 140.

Saccharine foods: Maple Syrup and sugar

“The sap of the maple, birch, and several other trees

was employed prehistorically. Besides its use as a bever¬

age, it was boiled and thickened somewhat, though its

manufacture into sugar must have been exceedingly diffi¬

cult, if not impossible, with the crude utensils at hand.”

214. Schafer, Joseph 1922

The Yankee and the Teuton in Wisconsin. Wiscon¬

sin Mag. Hist., 6, 129 (1922).

(Maple forests in Wisconsin)

“And here we find that the distinguishing fact marking

off the region in which Germans abounded from most of

the other settled or partially settled areas of the state

was its originally thickly wooded character. In a way

almost startling, and superficially conclusive, the German

settlements coincided with the great maple forest of south¬

eastern Wisconsin, spreading also through the included

pine forest on Lake Michigan south of Green Bay.”

215. Smith, Huron H. 1923

Ethnobotany of the Menomini Indians. Bull . Public

Museum Milwaukee, 4, 61 (1923).

(An important Menomini food)

“Hard Maple ( Acer saccharum Marsh.) ‘sopoma tik’,

. . . Maple sugar, ‘sopoma tik sopomo’, is one of the most

important Menomini foods. ...”

216. Smith, Huron A. 1928

Ethnobotany of the Meskwaki Indians. Bull . Public

Museum Milwaukee, 4, 255, (1928).

(Sugar making in Wisconsin by the Meskwaki

Indians)

“Sugar Maple ( Acer saccharum Marsh.), ‘sena mislT

(cold timber). There are not many sugar trees on the

Schuette and Ihde— -Maple Sugar

179

Meskwaki reservation; hence but little sugar is made,

but they recall with considerable longing the sugar that

they used to make in Wisconsin. Most of their cooking,

even of meats, in the olden days was done with maple

sugar as the seasoning instead of salt, but now they have

to depend mostly upon salt.”

217. Jenness, Diamond 1929

The Ojibwa Indians of Parry Island, Their Social and

Religious Life. Canada Dept, of Mines, National Museum

of Canada, Bull 78, Anthropological Series , No. 17, 1935,

(a) p. 12, (b) p. 13.

a. Sugar-making moon

sizbakudikegizis, sizubakudikegizis

b. (Indian's method of concentrating sap)

“ . . . The Indians then packed their possessions to

the maple groves and tapped the trees for their syrup.

. . . The women had few idle hours though their work

was comparatively easy. They directed the flow of the

maple sap over a large sheet of birch bark, where the

warm sun hardened it to the consistency of treacle. To

harden it still further they used several methods. A hot

sun alone would reduce it to a sort of toffee, or it could

be evaporated at night before the fire. More often, per¬

haps, the women boiled it in clay pots directly over the

fire, or else in vessels of birch bark by the use of hot

stones. Impatient members of the family sometimes

dipped into the syrup heated cones made of a soft, green¬

ish stone, when the syrup crystallized on the stone and

could be scraped away with a knife. . . . ”

218. Smith, Huron H. 1932

Ethnobotany of the 0 jib we Indians. Bull. Public Mu¬

seum Milwaukee, 4, 394 (1932).

(Maple sugar and the Ojibwa Indians)

“Sugar Maple ( Acer saccharum Marsh.), Tnena tig'

(indian tree) and ‘adjagobi min.’ Both names come from

the Pillager 0 jib we, and although the trees were scarce

on the Flambeau Reservation, they also call it Tnena tig/

and gather quantities of the sap somewhere south of the

reservation. Maple sugar is one of their most important

180 Wisconsin Academy of Sciences, Arts and Letters

foods and is used in almost every kind of cookery. Maple

sap is saved to drink as it comes from the tree, some¬

times with the added sap of the Box Elder or Yellow

Birch. Again it is allowed to become sour to make a

vinegar ‘ciwabo’ used in their cookery of venison, which,

when afterwards sweetened with maple sugar, corresponds

to the German fashion of sweet-sour meat. Before they

had the salt of the white man, maple sugar took its place

and still does when they can get it. There are many in¬

teresting legends about the tree, its discovery and sugar

making, as related in Mr. Alanson Skinner’s ‘Material

Culture of the Menomini.’ The Ojibwe garner their sugar

crop much the same way as they did years ago, except

that they have used large iron kettles since the coming

of the white man. The sugar camps are rather perma¬

nent affairs, and the framework of the boiling house with

its upright poles around the fire place to hold the kettles

is left intact. A bark-covered wigwam is used to store the

tools of sap gathering, and granulation. Most of the sap

vessels and storage vessels are made of birch bark, sewed

with boiled basswood fiber or the core of the Jack Pine

root. The vessels are rendered waterproof by the appli¬

cation of pitch secured by boiling Jack Pine cones.

“In early April, the Ojibwe visit their camps, the men

to repair the camps and the storage vats of hollowed

logs, and to cut fire wood, the women to see that the sap

buckets and mokoks are scrupulously clean and watertight.

If some can not be repaired, rolls of birchbark are there to

make new ones. The whole family then move to the camp

and live in the large wigwam, while they make sugar for a

month. During the sap how, a man can chop holes and

set taps into from two to three hundred trees a day.

The first flow of sap is best, and it gets to be of a rather

poor quality by the end of the flow. The Ojibwe will not

use the night flow of the sap, which they say is bitter, so

they cease collecting an hour before dark. Gathered sap

is stored in hollowed basswood log vats, and covered over

with birch bark to keep it clean. Boiling in the iron kettles

birch bark to keep it clean. Boiling in the iron kettles

is done much as the white man does it, except that foam

Schuette and Ihde — -Maple Sugar 181

is dissipated by stirring with a fresh brush of a spruce

branch. The syrup is strained through a cloth and re¬

cooked in two or three quart quantities until it is ready

to sugar. Then, while still warm, it is poured into a wood¬

en trough, where it is pounded and crushed with a heavy

wooden paddle as it hardens. It is stored in covered birch

bark baskets called mokoks, of from twenty-five to seventy-

five pounds capacity. The sugar is graded according to

whiteness and stored away. Sap is often added to the

dregs in the kettles and a second grade sugar is secured.

To waste or spill any of the sap is considered an affront

to their deities, who punish such an act by causing the

sugar to shrink after it is made.”

219. Smith, Huron H. 1988

Ethnobotany of the forest Potawatomi Indians. Bull .

Public Museum Mihvaukee, 7, (a) p. 92, (b) p. 93 (1933).

a. (Potawomi name for maple)

“ ‘Kisinamic’ (cold tree of timber). This name con¬

notes medicinal rather than food use. The name of the

tree when it is spoken of as food, is ‘inina tig’ (Indian

tree). . . . The sugar maple and the black sugar maple

are found all over Wisconsin and were the most valuable

trees to our aboriginal brothers of any in the forest be¬

cause they furnished them their seasoning material. . . .

“In February or March sugar camp among the Indians

was one of the high spots of the year. While everybody

had to work, they all derived a good deal of pleasure from

it, especially the children who made taffy as the white

children do, cooling it in the snow, ...”

b. (Maple sap boiled in birch bark vessels)

“ . . . The boiling of sap in birch bark vessels was

quite a difficult thing to do. In those days, the original

fire had to be fed with bark of the tamarack tree. . . .

The flame must never be allowed to come in contact with

the birch bark, but the intense heat of the coals made

the sap boil.

“Indian pottery was not much better than the bark

‘mokoks/ for it was rather fragile and would not stand

rough handling or overheating.”

182 Wisconsin Academy of Sciences , Arts and Letters

SUBJECT INDEX, 7

Adulteration of maple syrup and sugar, 202

Mapleine, 202

With cane sugar, 198, 202

With flour, 97, 139

With glucose, 198, 202

With hickory bark extract, 197, 198, 202

With maple wood extract, 197

Chemical technology in sugar making

Adsorption of color by flannel, 187

Clarification with

blood, 119

egg whites, 97, 119, 120, 154, 171, 180,

187, 194, 199

egg yolks, 118

lime, 119, 120, 171

milk, 120, 180, 187, 190, 194, 199

saleratus, 180, 187, 190

Foam inhibition with

butter, 120, 143

lard, 120, 143

sweet cream, 195

tallow, 120

Precipitation of “nitre,” 194, 195, 199

Commerce in maple sugar

Barter article, 116, 144, 162

Competition with cane sugar unfavorable,

174

England forbids importation, 117

Exports from Michigan territory, 163

French production suggested, 112

Rents paid in sugar, 187

Suggested as revenue source for govern¬

ment, 170

Proposed as source for British sugar, 113

Traded by fur-traders, 101, 109, 121, 131,

135, 137, 160

Concentration of maple sap and syrup

By boiling (see sap gathering and boiling)

By freezing, 97, 120, 195

By hot stones, 217

By spontaneous evaporation, 120

By the sun, 217

Directions for, 97, 120, 143, 180, 194, 199

In birch-bark vessels, 219

In pottery, 217, 219

Temperatures at which to stop boiling sap,

199

Test for granulating point, 143

Dietary virtues of maple sugar

Antiscorbutic, 161

Corrective for excess pork in diet, 139

Might lessen desire for strong drink, 120

More wholesome than cane sugar, 120

Geographical location of maple trees

Canada, 73, 74, 75, 76, 78, 80, 118, 119, 138,

168, 192, 200

Connecticut, 117

Illinois, 174

Kentucky, 174

Maine, 138, 184

Massachusetts, 172, 175

Michigan, 125

Minnesota, 177

Missouri, 174

New Hampshire, 155, 159

New York, 118, 120

Ohio, 174

Pennsylvania, 91, 116, 118, 120, 125, 128,

179

Tennessee, 174

Vermont, 111, 127, 136, 181

Virginia, 77, 123, 138

Wisconsin, 92, 144, 145, 146, 169, 191, 214,

219

Indian festivals

Honoring Lahontan, 87

Post-season thanks, 185

Seneca, 182

Sugar-making festival, 186, 189, 203

Indian legends

Menomini, 203, 209, 211

Ojibwa, 166, 205

Wyandot, 208, 210

Indian names

adjagobi min (maple tree— Ojibwa) , 218

bakwateneRan (cake sugar — Menomini),

203

gi-wa-go-mis-i-gan (maple syrup), 206

goste do (boiling sap — Iroquois), 207

ha ge o ta (he taps — Iroquois), 207

inena tig (maple tree— Ojibwa, Potawa-

tami), 218, 219

kisinamic (maple tree— Potawatami), 219

Man-i-to au-kick (God’s kettle), 206

mocock (sugar basket), 129, 131, 140, 162

mocout (sugar basket), 135

mokok (sugar basket), 218

muk-kuk (sugar basket), 176

nio geoda kwa (sap spout — Iroquois), 207

o ga not (sap time — -Iroquois), 207

on-si-ban sho-po-maw (maple sap into

sugar) , 206

owa no (sugar — Iroquois), 207

owa no gi (sap, syrup — Iroquois), 207

ozeketa (maple tree), 118

sena mish (maple tree — Meskwald), 216

sheshikima (hard maple — Menomini), 203

shopomakwopo (maple sap — Menomini),

203

sizbakudikegizis (sugar-making moon—

Ojibwa) , 217

sopomakwin keso (sugar-making moon —

Menomini) , 209

sopoma tik (maple tree — Menomini), 215

sopoma tik sopomo (maple sugar — Menom¬

ini), 215

wat do (maple — Iroquois), 207

wattap (cedar root thread for baskets),

162

Indian tribes making sugar

Attawawas, 104

Brothertown, 169

Chippewa, 176

Iroquois, 189, 207, 213

Menomini, 132, 149, 169, 209, 215

Meskwaki, 216

Ojibwa, 137, 203, 218

Potawatomi, 219

Sioux, 186

Souties, 104

Wyandot, 150

Medicinal virtues of maple sap

Diuretic, 97

For indisposition, 85

Pectoral, 102, 104

Restorer of strength, 177

Senica remedy, 182

Stomachic, 85, 106

Tonic, 81, 177

Medicinal virtues of maple sugar

Blood purifier, 187

Chin-cough treatment, 100

Corrective for excess fat pork in diet, 139

Emollient, 97

Good for lungs, 94

Heartburn never caused by, 94

Malignant fever treatment, 120

Pectoral, 97

7 This index covers not only the quotations cited but unquoted portions of the reference

as well.

Schuette and Ihde — Maple Sugar

183

Preventative of worm-bourne diseases, 120

Proposed to lessen desire for strong drink,

120

Medicinal virtues of maple syrup

Antiscorbutic, 161

Used with maiden-hair fern, 161

Miscellaneous

Lines to the maple tree, 88, 148

Patent for making imitation maple syrup,

197

Preservation of syrup by burying, 140

Tree venerated by Iroquois, 207

Year-round basis for production of maple

sugar, 187, 194, 204

Miscellaneous products made from maple sap

Beer, 119, 120, 138, 143

Cider, 94, 118

Distilled spirit, 119, 120, 196

Maple wax (lock-jaw), 140, 192, 212, 219

Vinegar, 94, 97, 102, 104, 118, 119, 120,

138, 143, 192, 195, 218

Names used for the maple tree

Acer Canadense, folio tridendato, 97

Acer Canadense, floribus rubris, foliis ma¬

jor ibus superne viridibus, subtus ar-

genteis, lanuginosis, 97

Acer Canadense Sacchariferum, fructi mi-

nori D. Sarrazin, 90

Acer eriocarpum (river or white maple),

143, 148

Acer nigrum (black sugar maple), 143,

168, 179

Acer rubrum (red or swamp maple, plaine

or plane tree), 86, 143, 168

Acer saccharinum (see note 8), 99, 118, 120,

139, 143, 148, 152, 168, 171, 179, 192,

203

Acer saccharum (sugar, hard, rock maple

— see also A. saccharinum), 215, 216,

218, 219

Arable, 74, 75

Cotony, 76

Erable, 78, 86, 90, 97, 168

Herable, 82

Indian (see Indian names)

Maypole, 173

Sugar-wood, 99

Prices

Maple sugar, 96, 118, 128, 147, 162, 187,

194, 199

Maple syrup, 199

Priority in maple sugar production

French learned from Indians, 201

French taught Indians, 96

Indians made sugar prior to colonization,

167

Indians made syrup before whites came,

208

Production figures for maple sugar

By families, 112, 116, 120, 144, 167, 187,

194, 196

In New York, 187, 194

In Vermont, 194

In Wisconsin, 149, 191

In United States, 120, 143, 187

Rank, by states, 195

Properties of maple sap

Appeals to animals, 143

Appeals to woodpecker, 120

Components

acids, 199

albuminoids, 199

ash, 199

salts, 194

sucrose, 199, 200

starch absent, 199

Influence of environment of trees on, 194

Late flow inferior, 97, 143, 179, 218

Old trees yield stronger sap, 137

Sugar content

low in late sap, 143

low in sap of soft maple, 167

Properties of maple sugar

Comparable to

cane sugar, 97

Havana sugar, 162

Jamaican sugar, 100

Muscavado sugar, 119, 154

West Indian sugar, 85, 97, 112, 119, 120

Easily digested, 94

Flavor resembles horehound, 139

Flavoring constituent not known, 202

Insect residues absent, 120

Invert sugar present, 198

Red maple tree produces inferior sugar,

168

Refinable to white sugar, 97, 105, 119, 139,

143, 161, 171, 187, 194, 200

River maple tree gives superior sugar, 148

Variable quality, 194

Properties of maple syrup

Ash content, 202

Comparable to West Indies molasses, 100,

143

Invert sugar present, 198

Spoilage, 194

Weight per gallon, 194, 195

Sap flow

Conditions favoring, 194

Earlier in white maple, 143

Experimental observations, 97

In burning wood, 97

Influence of weather, 90, 94, 97, 99, 105,

117, 120, 143, 162, 167, 204

Rate, 120, 143, 190

Snow necessary, 90, 97, 167

South side of tree, 97, 120, 143, 190

Time, 86, 97, 99, 105, 112, 118, 120, 124,

130, 139, 143, 179, 187, 190

Sap gathering and boiling

Description, 88, 90, 100, 104, 117, 119, 139,

140, 143, 167, 170, 171, 178, 179, 187,

190, 192, 194, 195, 199

Directions for making poor sugar, 194

Duty of squaws, 150, 162, 170

Foaming stopped with hemlock or spruce

branch, 140, 218

Hanging the kettle, 140

Night flow not used by Ojibwa, 218

Progress in manufacture, 194

Straining through flannel, 117

Source of maple sugar

Geographical

Canada, 93, 139

Connecticut, 117

Delaware, 112

Illinois, 156

Indiana, 156, 171

Iowa, 164, 165

Maine, 148, 184

Massachusetts, 175

Michigan, 162

New Hampshire, 95

New York, 187, 194, 195

Ohio, 114, 195, 202

Pennsylvania, 116

8 The name Acer saccharinum, L. is used to designate the soft, white, or silver maple.

The correct term for the hard, rock, or sugar maple is Acer saccharum. Marsh, first used by

Humphry Marshall in his Arbustrum Americanum: The American Grove (Philadelphia, 1785).

Acer saccharinum has been erroneously used to designate the true sugar maple as well as

the silver maple and the references cited apparently refer to the true sugar maple.

184 Wisconsin Academy of Sciences , Arts and Letters

Vermont, 95, 122, 124, 127, 136, 194, 195,

196

Wisconsin, 87, 92, 103, 121, 126, 129, 133,

144, 145, 146, 147, 149, 151, 154, 160,

165, 169, 191, 216, 219

Species other than sugar maple tree yielding

sweet sap

Ash, 94

Ash-leaved maple, 86, 200, 218

Basswood, 187

Birch, 93, 105, 193, 213, 218

Box elder (see ash-leaved maple)

Black maple, 143

Butternut, 198

Hickory, 187

“Other trees,” 108

Red maple (plane tree), 94, 143

River maple (white maple), 143, 148

Walnut, 94, 187

Wild cherry, 94

Sugar camps

Description, 140, 154

Easter celebrated at, 154

Family rights to (Indian), 137

Festivities, 140

Indian, 132, 137, 169, 218

Life at, 154

Menomini-Brothertown Indian disagree¬

ment over, 169

Sugar making accessories

Birch-bark vessels, 99, 140, 150, 154, 218

Care of, 194, 195

Casseau (bark sap receptacle), 140

Draw tub, 194, 199

Elm bark tubs, 125

Evaporators, 194, 199

Faucets, 122

Flannel for straining syrup, 140, 143, 199

Gauge (yoke for carrying two sap pails),

140

General, 194, 195

Gouge, 148

Gouttiere (basswood spout), 140

Moose-skin sap vats, 99

Moulds, 105, 140, 143

Russia iron sap pans, evaporators, 195, 199

Spile, 105, 187

Tin buckets, 194

Wooden buckets, 187

Sugar making a “home” industry

Means of destroying slavery, 112, 116

Not made by slave labor, 120

Off-season agricultural product, 116, 119,

195, 199

Potential sugar supply for U. S., 116, 187

Production economically superior to West

Indies sugar, 120

Production as substitute for cane sugar,

112

Sugar making season

Time of, 102, 143, 158, 167, 209, 218, 219

Technique of sap gathering

Collecting sap, 119, 187, 194

Conducting troughs, 119, 187

Tapping the tree, 94, 97, 99, 102, 103, 105,

118, 119, 120, 143, 148, 178, 187, 192,

194, 195, 199

Use of maple sap by the Indians

Beverage, 177, 213, 218

Hot sap as a weapon, 207

Use of sugar by Indians

Beverage preparation, 157

Burial with dead, 203, 209

Festival dish, 96, 182, 186

Seasoning of meats, 96, 125, 135, 216, 218

Sweetening of bread, parched corn, etc.,

91, 125, 141, 183, 188

Sale or trade, 103, 118, 126, 176

Traded for whiskey, 147, 149

War ration, 150

Use of maple sugar and syrup by white men

Celebrations with, 115

Emergency food. 111

To sweeten

buckwheat cakes, 187

coffee, 143

crepes susettes, 140

puddings, 187

strawberries, 88

tea, 143

wild rice, 137

Use of syrup by Indians

For flavoring raccoon meat, 150

Taffy made by cooling on snow, 219

“Wash” for meat, 87

Wisconsin localities mentioned

Fox River, 169

Green Bay, 87, 103, 144, 147, 149, 151, 154,

160

Jaune (Yellow) River, 129

Little Kaccalin, 169

Milwaukee, 121, 126

Oconto Falls, 169

Prairie du Chien, 133, 165

Rice Lake, 145

Sandy Lake, 146

Swamp River, 92

Yield

Of sap from tree, 94, 97, 100, 112, 120, 143,

167, 170, 187, 190

Of sugar from sap, 97, 112, 117, 148, 161,

170, 179

Of sugar from tree, 105, 116, 118, 119, 120,

143, 148, 167, 171, 187

ACIDITY OF SOIL AND WATER USED IN

CRANBERRY CULTURE

Neil E. Stevens

Department of Botany , University of Illinois

That cranberries are grown in Wisconsin under a much

wider variety of conditions than in the eastern states has long

been recognized. In no way is this more strikingly evidenced

than in the range of acidity of the soil and water used in their

culture. This is clearly shown in Figure 1 which is based in

part on data assembled by the late L. M. Rogers. It will be

noted that the pH of soil planted to cranberries ranges from

3.6 to 6.8 and that of the water used in flooding from 5.0 to

8.5. The chart also shows that while in general acid soil and

acid water tend to be found together, there are some exceptions.

In one case a marsh having soil of pH 4.0 used water with pH

7.8. Another marsh (only a few years old) having soil with

pH slightly over 5.0 uses water with a pH above 7.5.

In striking contrast to this C. S. Beckwith, who was for

many years in charge of the Cranberry Substation of the New

Jersey Experiment Station, informed me in a personal con¬

versation in April, 1943, that the pH of 90 per cent of New

Jersey's cranberry soils is between 4.0 and 5.0 with a maximum

alkalinity of about 5.5 and that the water used for flooding

cranberries in that state is very uniform at about 4.5.

In 1940 attention was called to the apparently harmful ef¬

fects of the use of alkaline water on cultivated cranberries in

Wisconsin (1). Continued observation in that state added fur¬

ther evidence of the importance of this relation (2). The eco¬

nomic importance in Wisconsin and the general scientific inter¬

est of this subject should assure a serious experimental study

of the whole problem. A desirable preliminary, however,

seemed to be to determine whether any comparable conditions

exist in southeastern Massachusetts, by far the most important

cranberry producing region in the world. For this purpose a

185

186 Wisconsin Academy of Sciences , Arts and Letters

survey of water supplies used in flooding cranberries in Massa¬

chusetts was made during July and August, 1945. Through the

generous assistance of the workers at the Cranberry Station,

H. F. Bergman, H. J. Franklin, and J. L. Kelley, and the officers

of the New England Cranberry Sales Company, it was possible

to reach bogs widely scattered through the cranberry growing

area even during the period of gasoline rationing.

SOIL pH

3.5 4 4.5 5 5.5 6 6.5

Fig. 1. — The approximate pH of soil and flooding water of Wisconsin

cranberry properties.

Stevens— Cranberry Culture

187

TABLE 1

Comparison of Flooding Water Used on Cranberries in

Wisconsin and Massachusetts

During the summer water from about 150 sources was

tested. The number of cranberry bogs on which the water

is used would be much larger since water from certain ponds

and streams is used on several different bogs. Information

gathered during the past seven years was available regarding

flooding water used in approximately 100 cranberry properties

in Wisconsin. Direct comparison between these and the first

100 samples taken in Massachusetts is made in Table 1. (3) It will

be noted in columns 1, 2, and 3 that the largest number in both

regions falls in the category having pH 8.1 to 7.0. The simi¬

larity ends there, however, for while all but four of the water

samples in Massachusetts have a pH of 7.0 or below, the re¬

maining waters in Wisconsin are almost equally distributed

between those below pH 6.1 and those above pH 7.0. In all

fairness it should be said that a number of the Wisconsin bogs

listed as having water of high alkalinity have been abandoned

for some years.

The contrast in water between eastern Massachusetts and

Wisconsin is even more strikingly shown in columns 4 and 5

which give the range of bound carbon dioxide (4) in the water

of the two regions. It is obvious that all the flooding water thus

far tested in southeastern Massachusetts is very low in car¬

bonates. Exactly one half of the first 100 water supplies on

which these Massachusetts data are based showed a bound carbon

dioxide content of three parts or less per million. All except

11 have five parts per million or less and would thus fall in the

188 Wisconsin Academy of Sciences , Arts and Letters

category 'Very soft/’ as set up by Professor Juday, and widely

used in Wisconsin.

Two comments may be added. Water sources falling in

Juday’s classes "medium hard” and "hard” were found in

southern Vermont and adjacent Massachusetts but so far as

could be learned these have never been used in cranberry

culture.

The foregoing information tends to show why none of the

cranberry problems which are believed to be associated with the

use of hard water in Wisconsin have been recognized in Massa¬

chusetts.

Literature Cited

1. Stevens, N. E., L. M. Rogers, and H. F. Bain. 1940. Alkaline flooding

water in cranberry growing. Trans. Wisconsin Acad. Sci. 32:351-360.

2. Stevens, N. E. 1944. Further observations on alkaline flooding water

in cranberry growing. Trans. Wisconsin Acad. Sci. 36:395-397.

3. Stevens, N. E. 1946. Observations on flooding waters used in cranberry

culture. Mass. Agr. Exp. Sta. Bull. 433:37-51.

4. Birge, E. A. and Chancey Juday. 1911. The Inland Lakes of Wiscon¬

sin. Wisconsin Geological & Natural History Survey. Bull. 22, 259 pp.

PRELIMINARY REPORTS ON THE FLORA OF

WISCONSIN. XXXIII.* RANUNCULACEAE

Norman C. Fassett

These data are based on the material in the herbaria of the

University of Wisconsin and the Milwaukee Public Museum, and

much of the Wisconsin material in the Chicago Museum of Nat¬

ural History. Another source of information has been also used

for the aquatic species; Mr. Neil Hotchkiss has sent me the rec¬

ords of his collections in Wisconsin made in connection with his

work for the Fish and Wild Life Service. On a few maps open

circles indicate sight records by Mr. Hotchkiss.

The numbering of genera follows that of dalla Torre & Harms.

Much of the work in preparing this paper was done under a

grant from the Wisconsin Alumni Research Foundation.

Nymphaeaceae. Water Lily Family

a. Petiole attached at the middle of the blade

6. Blades round, 20~40 cm. in diameter . 2508. Nelumbo.

b. Blades oval, 10 cm. or less in diameter . 2510. Brasenia.

a. Petiole attached at the base of a deep notch

e. Flowers white or pinkish; veins of leaf radiating

from attachment of petiole . 2513. Nymphaea.

o. Flowers yellow; veins coming from the midrib . 2514. Nupha/r.

2508. Nelumbo. American Lotus

N. lutea (Willd.) Pers. In sloughs and pools along the Mis¬

sissippi River north to the mouth of the Chippewa River, and

elsewhere in ponds in the southern and western part of the

state. The most northern station, in Polk County, is not along

the St. Croix River but on East Lake some five miles from the

river. The range in Wisconsin has doubtless been extended by

the Indians. Map 1.

* Mr Kruschke’s Preliminary Report on the Boraginaceae in Volume 36 was erroneous¬

ly numbered XXXI ; the number should have been XXXII.

189

190 Wisconsin Academy of Sciences, Arts and Letters

The name Nelumbo pentapetala (Walt.) Fernald, Rhodora

36 :23. 1934, has recently been substituted for N. lutea. The Lotus

in Wisconsin has many petals and they are so definitely yellow

that it is easy to recognize a colony of Lotus in flower from a

distance of several miles by color alone, when seen from a bluff

overlooking the Mississippi River; if Walter’s Nymphaea pent a-

petala, with 5 white petals, was the same species as that of the

Middle West it must have been, as Jones, FI. Ill. 132. 1945, has

suggested, based on a monstrosity.

2510. Brasenia. Water Shield

B. Schreberi Gmel. In lakes, mostly northward, coming

south in the eastern half of the state. Map 2.

2513. Nymphaea. Water Lily

Nymphaea is the white-flowered Water Lily, called Castalia

in the seventh edition of Gray’s Manual and in Britton &

Brown’s Illustrated Flora; the genus called Nymphaea in those

works is now known as Nuphar. Much of our material from

Wisconsin has been identified by Professor H. S. Conard.

a. Flowers mostly 12 cm. or less broad, fragrant, with

petals more or less pointed; sepals and lower leaf-

surfaces often purple; petiole not streaked with pur¬

ple . 1. N. odorata.

a. Flowers larger, not fragrant, with petals pointed;

sepals and leaf-surfaces without purple; petioles with

several purple streaks . 2. N. tuberosa.

1. N. odorata Ait. Throughout the state in quiet water of

lakes and streams. Map 3.

2. N. TUBEROSA Paine. Apparently throughout the state and

as common as the other species. Map 4.

2514. Nuphar. Spatterdock

This is the yellow Water Lily or Cow Lily, called Nymphaea

in Gray’s Manual and the Illustrated Flora; it also appears in

the literature as Nymphozanthus.

a . Anthers shorter than the filaments; blades of leaves

mostly less than 20 cm. long

b. Flower 2 cm. or less wide; stigma with less than

10 rays; young fruit without a ring of decaying

stamens; depth of sinus about 2/3 the length of

the midrib . . . 1. N. microphyllum.

Fassett- — Flora of Wisconsin. Ranunculaceae

Nymphaea tuberosa

Nuphar microphyllum

Nuphar rubrodiscum

Nuphar advena

192 Wisconsin Academy of Sciences, Arts and Letters

b. Flowers 3 cm. or more wide; stigma with more

than 10 rays; young fruit with a ring of decay¬

ing stamens; depth of sinus about 1/2 the length

of the midrib . 2. N. rubrodiscum.

a. Anthers as long as the filaments or longer; blades of

leaves mostly 20 cm. or more long

c. Leaves usually erect, the petiole oval in cross-

section, blade with a widely open sinus; flowers

usually not marked with red . 3. N. advena.

c. Leaves usually floating, the petiole flattened on the

upper surface and with 2 narrow wings, blade with

a narrow sinus or overlapping lobes; base of sepals

and fruit often marked with red . . . 4. N. variegatum.

1. N. microphyllum (Pers.) Fernald. In small lakes with

muck or Sphagnum margins, northern Wisconsin. Map 5.

2. N. rubrodiscum Morong. Northern Wisconsin. Map 6,

dots.

3. N. advena Ait. This species appears to be rare in south¬

eastern Wisconsin and to grade into the next. Two collections

from Whitewater, Walworth County, have rounded petioles; in

one the sinus is open and in the other the basal lobes overlap,

while in both the sepals are marked with red. Some of the leaves

were erect, others were floating. Likewise, a collection from

Cedarburg, Ozaukee County, has rounded petioles and wide open

sinuses, but the sepals are marked with red. The species is re¬

ported (as Nymphaea advena) from Delavan, Walworth County,

and Green Bay, Brown County, by Miller & Standley, Contrib.

U. S. Nat. Herb. 16, pt. 3 : 86. 1912. Map 6, crosses.

4. N. variegatum Engelm. Common except in the Driftless

Area, where it is abundant only along the Mississippi River.

Map 7.

CERATOPHYLLACEAE. Hornwort Family

2516. Ceratophyllum. Coontail

a. Divisions of leaves ribbon-like, from very fine to al¬

most 1 mm. wide, toothed . 1. C. demersum

a. Divisions of leaves thread-like and tapering to a

hair-like tip, scarcely or not at all toothed . 2. C. echinatum

1. C. demersum L. Usually in hard water, throughout the

state except perhaps in some parts of the Driftless Area. Map 8.

Fassett— Flora of Wisconsin * Ranunculacme

193

2. C* ECHINATUM Gray. C, demersum var. echinatum of

Gray's Manual; see Muenscher, Am. Journ. Bot. 27: 231-233.

1940, and Fernald, Rhodora 43: 551-552. 1941. Uncommon but

widespread in the state. Map 9.

RANUNCULACEAE. Buttercup Family

The species of this family were mapped in 1930 by Lois

Almon, Preliminary Reports XI., Trans. Wis. Acad. 25 : 205-214.

The maps are not repeated here, but changes in nomenclature

and additions to our knowledge of their occurrence in the state

are noted.

a. Plants aquatic or in wet places

b. Petals present, yellow or white; sepals green. .. .2546. Ranunculus .

b. Petals absent; sepals yellow or whitish . . 2524. Caltha.

a. Plants of uplands

c. Leaves simple or once compound

d. Herbs, not climbing

e. Flowers yellow

/. Leaves simple, or compound with leaf¬

lets longer than broad ............... 2546. Ranunculus.

f. Leaves with 3 leaflets, each as broad

as long .... - ......................... 2534. Coptis.

e. Flowers white, green or blue

g. Flowers radially symmetrical

h. Flowers borne on naked scapes

i Leaves as broad as long, 3-lobed . 2541B. Hepatica.

i. Leaves grass-like or tongue-like . 2543. Myosurus.

h. Flowering stems with leaves or a

whorl of leaf-like bracts

j. Leaves or their divisions pointed

at tip

k. One leaf sessile and close to

the flower, the other petioled

and at some distance below. 2522. Hydrastis,

k. Leaves or bracts whorled, or

sessile and solitary . . 2541. Anemone

j. Leaves rounded at tip ....... 2541A. Anemonella.

g. Flowers bilaterally symmetrical

l Flowers with a spur .............. 2539. Delphinium .

1. Flowers with a hood ............... 2540. Aconitum.

d. Stems climbing or trailing _ ................ 2542. Clematis .

194 Wisconsin Academy of Sciences , Arts and Letters

Ranunculus reptans

R. reptans v. ovalis

Ranunculus Gmelini

X var. terrestris

-ff. Purshii

Jeffej^spnie diphylla

Fassett— Flora of Wisconsin. Ranunculaceae

195

c. Leaves twice or more compound

m. Petals with long spurs . 2538. Aquilegia.

m. Petals without spurs, often absent

n. Flowers many in a raceme or panicle

o. Leaflets with many sharp teeth . 2537. Actaea.

o. Leaflets entire or with a few teeth or

lobes . 2548. Thalictrum.

n. Flowers solitary or a few together

p. Divisions of leaves ribbon-like . 2541. Anemone.

p. Divisions of leaves not ribbon-like

q. Stem with a whorl of leaves . 2541A. Anemonella.

q. Stems with scattered leaves . 2532. Isopyrum.

2522. Hydrastis. Goldenseal

H. canadensis L. Probably once more abundant, but now

rather rare in rich woods in the southern half of the state.

2524. Caltha. Marsh Marigold

a. Plants tufted with nearly erect stems and bright

yellow flowers . . . . . 1. C. palustris.

a. Plants with floating stems and white or pinkish

flowers . . 2. C. natans.

I. C. palustris. Throughout the state in swamps and marshes.

2. C. natans Pall. Foxboro, Douglas County.

2532. Isopyrum. False Rue Anemone

I. biternatum (Raf.) T. & G. Rather local in rich woods in

the southern half of the state. This species ordinarily has the

lobes of the leaflets rounded ; f . acutilobum Fassett,1 with point¬

ed lobes, has been collected in Dodge County.

2534. Coptis. Goldthread

C. trifolia (L.) Salisb., var. groenlandica (Oeder) Fas¬

sett.2 In bogs and moist woods, of general occurrence except

southeast of a line drawn from Polk to Walworth Counties.

I I, biternatum (Raf.) T. & G., f. acutilobum, n.f., lobis foliolarum acutis. — Lime¬

stone ledge, Horicon, Dodge County, Wisconsin, June 8, 1929, Fassett no. 8393 (TYPE in

Herb. Univ. of Wis.).

2 In 1929 Professor Fernald pointed out that the typical C. trifolia is a plant of

northeastern Asia and Alaska while the plant of northeastern America and Greenland

differs in what Dr. Hulten later characterized as “several although not very striking

respects." The chiefly American plant was accordingly called C. groenlandica (Oeder)

Fernald, but Fernald commented: “Completely isolated as they are, they constitute two

very strong geographic varieties or, presumably, two distinct species. Until more transi¬

tional material than we yet know comes to hand it seems more reasonable to treat

them as two species, which, however, may eventually be merged.” In 1937 Dr. Hulten

fulfilled this prediction by calling attention to transitional material in southeastern

Alaska and reducing C. groenlandica to a subspecies. In the varietal category it becomes :

C. trifolia (L.) Salisb., var. groenlandica (Oeder) n.comb. Anemone groenlandica

Oeder, FI. Dan. iv. fasc. x. 5, t. dlxvi. 1770. Coptis groenlandica Fernald, Rhodora 31:

142. 1929. C. trifolia subsp. groenlandica Hulten, FI. Aleutian Is. 178. 1937.

196 Wisconsin Academy of Sciences , Arts and Letters

2537. Actaea. Baneberry

a. Fruiting pedicels slender; summit of ovary, in flower,

narrowed to about one half the diameter of the stigma

b. Mature berries red . 1. A. rubra.

b. Mature berries white . /. neglecta.

a. Fruiting pedicels thickened; summit of ovary, in

flower, narrowed only to 3/4 or more the diameter

of the stigma

c. Berries white . . . 2. A. pachypoda.

c. Berries red or pink

d. Berries and pedicels dark red . . . f. rubrocarpa.

d. Berries and pedicels pink . f. microcarpa.

1. A. rubra (Ait.) Willd. Common in woods throughout the

state. F. neglecta (Gillman) Robinson often occurs with the

red-berried form. A. rubra var. gigantea Gates was reported

from Eau Claire County by Miss Almon; A. rubra shows a gra¬

dation in size of leaflets from those only 3 cm. long to those

more than 10 cm. long, but there seems to be no “break” in the

series. The larger ones tend to be truncate to cordate at base

and the smaller ones are usually cuneate, but there is no very

good correlation.

2. A. pachypoda Ell. A. alba of Preliminary Report and of

many authors ; see Mackenzie, Torreya 28 : 53. 1928, and Fernald,

Rhodora 42 : 260-265. 1940. Widely distributed in the state but

somewhat local. F. microcarpa (DC.) Fassett3 has been collected

in Iowa County. F. rubrocarpa (Killip) Fernald is presumably

the identity of a specimen from Grant County with thickened

pedicels but labelled “Red Baneberry” by the collector.

2538. Aquilegia. Columbine

A. canadensis L. Throughout the state in woods, on cliffs

and shores, etc. F. flaviflora (Tenney) Britton, with yellow

flowers, has been collected in woods on a hillside bordering the

Wisconsin River in Dane County.

3 A. pachypoda f. microcarpa (DC.) n. comb. A. bra-chypetdla DC., 8 microcarpa

DC.. Syst. 1: 385. 1817; Fernald, Rhodora 42: 261. 1940. This seems to be the plant re¬

ported by the writer, in Rhodora 34: 96. 1932, as A. alba f. rubrocarpa, with the follow¬

ing comment: “T**e berries, rhacbis and pedicels are pink, and the thickness of the

pedicels is intermediate between that of A. alba and that of A. rubra. The characters

of the plant strongly suggest that it is a hybrid between the two species.”

197

Fassett — Flora of Wisconsin . Ranunculaceae

2589. Delphinium. Larkspur

a. Introduced annuals of vacant lots, etc.; pistil one

b. Pedicel shorter than bract; fruit glabrous . 1. D. Consolida.

b. Pedicel longer than bract; fruit pubescent . 2. D. Ajacis.

a. Native perennial on northwestern prairies; pistils

three . . . . . 3. D. virescens.

1. D. Consolida L. Milwaukee and Racine and doubtless

elsewhere, escaping* from cultivation.

2. D. Ajacis L. Milwaukee and Janesville and doubtless else¬

where, escaping from cultivation.

8. D. virescens Nutt. D. Penardi of Preliminary Report; see

Perry, Rhodora 39 : 20-22. 1937. Sand prairies and prairie hill¬

sides, Douglas to La Crosse and Monroe Counties. While this

species is very variable where its range overlaps that of D. Caro¬

linians (see Steyermark, Spring Flora of Missouri, p. 203),

it is quite constant in its characters in Wisconsin, the stem be- ,

ing always glandular-pubescent above and crisp-pubescent to¬

ward the base.

2540. Aconitum. Monkshood

A. NOVEBORACENSE Gray, var. quasiciliatum Fassett, Rho¬

dora 31 : 49. 1929. In addition to the two localities in Wisconsin

cited in the original description this has now been collected

at Parfrey’s Glen in Sauk County a few miles within the glaci¬

ated part of the Baraboo Hills, and at Coon Valley in Vernon

County near the center of the Driftless Area. A. uncinatum is

reported from Wisconsin in the seventh edition of Gray’s Manual

and in Britton & Brown’s Illustrated Flora; there are no speci¬

mens from this state and there is in the Gray Herbarium a note

from Lapham stating that his report of A. uncinatum from Wis¬

consin was an error.

2541. Anemone. Anemone

a. Segments of the involucre ribbon-like, not toothed

6. Involucre and stem densely covered with long

silky hairs . 1. A. patens var. Wolfgangiana.

b. Involucre and stem with a few scattered hairs. . .

. . . 2. A. multi fida var. hudsoniana.

a. Segments of the involucre toothed

c. Sepals 10-20

3 .A. caroliniana.

198 Wisconsin Academy of Sciences , Arts and Letters

c. Sepals 4-7

d. Sepals hairy on the back

e. Involucral leaves petioled

/. Divisions of involucral leaves narrow,

toothed only above the middle; fruiting

head several times as long as thick;

pistils densely woolly in flower so that

only the tips of the stigma are visible;

peduncles usually naked above the in¬

volucre . . . 4. A. cylindrica.

/. Divisions of involucral leaves ovate,

toothed to below the middle ; fruiting

heads not more than twice as long as

thick; pistils silky in flower with the

whole style plainly visible; all but the

earliest peduncle with secondary invo¬

lucral leaves near the middle

g. Anthers 0.7-1. 1 mm. long; fruiting

, head 7-11 mm. thick with styles

ascending at about 45 degrees

h. Sepals white and petaloid . 5 .A. riparid.

h. Sepals greenish and leathery . f. inconspicua.

g. Anthers 1. 2-1.6 mm. long; fruiting

head 12-15 mm. thick with styles

spreading at about 90 degrees

i. Sepals greenish or yellowish and

leathery . . . 6. A. virginiana.

i. Sepals white and petaloid . f. leucosepala.

e. Involucral leaves not petioled . 7. A. canadensis.

d. Sepals not hairy . 8 .A. quinque folia var. interior.

1. A. PATENS L., var. Wolfgangiana (Bess.) Koch. Basque

Flower. Locally abundant on original prairie southwest of a

line drawn from St. Croix County to Milwaukee County, and

rarely northwestward to southern Douglas County.

2. A. multifida Poir., var. hudsoniana DC. ; Fernald, Rho-

dora 19 : 141. 1917. Collected at Elkhart Lake, Sheboygan Coun¬

ty, in 1909.

3. A. caroliniana Walt. Rare on sand terraces in Eau

Claire, Pierce and La Crosse Counties.

4. A. cylindrica Gray. Candleweed. Open woods or prairies

and sand plains in the southern half of the state, rarely north¬

ward to Washburn and Vilas Counties.

5. A. riparia Fernald. Most commonly in the north but some¬

times occurring in southern Wisconsin. F. INCONSPICUA Fernald,

Rhodora 19: 140. 1917, is rarely found.

Fassett— Flora of Wisconsin. Ranunculaceae 199

6. A. virginiana L. Thimbleweed. Throughout the state,

perhaps a little more abundant southward. F. leucosepala

Fernald, l.c., is rarely found.

7. A. canadensis L. Roadside turf and moist open ground

throughout the state.

8. A. quinquefolia L., var. interior Fernald, Rhodora 37:

260. 1985. Wood Anemone. In woods throughout Wisconsin.

2541A. Anemonella. Rue Anemone

A. thalictroides (L.) Spach. In woods southwestward,

northeast to Pierce, Dunn, Eau Claire, Monroe, Columbia,

Dodge and Milwaukee Counties.

A. thalictroides f. Favilliana Bergseng4, with the stamens

and pistils all replaced by sepals, is rare. The specimen from

Lake Mills has pink flowers, while it has been reported with

white flowers near Pewaukee, Waukesha County.

2541B. Hepatica. Hepatica

One species is usually abundant locally to the exclusion

of the other; for example, in Dane County only H. acutiloba

is found in the vicinity of Madison, while H. americana occurs

in the northwest corner of the county. An occasional individual

appears to be intermediate.

1. H. acutiloba DC. Sharplobe Hepatica. Common in woods

north to Barron, Marathon, Shawano, Marinette and Door

Counties.

2. H. americana (DC.) Ker. Roundlobe Hepatica. H. tri¬

loba of American authors; see Fernald, Rhodora 19: 45. 1917.

Locally abundant throughout the state.

2542. Clematis. Clematis

a. Flowers in a compound inflorescence, less than 1 cm.

long, whitish . 1. C. virginiana.

a. Flowers solitary, 4-5 cm. long, purple . 2. C. verticillaris.

1. C. virginiana L. Virgin’s Bower. Thickets and open

ground, often climbing over shrubs.

4 In May, 1946, Mr. Stoughton Faville of Lake Mills sent to Mrs. M. S. Bergseng,

Herbarium Assistant at the University of Wisconsin, a double-flowered Anemonella for

which she supplies the following description : Anemonella thalictroides f. Favilliana, n.f.,

floribus plenis.— -Fence corner in field, Lake Mills, Wisconsin, May 1, 1946, Stoughton

Faville (TYPE in Herb. Univ. of Wis.).

200 Wisconsin Academy of Sciences, Arts and Letters

2. C. VERTiciLLARis DC. Purple Clematis. Woods and rocky

places, not very common, in the northwest part of the state

southward to St. Croix, Clark, Marathon, Lincoln and Oneida

Counties, and locally southward to the Dalles of the Wis¬

consin River and Parfrey’s Glen in Sauk County, and Blue

Mounds in Dane County.

2543. Myosurus. Mousetail

M. minimus L. Probably adventive, Pharmacy Garden of

University of Wisconsin, Madison.

2546. Ranunculus. Buttercup

a. Plants floating in the water, or prostrate on the mud

and rooting at the nodes, or connected by horizontal

rootstocks

b. Stem-leaves compound or deeply cleft

c. Flowers white; leaves with thread-like divisions

d. Leaves very limp and collapsing when taken

from the water, with a petiole between the

stipule and the division into 3 parts .... 1. R. trichophyllus.

d. Leaves stiff and keeping their shape when

taken from the water, without petioles and

dividing into 3 parts at the stipule . . . 2. R. longirostris.

c. Flowers yellow; divisions of leaves flat (but

sometimes ribbon-like and very narrow)

e. Divisions of leaves narrowly ribbonlike, or,

if broader, with the base of each division

of the same texture as the rest

/. Petals 6-17 mm. long; fruits (including

the beak) 2. 5-3. 5 mm. long, with a

wing-margin; blades of leaves 1.5-10 cm.

long

g. Leaves dissected into many narrow

segments 2 mm. or less wide; plants

not hairy; submersed forms . 4 . R. flabellaris.

g. Leaves with broader divisions; plants

often hairy; forms stranded on the

. f. riparius.

f. Petals 3.5-5 mm. long; fruits (including

the beak) 1.5-3 mm. long, not wing-

margined; blades of leaves 1-2 cm. long

h. Leaves with divisions several mm.

wide; plants often hairy; forms grow¬

ing on the mud . 5. R . Gmelini var. terrestris .

F ass e tt— Flora of Wisconsin . Ranunculac eae

201

h. Leaves with divisions 1 mm. or less

wide; plants not hairy; submersed

forms .................................. f. Purshii .

e. Leaves divided into 3 leaflets each of which

has a distinct stalk of different texture

from the flat part of the blade

i. Stems hairy

j, Hairs appressed ..................... 13. R. repens,

j. Hairs spreading ...................... var. villosus.

i. Stems glabrous or nearly so . . var. glabratus.

b . Leaves simple, flat or thread-like

k. Blades of leaves truncate or slightly heart-

shaped at base, with toothed margins

l. Stems without hairs . . 3. J?. Cymbalaria.

L Stems closely hairy . . . .... f. hebecaulis.

k. Blades of leaves thread-like, or if flat tapered

at base, with entire margins

m. Leaves thread-like . . . . 6. R. reptans,

m. Leaves with flat blades 2 mm. or more

wide . . var. ovalis.

Plants not connected by runners, with basal leaves,

and stems which are usually erect (prostrate but not

rooting in one species)

n. Petals inconspicuous, shorter than or barely ex¬

ceeding the sepals

o. Styles hooked

p. Stems with spreading hairs ............ 10. R. recurvatus.

p. Stems glabrous . . f . laevicaulis.

©. Styles not hooked

q. Basal leaves simple (occasionally one of

them 8-parted) ; head of fruits globular

r. Stems and peduncles glabrous ............ 9. R. ab&rtivus.

r. Stems and peduncles closely pubescent . . . var. acrolasius.

q . Basal leaves compound or deeply lobed;

head of fruits much longer than thick

r. Stems bristly-hairy; achenes with a

beak-like style .................. 14. R. pensylvanicus.

r. Stems smooth or with inconspicuous

hairs; achene beakless

s. Face of mature achene with faint

transverse ridges; the 3 lobes of the

stem-leaves usually toothed or shal¬

lowly lobed . . . 8. R. sceleratus .

s. Face of achene without transverse

ridges; the 3 lobes of the stem-

leaves deeply lobed or divided _ _ var. multifidus .

202 Wisconsin Academy of Sciences , Arts and Letters

n. Petals conspicuous, several times as long as the

sepals

t. Basal leaves oval, not lobed or divided .... 7. R. rhomb oideus.

t. Basal leaves deeply cleft or divided

u. Style 1 mm. or more long, erect or ascend¬

ing, beak-like

v. Boots thickened and fleshy; lateral leaf¬

lets rarely stalked; plants without long

prostrate branches . . 11. R . fascicularis.

v. Roots not thickened and fleshy; lateral

leaflets with stalks 5 mm. or more long;

plants soon developing long prostrate

branches . . . 12. R. septentrionalis.

u. Style very short, recurved, not beak-like . 15. R. acris.

1. R. trichophyllus Chaix. Water Crowfoot. R. aquatilis

var. capillaceus of Preliminary Report; see Drew, Rhodora

38: 17. 1936. In lakes and streams throughout the state.

2. R. longirostris Godron. Stiff Water Crowfoot. R. cir-

cinatus of Preliminary Report; see Drew, Rhodora 38: 42.

1936. Southeastern Wisconsin, rarely northwestward to Craw¬

ford, Waushara and Brown Counties; Lake Superior region

in Ashland County.

3. R. Cymbalaria Pursh. Seaside Crowfoot. Very rare

along Lake Michigan shore in Manitowoc, Racine and Kenosha

Counties, inland only at Lake Geneva in Walworth County

where collected in 1885. Also adventive in Central Park in

Superior, Douglas County. Much of the material from Lake

Michigan shore is f. hebecaulis Fernald, Rhodora 16: 162.

1914, and the plants from Superior approach this form. Some

sheets have both glabrous and pubescent plants.

4. R. elabellaris Raf. Yellow Water Crowfoot. R. del-

phinifolius of Preliminary Reports; see Fernald, Rhodora 38:

171. 1936, and Benson, Bull. Torrey Club 69: 315. 1942. In

quiet water throughout the state, perhaps most common south¬

eastward. F. riparius Fernald, l.c., (R. delphinifolius var.

terrestris of Gray’s Manual) is occasionally found on muddy

shores.

5. R. Gmelini DC., var. terrestris (Ledeb.) Benson. R .

Purshii of Gray’s Manual; see Hara, Rhodora 41: 386. 1939,

and Benson, Bull. Torrey Club 69: 313. 1942. Apparently rare

in northern Wisconsin (Map 12, x’s). F. Purshii (Richards.)

Fassett — Flora of Wisconsin . Ranunculaceae 203

Fassett5 has been collected in a cold spring-fed brook near

Jacksonport in Door County and near Ashland in Ashland

County (Map 12, crosses).

6. R. reptans L. Creeping Spearwort. R. Flammula var.

filiformis of Gray’s Manual; see Rhodora 19: 135-137. 1917 ; R.

Flammula var. filiformis (Michx.) Hook.; Benson, Bull. Torrey

Club 69 : 306. 1942. Very common on sandy lake shores of north¬

western Wisconsin and at Green Bay in Brown County (Map

10, dots). The leaves vary from threadlike to flat and about

1.5 mm. wide, in the latter case approaching var. ovalis . Var.

OVALis (Bigel.) T. & G. (R. Flammula var. reptans of Gray’s

Manual; R. Flamuia var. ovalis (Bigel.) Benson, l.c., p. 305),

with blades of leaves from 2-7 mm. wide, has been collected in

Door County (Map 10, crosses).

7. R. rhomroideus Goldie. Dwarf Buttercup. Prairies and

sand, from Pierce, Dunn, Eau Claire, Portage and Shawano

Counties southward; not collected but probably to be ‘expected

in the southwest corner of the state in Vernon, Crawford, Rich¬

land, Iowa and Lafayette Counties.

8. R. sceleratus L. Cursed Crowfoot. On muddy shores

and in shallow water from Dane, Columbia, Brown and Door

Counties southward and eastward. Var. multifidus Nutt, is

approached by plants with achenes not wrinkled and leaves slight¬

ly more divided than usual, from Fox Lake in Dodge County

and Dousman in Waukesha County, and less strongly ap¬

proached by a collection from Fish Creek in Door County,

which has smooth achenes but leaves like the ordinary R. sceler¬

atus .

9. R. abortivus L. Small-flowered Buttercup. In woods,

pastures, etc., throughout the state. The generally more north¬

ern var. ACROLASIUS Fernald, Rhodora 40: 418. 1938, has been

collected in 13 counties from the northern to the southern bor¬

ders of Wisconsin, but its validity as a geographic variety is

shown by the fact that while in northern Wisconsin it may

exist as the major element in many populations its occur¬

rence southward is only as an occasional pubescent individual

among a great majority of glabrous ones; moreover, north

5 R. Gmelini DC., var. terrestris (Ledeb) Benson, f. Purshii (Richards.) n. comb.

R. Purshii Richards.^ Bot. App. Frankl. 1st. Jour. ed. I, 741. 1823. R. Gmelini var. Pur-

shii (Richards.) Hara, Rhodora 41. 386. 1939. Hara recognizes the submersed state

as var. Purshii and the emersed one as var. limosus, while Benson treats the whole range

of ecological forms as one geographic variety to which the earliest name terrestris must

unfortunately be applied.

204 Wisconsin Academy of Sciences, Arts and Letters

of Wisconsin var. acrolasius becomes the only representative

of the species, while southward it disappears. See Fassett,

Amer. Midi. Nat. 27: 512-522. 1942.

A peculiar individual from Pine Hollow near Poynette in

Columbia County appears to be closely related to R. abortivus.

It has hooked styles a little longer than in that species, while

some of the basal leaves have the shape and pubescence char¬

acteristic of R. rhomb oideus. Dr. Lyman Benson, monographer

of North American Ranunculus, has examined the specimen and

commented: “Petals 4.5 mm. long, otherwise a mixture of

characters of R. abortivus and R. micranthus ” It was collected

by the late Dr. A. L. Frolick on May 24, 1935, and all at¬

tempts to find more plants have failed. R. micranthus is not

known from Wisconsin. Pine Hollow is only a few miles east

of the Driftless Area, where many endemics and isolated spe¬

cies are known.

10. R. recurvatus Poir. Hooked Buttercup. Rather com¬

mon in rich woods throughout the state. F. laevicaulis Weath-

erby, Rhodora 31: 164. 1929, was collected in company with

the common pubescent type, at Pine Hollow during one of the

unsuccessful searches for the peculiar plant described in the

preceding paragraph.

11. R. FASCICULARIS Muhl. Early Buttercup. Open ground,

north to Pierce, Dunn, Juneau, Waupaca and Brown Counties.

12. R. septentrionalis Poir. Marsh Buttercup. Through¬

out the state in marshes and wet woods. The stems may be

glabrous, or may have appressed, spreading or reflexed hairs.

13. R. repens L. Creeping Buttercup. Rarely adventive in

the eastern part of the state. Several varieties have been recog¬

nized (Fernald, Rhodora 21: 169. 1919). The typical form and

var. glabratus DC. have been collected at Sturgeon Bay in

Door County; var. villosus Lamotte has been collected in a

park at Madison in Dane County, in a tamarack swamp at

Addison in Washington County, and on a lawn in Wauwa¬

tosa, Milwaukee County; var. glabratus DC. has been collect¬

ed at Milwaukee, Milwaukee County, and Shawano, Shawano

County.

Young individuals of this species sometimes lack the creep¬

ing stems and may be difficult to distinguish from young plants

of R. septentrionalis lacking prostrate stems. There are char-

Fassett — Flora of Wisconsin . Ranunculaceae 205

acters in the young carpels and fruits as follows: R. repens

has the carpels abruptly narrowed to a curved beak 0.5 mm.

long in flower and about 1 mm. long in fruit, and the mature

nutlet is plump, scarcely keeled, 2-2.5 mm. long, while R . sep -

tentrionalis has the carpel gradually tapered to a stout straight

or slightly curved beak which is 1 mm. long in flower and 2

mm. or more long in fruit, and the mature nutlet is flattened

on the sides and broadly keeled, 3-3.5 mm. long.

14. R. PENSYLVANICUS L. f. Bristly Crowfoot. Common

throughout the state in open wet places, bogs, etc.

15. R. acris L. Tall Buttercup. This introduced species

is found across northern Wisconsin, south to Washburn, Tay¬

lor, Marathon, Brown and Kewaunee Counties, and in the

southeastern part of the state from Monroe County southeast¬

ward. The first collection in Wisconsin was in 1858 in the

Apostle Islands in Ashland County, a seat of early settlement.

It was found by Lapham in Milwaukee, and collected at Ra¬

cine in 1879. The first collection in Dane County was in 1886,

but here it is not yet very abundant.

2548. Thalictrum. Meadowrue

a. Stem-leaves with a petiole between the sheath and

the division into 3 parts

b. Plants with a short stout rootstock; leaflets thin,

smooth, the edges not revolute; flowering in May;

achenes bluntly pointed equally at both ends . 1. T. dioicum.

b. Plants with long rootstocks; leaflets thick, ru¬

gose, the margins revolute; flowering in June and

July; achenes rounded to a stipitate base and

long-tapered to the tip

c. Stigmas 1. 0-2.0 (rarely 2.5) mm. long; ma¬

ture achenes with body 3.0-4.0 mm. long and

2.4 mm. or less thick; anthers 2.0-3.5 mm. long

with an apiculate tip about 0.1 mm. long ... 2. T. venulosum.

c. Stigmas 2.Q-4.0 mm. (rarely more) long; ma¬

ture achenes with body 4. 0-6.0 mm. long and

2.5 mm. or more thick; anthers 3.0-4. 0 mm.

long with an apiculate tip about 0.4 mm. long . . 3. T. confine,

a. Stem-leaves sessile, dividing into 3 parts directly

above the sheath

d. Leaflets not glabrous beneath

e. Lower surfaces of leaflets covered with minute

stalked glands . 4. T. revolutum.

e. Lower surfaces of leaflets with jointed hairs . 5. T. dasycarpum.

d. Leaflets perfectly glabrous beneath . 6. ?

206 Wisconsin Academy of Sciences, Arts and Letters

1. T. dioicum L. Early Meadowrue. Rich woods through¬

out the state, perhaps less common northward.

2. T. VENULOSUM Trel. A collection is cited from R. R.

right-of-way north of Hayward, Sawyer County, by Boivin,

Rhodora 46 : 439. 1944.

3. T. CONFINE Fernald. On sandy soil, Wisconsin Point,

Superior, Douglas County.

4. T. revolutum DC. Rather rare in the southeastern cor¬

ner of the state from Washington and Rock counties southeast¬

ward.

5. T. dasycarpum Fisch. & Lall. Purple Meadowrue. Most¬

ly in meadows, throughout the state.

6. From southeastern Wisconsin there are six collections of

a plant resembling the two preceding species but with the firm

rugose revolute-margined leaves absolutely glabrous beneath.

The leaflets of all of our T. revolutum have the same texture, as

do some of those of T. dasycarpum . There are similar plants

collected in Iowa and northern Illinois, while from Missouri

Dr. Steyermark has labelled several collections T. revolutum f.

glabrum Pennell. The anthers range from 1-2 mm. long in

the Wisconsin plant, and Dr. Boivin, Rhodora 46: 471. 1944,

describes the anthers of T. revolutum as being 1.7-2. 8 mm.

long. Boivin recognizes a glabrous variety of T. dasycarpum

but describes it as having membranous leaves and anthers

2. 2-3. 2 mm. long.

BERBERIDACEAE. Barberry Family

a . Plants herbaceous; leaves 5 cm. or more long, com¬

pound or deeply lobed

b. Leaves palmately deeply lobed . 2558. Podophyllum.

b. Leaves compound

c. Leaves cut into 2 leaflets . 2559. Jefferso>nia.

c. Leaves cut into many leaflets . 2565. Caulophyllum.

a. Plants shrubby; leaves less than 5 cm. long, appear¬

ing simple . , . . . 2566. Berberis.

2558. Podophyllum. Mayapple

P. peltatum L. Common in woods in the southern half

of Wisconsin. Map 11. A plant answering the description of

f. aphyllum Plitt, Rhodora 33: 229. 1931, was reported by

Fassett— Flora of Wisconsin . Ranunculaceae

207

Wadmond, FI. Racine & Kenosha Counties, Trans. Wis. Acad.

16: 838. 1909.

2559. Jeffersonia. Twinleaf

J. diphylla (L.) Pers. Rare and local in rich woods of

southern Wisconsin. Map 12, dots.

2565. Caulophyllum. Blue Cohosh

C. thalictroides (L.) Michx. Not uncommon, and charac¬

teristic of rich woods throughout the state. Map 13.

2566. Berberis. Barberry

The 3 species most apt to be found in Wisconsin are keyed

as follows by Beam in the Flora of Indiana:

a. Leaves entire; spines generally simple; flowers in

fascicles of 2-6; petals notched . B. Thunbergii.

a. Leaves not entire, the margins more or less serrate;

spines generally 3-pronged

b. Margins of leaves regularly bristly-serrate; ra¬

cemes many-flowered; petals entire; two year

old branches grayish . . . . . . . B. vulgaris .

b. Margins of leaves irregularly serrate, the teeth

not bristly-pointed; racemes few-flowered; petals

notched ; two year old branches reddish brown . . . B. canadensis.

208 Wisconsin Academy of Sciences, Arts and Letters

B. Thunbergii DC., the Japanese Barberry, is commonly

planted but rarely escapes; B. vulgaris L., the European Bar¬

berry, was formerly planted and widely escaped, and is still

found occasionally in woods and pastures despite the campaign

for its eradication; B. canadensis Mill., the Allegheny Bar¬

berry, is rarely planted.

MENISPERMACEAE. Moonseed Family

2567. Menispermum. Moonseed

M. canadense L. Rather local, mostly in wooded alluvial

soils northward to Pierce, Dunn, Eau Claire, Clark, Taylor,

Marathon, Shawano and Manitowoc Counties. Map 14.

LAURACEAE. Laurel Family

2795. Sassafras. Sassafras

The writer has never seen a specimen of Sassafras from

Wisconsin. Dr. R. P. Hoy, History of Walworth County, Wis¬

consin, p. 133. 1882, wrote: “Sassafras officinale is a small

tree of fine appearance with fragrant leaves and bark. Grows

in Kenosha county. Should be cultivated.” Mr. S. C. Wadmond

did not list it in his Flora of Racine and Kenosha Counties,

but he recently wrote me as follows : “My memory is that Dr.

Hoy used to find Sassafras in an early day along Pike Creek

(Kenosha County) near its outlet into Lake Michigan, but I

cannot recall definitely whether Dr. Davis [Dr. J. J. Davis,

well known in the field of mycology] had ever encountered it.

I am sure that if a station for it had existed when I was com¬

piling my Flora, Dr. Davis would have directed me to it, as

was his general custom.”

Since Deam’s Flora of Indiana records both S. aibidum

(Nutt.) Nees and S. aibidum var. molle (Raf.) Fernald (see

Rhodora 15: 14-18. 1913, and 38: 178-179. 1936) from the

counties in Indiana bordering Lake Michigan, it is not un¬

reasonable to expect Sassafras in Kenosha County, Wisconsin.

Less expected, however, are reports from two adjacent counties,

Marquette and Green Lake, in central Wisconsin. On October

25, 1857, William P. Huntington, U. S. Deputy Surveyor, mak¬

ing the original land survey, laid out the line between Sections

22 and 23 in Township 16 North, Range 9 East, in what is now

Marquette County, and described the ground cover, in part, as

Fassett — Flora of Wisconsin. Ranunculaceae 209

follows: “Undergrowth red & black willow, alder, green-briar,

prickly ash, barberry, gooseberry, blackberry, Poison Sumach,

sassafras.” Of course, he might have been mistaken about

the sassafras. The writer has done considerable collecting in

the region, and has never encountered it.

Just 15 miles southeast of the point where Surveyor Hunt¬

ington reported Sassafras is the town of Marquette, in Green

Lake County on the south shore of Lake Puckaway. In early

1929 Mr. F. M. Uhler wrote me about some plants noted by

Mr. W. L. McAtee at Marquette, as follows : “All three of these

plants were recorded by Mr. McAtee at Marquette, Wisconsin,

during the period between October 21 and November 6, 1908.

He collected a specimen of Tanacetum huronense at that point

on November 4, 1908, and a specimen of Opuntia Rafinesquii 'on

granite among red cedars’ on November 5, 1908. Sassafras

variifolium was merely recorded in his list for that period

and no specimen preserved.” Tanacetum huronense, it may be

remarked, is known in Wisconsin only as a rare plant of the

Lake Michigan shore in Door County. In September, 1929, Mr.

Uhler and I spent a day collecting in the region of Marquette;

we found Opuntia Rafinesquii on granite among red cedars,

but no Tanacetum huronense, and no Sassafras.

.

A CYTOLOGICAL STUDY OF

THE DEVELOPMENT OF THE OOSPORE OF

SCLEROSPORA MACROSPORA (SACC.)

E. S. McDonough*

Biology Department , Marquette University

Milwaukee, Wisconsin

The author became acquainted with the fungus known as

Sclerospora macrospora (Sacc.) in the summer of 1941 while

at the Biological Laboratories of Harvard University. Dr.

William H. Weston kindly made available his own exsiccate and

gave detailed descriptions of the territory in Tennessee and

Kentucky where he had previously collected (17), which made

possible the securing of fresh material for study. This was

especially fortunate because, so far as the author is aware, in¬

vestigations of the cytology of members of the genus

Sclerospora have been limited to the single species, S. gramini-

cola (Sacc.) Schroet (2,8,7,8,13).

It is the purpose of the writer to set forth in this paper

the main features of the development of the sex organs and

oospore of Sclerospora macrospora (Sacc.), [Phytophthora

macrospora (Sacc.) S. Ito and I. Tanaka] and to show in a

preliminary way how these cytological characteristics may in¬

fluence the classification of the fungus. The writer hopes that

later Weston, who has been intermittently working on this

oomycete for a long time, will publish further on its identity

and taxonomic affiliations as well as on its geographic distribu¬

tion and range of parasitism.

* This investigation was carried out with the aid of a grant from the A.A.A.S. re

ceived through the Wisconsin Academy of Science, Arts and Letters,

It is a pleasure to acknowledge the inspiration and counsel supplied by Dr. William

H. Weston in the inception and during the course of the investigation.

Many thanks go to Miss Frances Bielinski who helped make the slides.

211

212 Wisconsin Academy of Sciences , Arts and Letters

McDonough — Sclerospora Macrospora (Sacc.) 218

Description of Figures 1-8

All drawings were made of sectioned material with the aid of a

camera lucida. Magnifications are given for each drawing as reproduced.

1. A young antheridium attached to a young oogonium. The early

peripheral distribution of the oogonial nuclei is indicated. X653.

2. An oogonium showing the thick wall. X 653.

3. A female gametangium projecting into the antheridium. X653.

4. The first mitosis of nuclei in the oogonium. X1460.

5. The two sex organs showing the fertilization tube with several

nuclei. “Zonation” and migration or dissolution of all oogonial

nuclei but one has taken place. X653.

6. An oospore in which karyogamy is about to take place. X653.

7. An oospore showing nuclei in different phases of mitosis. X1460.

8. A resting spore showing the thick-walled oospore surrounded by

the oogonial wall. The eoenocytic condition is apparent. X 653.

214 Wisconsin Academy of Sciences , Arts and Letters

Materials and Methods

Collections of infected wheat containing oogonia were made

June 13, 1942 near Fulton, Kentucky. This location was in the

general vicinity where Weston reported the presence of Sclero -

spora macrospora on wheat in 1921 (17). The diseased leaf

tissue was fixed in F A A, Craf I (10), Flemming medium, and

Carnoy’s solution without chloroform (75 ml. anhydrous ethyl

alcohol and 25 ml. glacial acetic acid). The tissue fixed in Craf

I and Flemming was most useful. The material was run up into

paraffin and sectioned at 10 microns. It was stained with Flem¬

ming's triple stain, iron-alum hematoxylin, and crystal violet-

iodine.

Observations

The sexual organs were observed to have arisen intercellu-

larly and usually intimately associated with the vascular bun¬

dles of the wheat plant. It was possible to distinguish the

antheridial and oogonial primordia only after the oogonial ele¬

ment had attained a relatively large size (Fig. 1) and the hy-

phae were in contact. The cytoplasm, in both structures, was

at first uniformly and minutely vacuolate. Counts made in 17

of the male elements showed from six to nine nuclei to be

present in the antheridium after segmentation of the antheridial

hypha. A simultaneous division of the antheridial nuclei was

observed (Fig. 1) previous to a mitosis of the nuclei in the

female organ. Mitoses of nuclei in male organs attached to

oogonia which had differentiated further were found. This

seemed to point to the conclusion that there were two succes¬

sive, simultaneous divisions of nuclei in the antheridium. Fur¬

ther evidence in support of the theory that two divisions took

place was found when counts were made of the nuclei in fully

developed male gametangia, and compared with the counts

from the young organs, a range of from 16 to 28 nuclei being

observed in nine mature antheridia. However, a great vari¬

ation existed in antheridial size and the counts of nuclei ob¬

served did not absolutely prove that two successive mitoses

always took place.

The wall of the young oogone thickened soon after the

maximum size of the gametangium had been attained (Fig. 2).

The average thickness of the oogonial wall was found to be

two microns. A range of from 30 to 45 nuclei which tended to

McDonough— Selerospom Macrospora (Sacc.) 215

be peripheral in location was observed to be present in the

oogonia. These nuclei soon increased in size (Figs. 1,2,3) and

a simultaneous nuclear division took place ( Fig. 4) previous

to a definite “zonation” of the cytoplasm. After these mitoses,

differentiation of the cytoplasm into ooplasm and periplasm

and at the same time a degeneration of the nuclei in the peri¬

plasm were found to occur (Fig. 5). No second simultaneous

mitosis of nuclei was observed, but it was not uncommon to

find some of the nuclei in the oogonium, subsequent to the early

divisions, in the metabolic condition and at the same time dif¬

ferent phases of mitosis. Because of this degeneration of nu¬

clei and the great variation in size of the oogonia, comparison

of nuclear counts in young and old oogonia were not very re¬

liable in determining the number of divisions. However, a

range of from 52 to 120 nuclei was obtained when an attempt

was made to get maximum numbers. Almost from the start

few nuclei were found in the central region of the oogone

(Bigs. 1,2,3), and before fertilization all but one of the nuclei

originally present, or those produced later by division, migrated

to the periplasm or degenerated until eventually a single fe¬

male pronucleus existed in the center of the oosphere (Fig. 5).

A characteristic coenocentrum was not found. Some tend¬

ency for a small amount of dense cytoplasm to be present in

the center of the oosphere was observed (Figs. 5,6), but this

was never abundant and may have been caused by the failure

of the central part of the cytoplasm to have its vacuoles in¬

crease in size, a characteristic typical for most of the cytoplasm

(Figs. 1 to 5).

The wall of the oogone was observed to have remained thin

at the region of contact with the antheridium and the two

walls were so closely associated that a separation was not ap¬

parent. This intimate association of the sex organs was further

accentuated by a protrusion of the oogonial material into the

antheridium, the “receptive papilla” of Wager (16) and Stevens

(11) (Fig. 5). Typical also of this association was the eccentric

position of the ' oosphere wherein the egg was placed near the

side of the oogonium to which the antheridium was attached.

The fertilization tube was very conspicuous (Fig. 5) and was

observed to contain six or more nuclei, all evidence indicating

that at least most of these entered the oosphere. However, it

was clearly indicated that but one nuclear union took place,

216 Wisconsin Academy of Sciences , Arts and Letters

a small male pronucleus uniting with a slightly larger female

pronucleus, the remaining nuclei from the antheridium degen¬

erating (Fig. 6).

Due to the scarcity of oospores showing divisions of the

union nucleus and its daughter nuclei, it was assumed that

once the union was completed the nuclear divisions followed

rapidly. However, it was evident that nuclear divisions follow

soon after fertilization (Fig. 7) and that the resting spores

contained numerous nuclei (Fig. 8). From 38 to 44 nuclei

were found in each oospore located in the peripheral cytoplasm,

the center being occupied by a more or less homogeneous cen¬

tral body (Figs. 7,8) of reserve material.

During the maturation of the oospore the wall reached an

average thickness of three microns. There were clearly visible

at least two layers, an outer thin one and a comparatively

thick inner wall. A large percent of the oospores observed

in sectioned material had the thin outer coat minutely wrin¬

kled. This condition did not seem to be an artifact, since the

inner wall was definitely, in many instances, minutely rough¬

ened. However, not all oospores were observed to be so

roughened and this condition was not clearly distinguished in

unsectioned material. In a different manner the oogone wall

became roughened. Due apparently to a folding of this wall,

protrusions were formed (Fig. 8). Such foldings were caused

apparently by a shrinking of the v/all of the oogone and con¬

formation of the wall to the surrounding cells.

Discussion

The phycomycete considered here was first described by

Saccardo (9), who included it in the genus Sclerospora largely

on the basis of the thickened wall of the oogone, the imperfect

stage being unknown. The description of the voluminous, pap¬

illate, lemon-shaped sporangia as being born singly and ex¬

ternally on short peduncles by Peglion (5) and Peyronel (6)

brought into question the taxonomic position of Sclerospora

macrospora (Sacc.), since this condition was so unlike the

members of the genus. Tanaka (15) in 1940 described this

fungus as Phytophthora macrospora (Sacc.) S. Ito and I. Ta¬

naka and considered Sclerospora macrospora as a synonym.

McDonough — Sclerospora Macrospora (Sacc.) 217

A comparison of the cytological features of the fungus

herein referred to as Sclerospora macrospora with the published

reports of both amphigynous and paragynous Phytophthoras

casts doubt on the advisability of including the parasite in the

genus Phytophthora. The thick oogonial wall alone would

seem to exclude the fungus from the above genus, since the

published reports of Murphy in 1918 (4) for the amphigynous

Phytophthora erythroseptica and of Blackwell for the essen¬

tially paragynous P. cactorum (1) clearly indicate the thin

nature of this wall. To this discrepancy may be added other

cytological differences. It has long been considered that the

number of nuclei present in the mature oospore is a good cri¬

terion of the taxonomic position of a member of the Perono-

sporales, since all members of the Albuginaceae studied cyto-

Jogically have coenocytic resting spores while Sclerospora (as

exemplified by S. graminicola) , Plasmopara , Peronospora, and

Phytophthora have a delayed union of. the pronuclei, the ma¬

ture oospore being monocaryon.

The development of the oospore of Sclerospora macrospora

seems to be more like that of Albugo tragopogonis as described

by Stevens (12) than any other member of the Peronosporales

so far studied. In A. tragopogonis several nuclei enter the

oosphere from the antheridium and but one unites with the

single functional female nucleus. A. ipomoeae-panduranae, a

fungus similar to A. tragopogonis in many respects (14), has

been reported by Stevens as having a slight but very perceptible

thickening of the oogonial wall. However, there would seem

to be little justification for placing Sclerospora macrospora in

the genus Albugo on the basis of the asexual phase.

Summary

From six to nine nuclei were observed in the young anthe¬

ridium and from 30 to 45 nuclei in the oogonial initial. Simul¬

taneous mitoses were found in both sex organs, and at least

some of the nuclei so produced underwent a second division.

Several nuclei were found to enter the oosphere, one single male

nucleus uniting with the solitary female pronucleus, the rest

disintegrating. The mature oospore contained from 33 to 44

nuclei. The oogonial wall averaged two microns in thickness,

and the wall of the oospore consisted of a thin outer layer and

218 Wisconsin Academy of Sciences , Arts and Letters

thick inner layer, the combined thickness of the two averaging

three microns.

A preliminary consideration of the taxonomic position of

the fungus led to the conclusion that this species could not

easily be placed in any known genus. Further work now con¬

templated may well lead to the erection of a new genus.

Literature Cited

(1) Blackwell, Elizabeth. 1943. The life history of Phytophthora

cactorum (Leb. V Cohn) Schroet. Trans. Brit. Myc. Soe. 26:71-

89.

(2) McDonough, E. S. 1937. The nuclear history of Sclerospora grami-

nicola. Mycologia, 29:151-173.

(3) - - 1943. Studies on the cytoplasm and its inclusions in Sclero¬

spora graminicola. Amer. J. Bot. 30:809-813.

(4) Murphy, P. A. 1918. The morphology and cytology of the sexual

organs of Phytophthora erythroseptica Pethyb. Ann. Bot. 32:115-

152.

(5) Peglion, V. 1930. La formazione dei conidi e la germinazione delle

oospore della Sclerospora macrospora Sacc. Bolletino della R.

Stazione di Patologia vegetale di Roma, Anno 10, Nuova Serie:

153-164.

(6) Peyronel, B. 1929. Gli zoosporangi nella Sclerospora macrospora .

Bollettino della R. Stazione di Patologia vegetale di Roma. Anno

9, Nuova Serie: 353-357.

(7) Ruhland, W. 1902. Die Befruchtung der Albugo Lepigoni und ein-

iger Peronosporeen. Hedwigia, 41: 179-180.

(8) - 1904. Studien uber die Befruchtung der Albugo Lepigoni

und einiger Peronosporeen. Jahrb. Wiss. Bot. 39: 135-166.

(9) Saccardo, P. A. 1890. Fungi aliquot australienses. Hedw. 29: 154-

156.

(10) Sass, J. E. 1940. Elements of Botanical Microtechnique. McGraw-

Hill.

(11) Stevens, F. L. 1899. The compound oosphere of Albugo Bliti. Bot.

Gaz. 28: 149-176, 224-245.

(12) - 1901. Gametogenesis and fertilization in Albugo. Bot. Gaz.

32: 77-97, 157-169, 238-261.

(13) — - 1902. Studies in the fertilization of Phycomycetes. Bot.

Gaz. 34: 420-425.

(14) - — 1904. Oogenesis and fertilization in Albugo ipomoeae — pan-

duranae. Bot. Gaz. 38s: 300-302.

(15) Tanaka, I. 1940. Phytophthora macrospora (Sacc.) S. Ito and I.

Tanaka on wheat plant. Ann. Phytopath. Soc., Japan, 10: 127-

138.

(16) Wager, H. 1896. On the structure and reproduction of Cystopus

candidus. Ann. Bot., 10: 295-342.

(17) Weston, Wm. H. 1921. The occurrence of wheat downy mildew

in the United States. U.S.D.A. Dep. Circ. 186.

NOTES ON WISCONSIN PARASITIC FUNGI. VIII.

H. C. Greene

University of Wisconsin, Madison

In the University of Wisconsin Herbarium there is a specimen

of Albugo Candida (Pers.) 0. Ktze. on Thely podium wrightii

from Zion Park, Utah, collected June 25, 1926 by E. J. Kraus.

Although this is not Wisconsin material it is mentioned here for

the sake of a record, since so far as I can determine T. wrightii

has not been listed elsewhere as a host for this fungus.

Davis in his “Parasitic Fungi of Wisconsin” states that

Uncinula necator (Schw.) Burr, is common on Vitis vulpina

in Wisconsin. Over a ten-year period I have seen many thousand

plants of V. vulpina, but have observed U. necator in a single

case only. This may have been more abundant in past years,

but it is perhaps significant that there is but a single earlier

specimen on this host, collected 40 years ago, in the Wisconsin

Herbarium, and one in the Davis Herbarium, collected 56 years

ago.

Plants of the showy garden subject, Cleome spinosa, in a Uni¬

versity of Wisconsin greenhouse were lethally blighted by a

powdery mildew, but perithecia failed to develop.

The oidial stage of presumed Erysiphe cichoracearum DC.

has been found in abundance on Hieracium longipilum at Madi¬

son. Hieracium canadense is regularly thus infected, but mature

perithecia seem not to have been collected on Hieracium in Wis¬

consin.

Thompson (Phytopath. 31 : 241, 1941) finds a Mycosphaer-

ella to be the perfect stage connected with Septoria musiva Peck,

cause of the common leaf-spot disease of poplar. This he names

Mycosphaerella populorum. (Davis followed Petrak in con¬

sidering S. musiva Peck to be a synonym of S. populi Desm., and

the numerous specimens in the Wisconsin Herbarium are thus

filed.)

219

220 Wisconsin Academy of Sciences , Arts and Letters

Demaree and Wilcox (Phytopath. S3: 986, 1948) present

evidence to show that the organism causing the common leaf -spot

of raspberry, usually supposed to be Septoria rubi West., is

identical with Cylindrosporium rubi Ell. & Morg., and that the

latter name must be supplanted by that of the ascigerous stage,

Sphaerulina rubi Demaree & Wilcox. They question the connec¬

tion of Mycosphaerella rubi Roark (Roark worked in Wiscon¬

sin) with the leaf-spot of Rubus. In their artificial inoculations

the raspberry leaf -spot fungus could not be carried over to dew¬

berry and blackberry. They suggest the temporary retention of

the binomial Septoria rubi for designating the fungus causing

the leaf-spot in the Eubatus section of Rubus.

The ascigerous stage of Acanthostigma occidentale (Ell.

& Ev.) Sacc. occurs associated with the conidial stage in a collec¬

tion on Artemisia ludoviciana,' made at Madison, June 21, 1944.

Previous Wisconsin specimens on this host have had conidia only.

Mature Pleospora sp. has been found closely associated and

presumably connected with Alternaria on small, definitely de¬

limited arid spots on living leaves of squash. The asci are about

65 X 30/*, the ascospores about 25 X 13-15/*. It is questionable,

however, that the fungus is parasitic since the spots are of a

type that might be caused by insects.

Coleosporium solidaginis (Schw.) Thum. I occurred in great

profusion on needles of young trees of Finns resinosa planted

in a nursery in the University of Wisconsin Arboretum at Madi¬

son, May 26, 1944. A single small earlier collection on this host

was made by Davis from planted specimens in Peninsula State

Park, Door Co.

In Davis' “Parasitic Fungi of Wisconsin," p. 43, it is indicated

that telia only of Uromyces alopecuri Seym, have been found in

Wisconsin, but uredia have been collected also as shown by Davis'

own specimens on Alopecurus geniculatus var. aristulatus from

Haugen, Barron Co., and Big Bend, Waukesha Co.

In the summer and fall of 1943 Puccinia virgata Ell. & Ev.,

previously unreported from Wisconsin, occurred in vast abund¬

ance on Sorghastrum nutans in the University of Wisconsin Ar¬

boretum at Madison. In 1944, on the same plants, not a trace of

the rust could be found.

A Phyllosticta on leaves of Syringa vulgaris from Madison

does not meet the specifications of hitherto described Phyllostic-

tae on this host. The conidia are like those of Phyllosticta

Greene — Notes on Wisconsin Parasitic Fungi . VIII 221

syringae West., but the leaf spotting is quite different. In the

Madison material very large marginal areas are involved, up to

one fourth to half the leaf surface. The type of necrosis is that

frequently associated with Ascochyta, but microscopic examina¬

tion of many leaves shows no evidence whatever of septation in

any of the conidia. It. is possible that the fungus is secondary.

A Phyllosticta which may be of the type of Phyllosticta

decidua Ell & Kell, has been found on leaves of Siiphium integri-

folium collected near Eagle, Waukesha- Co. The leaves are heavily

rusted by Puccimia silphii and the epiphyllous pycnidia are re¬

stricted to arid, whitish regions above the telial cushions. The

hyaline, short-cylindrical conidia are 3-3.5 X 6-8/*.

Another in the series of indeterminate and dubiously para¬

sitic small-spored Phyllosticta© on astericolous and solidagini-

colous hosts has been observed on large gray spots on leaves of

Aster lucidulus. The pycnidia are large and very dark, with nu¬

merous hyaline, biguttulate conidia about 7X2/*. The spots ap¬

pear to have been caused, at least partially, by the infestation of

insects.

A problem of great importance to those interested in the iden¬

tification of fungi parasitic on plants has recently been brought

rather forcibly to my attention although, as all must who work

with these parasites, I have always been aware of its existence.

The problem is the old one of how much emphasis should be

placed on hosts as bases for establishment of species of plant

parasitic fungi.

In the fall of 1944 I collected a species of Ascochyta on the

leaves of what I identified as a large rayless species of Senecio.

This is firmly established as an escape on the grounds of a long-

abandoned farmhouse near Madison. Examination of the litera¬

ture showed that Petrak (Ann. Mycol. 22: 167, 1924) described

Ascochyta senecionicola on Senecio sarracenicus from central

Europe, a species which the Madison specimen matches almost

exactly in microscopic characters, both from the printed descrip¬

tion and from Petrak’s own specimens in the University Herba¬

rium. Although in the Wisconsin material the spots are some¬

what smaller and better defined, it was nevertheless placed in the

herbarium as Ascochyta senecionicola Petr. Later in the year,

following a more careful check of the host, I discovered that my

supposed rayless Senecio is the common mint geranium, Chrysan¬

themum balsamita L. var. tanacetoides Boiss. In the Gray’s

222 Wisconsin Academy of Sciences, Arts and Letters

Manual treatment Chrysanthemum and Senecio are in different

tribes, the Anthemideae and Senecioneae respectively, although

these tribes are placed next one another.

The question, then, is: should Ascochyta senecionicola,

described on Senecio, be reported as occurring on the not too

closely related Chrysanthemum, or should my specimen be de¬

scribed as new, since no similar Ascochyta appears to have been

described on Chrysanthemum? The problem thus posed is per¬

haps insoluble without cross inoculations between hosts, a thing

which is unfortunately often not feasible unless the hosts are of

considerable economic importance. In the present case it is prob¬

ably best to see whether the fungus can be collected again at the

same station and, if so, attempt to obtain cultures with which

to try infection of Senecio. In my experience, so sporadic is the

occurrence of most parasites that the chances are at least even

that this will not be found again.

An interesting Ascochyta on Solanum dulcamara was col¬

lected in small quantity at Madison, July, 1943. The fungus

occurs on numerous transparent spots on the leaf, and parasitism

is perhaps doubtful. When gathered, the impression was that it

was probably Phyllosticta decidua Ell. & Kell., although it

was noted that the pycnidia did not show the translucency so

often characteristic of Ph. decidua. However, microscopic ex¬

amination revealed an Ascochyta with small, black, indistinctly

ostiolate pycnidia, 60-90 p diam. The hyaline, uniseptate conidia

are 6-10x3. 5-4.5/x. This fungus does not seem to correspond to

Ascochyta dulcamarae Bubak, Phyllosticta dulcamarae

Sacc., or Phyllosticta perforans Ell. & Ev.

Certain of the Wisconsin specimens of Septoria on Alnus that

have been variously placed under Septoria alni Sacc. and Sep¬

toria alnifolia Ell. & Ev. have the pycnidial walls so imper¬

fectly developed it would seem they might better be referred to

Cylindrosporium. In particular is this true of a specimen col¬

lected at Madison in 1943 on Alnus incana. On the basis of mi¬

croscopic morphological characters alone this is a good Cylindro¬

sporium, but the type of host lesion, and one's knowledge of

bridging forms, suggest its inclusion with the alnicolous Sep-

torias.

As in the case of similar material placed in the herbarium by

Davis, a specimen of Septoria gei Rob. & Desm. on radicle leaves

of Geum canadense from Madison has imperfect pycnidia verging

Greene — Notes on Wisconsin Parasitic Fungi . VIII 223

on acervuli, and the conidia are up to 50/a long. This species seems

very close to Cylindrosporium.

Septoria dearnessii Ell. & Ev. has been found on Angelica

atropurpurea in 1943 in the University Arboretum at Madison.

The only other Wisconsin station for this species is Eagleville,

Waukesha Co.

Phleosfora anemones Ell. & Kell, on Anemone virginiana

from Madison was at first determined as Septoria cylindrica

Ell. & Ev. The latter species was described in 1900 on Anemone

cylindrica from Montana, and so similar is the description to that

of Phleosfora anemones that it appeared that these might be

conspecific. No specimen of Septoria cylindrica is in the Wis¬

consin Herbarium, but material of the Phleospora was sent to

Dr. Linder of the Farlow Herbarium who finds that the two are

in no way different and that both correspond to the type of

P. anemones. The latter name must stand unless this is con¬

sidered to be a Septoria, in which case S. cylindrica Ell. & Ev.

would be the designation. (J. J. Davis, in his “Parasitic Fungi

of Wisconsin,” p. 79, gives this the new name of Septoria pxjni-

CEA which is untenable, since S. cylindrica is the earlier name.

Dr. Linder points out that 0. Kuntze in 1898 transferred Phleq-

SPORA ANEMONES to SEPTORIA ANEMONES (E. &. K.) 0. Ktze.

This appears to be a later homonym, since Desmazieres applied

the same name to another and well-known fungus forty odd years

previously.)

An excipulaceous fungus, which perhaps should be assigned

to Patellina Speg., has been found at Madison on Penthorum

sedoides. The amphigenous, flesh-colored, cup-like fruiting struc¬

tures are sessile and firmly seated in the host tissue, but project

noticeably above the surface. The brown spots are very definite,

almost circular, about 5 mm. diam. The cups are 125/a or more

in diam., and the long, slender phores, some of which are

branched, are about 30-40 Xl-Sy. They are crowded in the cup

and tend to curve toward the center. The acrogenous, hyaline

conidia are ellipsoid to allantoid, about 5-8 X2y, and are produced

in great profusion.

An unusual dematiaceous fungus occurs on an undetermined

narrow-leaved species of Carex, collected at Eagleville, Wauke¬

sha Co., August 8, 1943. So far as I have been able to determine

this does not correspond in its characters to any previously de¬

scribed genus. The specimen is ample and in excellent condition,

224 Wisconsin Academy of Sciences, Arts and Letters

but it seems inadvisable to erect a new genus of parasitic fungi

where the host concerned is unknown as to species. However, the

following descriptive notes are given as a possible aid to others

who may find the same thing and wish data and material for

comparison. The specimen will be preserved and available for

examination. The fungus is hypophyllous on narrow, more or

less elongate, medium brown spots on the living leaves. The gre¬

garious, seriate tufts of conidiophores are produced through the

stomata in fascicles which are compact at the base, but in which

the individual conidiophores spread widely upwards. The phores

are pale brown below, rather deep brown in mass, fading into

hyaline toward the tips, usually obscurely once or twice septate,

more or less geniculate and tortuous, sometimes once branched

near the base, or sometimes with pseudo-branching which is the

result of anastomosis between adjacent conidiophores. There are

from about 10-20 phores (approx.) in a tuft. The more strongly

geniculate phores may have a number of spore scars, and the tips

are frequently denticulate with several scars in close proximity.

The phores are from 30-80 X 3-3. 5 /x, mostly about 45x3/x. The

conidia, by direct lighting of the dried material as collected, with

appropriate magnification, are seen to be several-catenulate, and

when the conidiophores are denticulate several chains of conidia

may be produced simultaneuosly from near the tip of a single

conidiophore. The conidia are continuous, hyaline, globose or

broadly ellipsoid or ovoid, from 5-13x4-6/x. Those which are

globose are of course not more than 6/x diam.

A very interesting species of Cladosporium has been found

on living leaves of Acer negundo at Madison, August 1944. This

species is characterized by the presence, free among the conidio¬

phores, of large numbers of germinated conidia which at first,

because of their uniformity, were taken to be spores of a Cerco-

spora type. (A specimen from the same tree taken a month

later corresponds exactly to that collected in August). These

germinated spores form structures which measure from about

50-115/x long and are 2- or 3-septate. They are strongly con¬

stricted at the septa and the primary, original cell which is itself

only rarely septate (as is so frequently the case in Cladosporium)

runs from about 15-21 X 9-11 /x. Examination of these cells as

they are borne on the conidiophores before dispersal shows them

to be of the same length, but considerably narrower, as one would

expect of ungerminated spores. There is often the appearance

Greene— Notes on Wisconsin Parasitic Fungi. VIII 225

of catena I at ion, although this may be due, in some eases at least,

to germination in situ on the conidiophore. The very dark coni-

diophores are composed of short-eylindrie, almost isodiametric

cells and are markedly several times geniculate and crooked.

They are frequently two to three times branched, at wide angles,

and are of stiff and awkward aspect. I do not find a previous

report of Cladosporium on box elder, and this may be a new

species. I shall be pleased to furnish material to any mycologist

who will undertake the specific determination of this fungus.

In my opinion Cladosporium aromaticum Ell. & Ev. and

Cladosporium mervalr Ell. & Bearn, on Rhus are the same

thing, and since the former name is the earlier it should be used.

One principal supposed difference between the two forms is that

in C. aromaticum the fungus is supposed to be epiphyilous, while

in C. nervals it is, by implication at least, hypophyilous. Recent

collections made near Madison on Rhus glabra show the fungus

to be amphigenous, somewhat diffuse on the upper surface and

with a fairly prominent development along the midrib on the

lower surface. Differences in conidiophore length do not seem

to be of diagnostic importance. Fungi Columbiani 2010, sup¬

posedly type material of C. NERVALE, has no conidiophores over

SO/A long, but in the description on the label they are given as

150 X 6/a. Also, the supposed differences in the shape of the

conidia appear not to be significant.

Cercospora DULCAMARAS (Peck) Ell. & Ev. on Solanum dul¬

camara (Jour. Myc. 1 : 55, 1885) as described has the fruit¬

ing forming indefinite, subviolaceous, or greenish-lead-colored

patches on the lower surface of the leaf, and also more sparingly

so above but without any distinct spots. In an earlier paper

(Trans. Wis. Acad. Sci. 32: 79, 1940) I mentioned a collection

made at Mazomanie, Wis. in which 'The conidiophores are borne

on definitely delimited, arid white spots with a wide, blackish-

brown border.” Two later collections made in successive years

at Madison show similar sharply defined spots, but a specimen

taken at Madison in 1944 has the effuse type of fruiting, as do

two earlier collections made by Davis. On the basis of macro¬

scopic appearance one would immediately conclude that two dif¬

ferent fungi are involved, but microscopically the differences

are not so apparent. The conidiophores are perhaps somewhat

longer and more closely fascicled in the form producing the defi¬

nite spots, but the conidia are much the same, and Professor

226 Wisconsin Academy of Sciences, Arts and Letters

Chupp stated that the Mazomanie specimen! was probably best

referred to C. dulcamarae. If the two forms should be con¬

stantly found over a long period of years, however, it would

seem that at least a varietal distinction might be justified.

In 1944 at Madison, on Senecio balsamitae , there was found

a Cercospora which is borne on very definite small spots, and in

which the fruiting, though amphigenous, is chiefly epiphyllous.

A specimen was sent to Professor Chupp for determination and

it is his opinion that this should be referred to Cercospora sene-

CIONICOLA J. J. Davis (Trans. Wis. Acad. Sci. 30: 10, 1987) on

the basis of similar microscopic characters, and indeed they do

correspond closely. In C. senecionicola, as described on Senecio

aureus , the spots are indeterminate and the fungus hypophyl-

lous and effused. It is interesting that Davis found this in the

effused condition on Senecio balsamitae , as well as on S. aureus.

In “Notes on Wisconsin Parasitic Fungi. Ill” (Trans. Wis.

Acad. Sci. 35: 120, 1943) mention is made of an Alternaria pos¬

sibly parasitic on leaves of Petalostemum candidum. A similar

fungus has recently been observed, this time on definite lesions

on green stems of the same host.

In the lists of additional hosts and species not hitherto re¬

ported for Wisconsin which follow, all collections, unless other¬

wise specified, were made at Madison, Dane Co.

Additional Hosts

The fungi listed have not before been reported as occurring

on these hosts in Wisconsin.

Peronospora parasitica (Pers.) Fr. on Dr aba nemorosa.

Marinette Co., Marinette, June 23, 1916. On a phanerogamic

specimen collected by the late Charles Goessl. This is presum¬

ably the form of P. parasitica which Gaumann sets apart as

Peronospora drabae.

Sphaerotheca humuli var. fuliginea (Schl.) Salm. on

Erigeron canadensis. October 17, 1944.

Microsphaera alni (Wallr.) Wint. on Lathyrus palustris

var. myrtifolius . October 22, 1910. Coll. J. R. Heddle. I do not

find any other specimens in the herbarium on this variety of

Lathyrus, which some authors set aside as a distinct species.

Greene — Notes on Wisconsin Parasitic Fungi . VIII 227

Microsphaera diffusa C. & P. on Desmodium acuminatum.

( D . grandiflorum of Gray’s Manual, 7th ed.) October 22, 1910.

Coll. J. R. Heddle. Not only new to Wisconsin, but seemingly

not reported elsewhere as a host for this fungus.

Erysiphe polygoni DC. on Pisum sativum . Grant Co., Platte-

ville, September 1, 1888. Coll, and det. by S. M. Tracy. This

specimen was distributed as one of the Reliquiae Seymourianae.

E. polygoni is not mentioned in Davis’ notes as occurring on

cultivated pea in Wisconsin.

If the development of spores on the living host is one of the

criteria distinguishing Erysiphe cichoracearum DC. from E.

galeopsidis DC., then a recent (1944) collection on Scutellaria

lateriflora must be assigned to E. cichoracearum. Davis at one

time listed his specimens on this host as E. cichoracearum, but

later filed them all as E. galeopsidis. Examination of these spe¬

cimens shows that they too have asci with spores, and they are

plainly not overwintered material. For this reason, therefore, the

record of E. galeopsidis on Scutellaria lateriflora in Wisconsin

seems to be incorrect.

Puccinia coronata Cda. II, III on Agropyron repens. Au¬

gust 10, 1944. Arthur’s Manual lists only Vermont as a host

locality for P. coronata on quack grass.

Puccinia extensicola Plowr. II, III on Carex stellulata var.

cephalantha. July 20, 1948.

Puccinia caricis (Schum.) Schroet. II, III on Carex stricta.

September 15, 1944.

Phyllosticta nebulosa Sacc. on Arenaria lateriflora. Sauk

Co., Town of Sumpter, Baxter’s Hollow, July 8, 1944. The spores

of the specimen on Arenaria are somewhat wider than the 2 fi of

the description, averaging about 2.5 /x, as do those of other Wis¬

consin specimens on Silene. As in most of the specimens on

Silene, the pycnidia are on dead portions of the leaves.

Phyllosticta decidua Ell. & Kell, on Geum strictum. July

30. On Monarda fistulosa, July 19. On Solanum dulcamara. July

22. All collected in 1943. Although this is doubtfully parasitic,

it is included because of previous mention on other hosts in

Wisconsin lists.

Phyllosticta phaseolina Sacc. on Soja max. Rock Co.

Janesville, July 14, 1944. Coll. E. E. Honey & J. G. Dickson.

228 Wisconsin Academy of Sciences , Arts and Letters

Ascochyta graminicola Sacc. on Muhlenbergia foliosa. Oc¬

tober 8, 1944. Seymour does not list M. foliosa as a host of this

fungus.

Septoria atropurpurea Peck on Solidago altissima . Septem¬

ber 28, 1944. In this specimen the flexuous sporules are from

60-90 X 2-2.5/x. Solidago latifolia is the only other species of

Solidago reported as bearing S. atropurpurea in Wisconsin. The

spots on S. altissima, paler in the center, are similar to those on

S. latifolia.

Hendersonia typhae Oud. on Typha angustif olia. Septem¬

ber 8, 1944. Doubtfully parasitic. Davis made an earlier collec¬

tion on this host species, but did not refer to it as such. A

Wisconsin specimen placed in the herbarium as STAGONOSPORA

typhoidearum (Desm.) Sacc. has deeply olivaceous spores and

seems to be H. typhae Oud.

Ramularia aequivoca (Ces.) Sacc. on Ranunculus pennsyl-

vanicus. July 12, 1944. Not listed by Seymour on this host.

Leptothyrium similisporum (Ell. & Davis) Davis on Soli¬

dago altissima. September 1, 1943. Apparently a new host for

this fungus, with no reports from Wisconsin or elsewhere.

Helminthosporium sativum Pamm., King & Bakke on Muh¬

lenbergia foliosa. October 8, 1944. Numerous single spore cul¬

tures were obtained and submitted to Professor J. L. Allison,

Associate Pathologist, U. S. D. A., stationed at Madison, who

states that these show the cultural growth characteristics of H.

sativum. Not listed in Seymour on this host.

Cercospora caricina Ell. & Dearn on Car ex interior. July 1,

1944. This is a slender and somewhat depauperate development

of what is usually a decidedly more robust species.

Cercospora variicolor Wint. on Paeonia officinalis. Septem¬

ber 30, 1944.

Cercospora cord at ae Chupp & Greene on Zizia aurea. Wal¬

worth Co., Pickerel Lake, September 19, 1941; Waukesha Co.,

Eagleville, October 6, 1941, September 3, 1942; Dane Co., Mad¬

ison, July 2, 1943, June 11, 1944. This species was described on

Zizia cordata from material collected by J. J. Davis at Racine,

Wis. in 1890, and an additional specimen on the same host was

collected on the Scuppernong Prairie near Eagle, Waukesha Co.

in 1941. On Zizia aurea, in some cases, the conidiophores are

Greene — Notes on Wisconsin Parasitic Fungi . VIII 229

longer than the 35 /x maximum of the description, running to

50/a or slightly more.

Cercospora blephiliae Chupp & Greene on Lycoyus uni -

florus. September 1, 1944. Determined by Professor Chupp.

Additional Species

The fungi listed have not before been reported as occurring

in Wisconsin.

Bifusella faullii Darker on Abies balsamea. Vilas Co.,

July 27, 1902. Coll. J. J. Davis. Det. Dr. G. D. Darker. The

effused or labyrinthine pycnidia of this species occur together

with Phacidium balsameae J. J. Davis. Dr. Darker states that

B. faullii is the primary pathogenic agent, and that the Phacid¬

ium is secondary.

Puccinia millefolii Fckl. on Artemisia frigida. Pepin Co.,

Stockholm, August 25, 1935. Coll. N. C. Fassett and R. I. Evans.

Arthur’s Manual lists no station east of North Dakota for this

lepto- form.

Phyllosticta salicicola Thum. on Salix. sp. July 13, 1944.

Septoria caricis Pass, on Carex stricta. July 15, 1944. This

appears to be truly parasitic and shows close microscopic cor¬

respondence with Sydow’s Mycotheca germanica No. 2199.

Septoria caraganae P. Henn. on Caragana arborescens

(cult.). Washburn Co., Spooner, August 14, 1944. Coll. E. E.

Honey. On the grounds of the University of Wisconsin Experi¬

mental Farm. Apparently not before reported from the United

States.

Septoria menthae (Thum.) Qud. on Monarda fistulosa,

September 1, 1944. Referred here with some doubt. The pycnidia

are most inconspicuous, and detectable only in sections. The

sporules are mostly from 40-55 X 1.5-2 /x. That S. menthicola

Sacc. & Let. is distinct from S. menthae seems questionable.

S. menthae is the earlier name. European material in the Wis¬

consin Herbarium labeled S. menthae has spores of the dimen¬

sions specified for S. menthicola and there do not seem to be

any well-marked differences in the host lesions. (An interesting

member of the Staurosporae occurs together with the Septoria

on some of the spots. The small, hyaline, 3-radiate conidia are

borne on almost obsolete conidiophores which are aggregated in

230 Wisconsin Academy of Sciences , Arts and Letters

rudimentary acervuli. This is probably not a parasite.)

Hyalothyridxum calamagrostidis sp. nov.

Maculis amphigenis, ellipsoideis vel fusiformibus, aliquoties

confluentibus, supra pallidioribus cum marginibus angustis,

fusco-purpureis, .25-1.5 cm. longis; pycnidiis elementariis, sub-

epidermidibus, dispersis, profunde immersis, amphigenis, globo-

sis, erumpentibus, ostiolatis, 110-135/a diam., muris cum cellis

pallido-brunneis, planis, compressis, fere concoloribus cum prox-

imis cellis hospitibus; conidiophoris non distinctis, sed cellis

fertilibus infra in pycnidiis ; conidiis hyalinis, cellis multis, vari-

atis, anguste vel late ellipsoideus, ovatis, vel obovatis, 30-45 X

16-23/a, plerumque apiculate, apicibus hyalinis, 3-13 (plerumque

4-6) X 1*5— 2/a.

Spots amphigenous, ellipsoid or fusiform, sometimes conflu¬

ent, paler above with a narrow, dark purple border ; .25-1.5 cm.

long; pycnidia rudimentary, subepidermal, scattered, deeply

sunken in the host tissue, amphigenous, globose, erumpent,

ostiolate, 110-135/a diam., wall of pale brown, flattened, com¬

pressed cells, almost concolorous with surrounding host cells;

conidiophores not differentiated, but sporiferous tissue occupy¬

ing the lower half of the inner surface of the pycnidium; con-

idia hyaline, many-celled, variable, narrowly to broadly ellipsoid,

ovate, or obovate, 30-45 X 16-23/a, usually with a pointed hy¬

aline apiculum 3-13 (mostly 4-6) X 1-5-2/a.

On living leaves of Calamagrostis canadensis. Madison, Dane

Co., Wis., U. S. A., August 18, 1944. Coll. H. C. Greene. The type

specimen is deposited in the University of Wisconsin Crypto-

gami c Herbarium.

The divisions in the conidia have occurred in many planes so

that the cells are not generally in even tiers, but are opposite

to one another after the fashion of roughly hewn stones cement¬

ed in a wall. The individual cells are variable in size and shape,

but are often oblong, or rhomboidal, or otherwise angled. Fre¬

quently as many as twenty are in view simultaneously on one

side of a conidium. So striking, large, and unusual are the conidia

that it was at first doubted that the organism was a fungus, but

a germination test showed the development of large numbers of

rapidly elongating germ tubes, one per cell, so there seems no

question of the true fungal nature of H. calamagrostidis. This

is the hyaline counterpart of Camarosporium.

Greene — Notes on Wisconsin Parasitic Fungi . VIII 231

Marsonia gloeodes sp. nov.

Acervulis hypophyllis, dispersis, subepidermidibus, immersis,

sed cum conidiis in massis erumpentibus, 100-150/x diam. ca.;

conidiophoris curtissimis, prope obsoletis; conidiis hyalinis, cyl-

indraceis, cum septis mediis distinctis, leviter constrictis septis,

cum exoperidiis statim fractis, sed constanter affixis, 19-23

X 6-7/x.

Acervuli hypophyllous, scattered, subepidermal in origin,

sunken in host mesophyll, but producing erumpent masses of

conidia, 100-150/x diam. ca. ; conidiophores very short, almost

obsolete, not plainly differentiated from non-sporiferous tissue;

conidia hyaline, cylindrical, with distinct median septum, slight¬

ly constricted at septum, with a soon ruptured, but persistently

adherent outer wall or envelope, 19-23 X 6-7/x.

On living leaves of Fraxinus americana. Madison, Dane Co.,

Wis., U.S.A. August 4, 1944. Coll. H. C. Greene. The type spe¬

cimen is deposited in the University of Wisconsin Cryptogamic

Herbarium.

The conidia of M. gloedes are most distinctive, and when

first examined seemed to have thickened hyaline caps at their

ends, with appendages radiating from the caps. However, more

careful and extended observation shows that the supposed ap¬

pendages are merely ragged, free-hanging portions of the outer

envelope which remains more or less firmly attached at both

ends of the conidium, producing the appearance of caps on the

apices.

When examined under a hand lens this fungus, in its gross

appearance, is extremely suggestive of one of the Marsonia

fraxini-Septoria besseyi intergrading series (see Davis, Trans.

Wis. Acad. Sci. 21: 289, 1924) and at the time of collection was

thought to be such. The conidia are, however, utterly different.

Cylindrosporium celtidis Earle on Celtis occidental is. Au¬

gust 4, 1944. The specimen on which the description is based

occurred on leaves of Celtis mississippiensis from Alabama. The

spore characters are given as “cylindric or clavate, guttate, at

length obscurely several septate, 20-25 X 3/x.” The further state¬

ment is made that this somewhat closely resembles Cylindro¬

sporium ulmicolum Ell. & Ev. which, however, has spores 45-65

X 4/x. In the Madison specimen most of the spores are from

30-35 X 3/x, with extremes of 22-42/x being noted. Many of the

spores show considerable curvature. They are frequently clavate

232 Wisconsin Academy of Sciences, Arts and Letters

and in other particulars the Wisconsin material shows rather

close correspondence to the description. (Heald and Wolf de¬

scribed Cylindrosporium defoliatum on southern hackberry,

but this involved large portions of leaves, or entire leaves. The

spores are given as 30-42 X 3/*, cylindrical, which is, to be sure,

more in the range of my specimen on C. occidentals. Possibly the

two are conspecific. If so, C. celtidis is the earlier name.)

Cladosporium lysimachiae sp. nov.

Amphigenis, effusis, indeterminatis, fuligineis, in foliis cauli-

busque; conidiophoris plerumque solitariis, sed aliquoties in

fasciis (minus quam decern), fusca infra, apicibus pallido-

brunneis, erectis, vel aliquoties angulosis et laxis, nonnihil sinu-

osis, dense denticulatis, continuis vel 1-3-septatis, plerumque,

50-105 X 5jul; conidiis pallido-fuscis, fusoideis-cylindraceis, apic-

ulatis, constanter 1-septatis, leviter angustatis septis, 17-23

X 5-7p.

Amphigenous in diffuse, indeterminate, blackish-brown patch¬

es on leaves and stems; conidiophores usually single, but oc¬

casionally in small tufts (less than ten), dark brown below, tip

pale brown, erect or sometimes angled and spreading, somewhat

sinuate, closely geniculate in the upper half, tip often dentic¬

ulate, continuous or 1-3-septate, mostly 50-105 X 5^; conidia

pale fuscous, fusoid-cylindrical, definitely apiculate, usually pro¬

nouncedly so at one end, regularly 1-septate, slightly constricted

at septum, 17-23 X 5-7 /i.

On living leaves and stems of Lysimachia terrestris. Madison,

Dane Co., Wis., U.S.A., August 24, 1943. Coll. H. C. Greene.

Similar material on the same host was collected in August 1944

at the same station. Both the type specimen and the later collec¬

tion have been deposited in the University of Wisconsin Crypto¬

gam ic Herbarium.

In thin, appropriately stained sections the scanty internal

mycelium appears to be exclusively intraepidermal. The place of

origin of the conidiophores seems to bear no relation to the sto¬

mata! openings.

It should be pointed out that while many species of Clado¬

sporium have been described, most of the earlier descriptions are

inadequate, and it is difficult and often impossible to tell whether

a species which comes under observation has been previously de¬

scribed. However, it is felt that less confusion is created by pro¬

viding an adequate description and a new name, than by placing

Greene — Notes on Wisconsin Parasitic Fungi . VIII 233

a newly found form arbitrarily under an old and dubious name

to disconcert future investigators. So far as I have been able to

determine, only one other species of Cladosporium has been de¬

scribed on Primulaceae and that on a genus which does not

occur in the New World. It is, furthermore, obviously quite dif¬

ferent from the species on Lysimachia terrestris .

A specimen of the saprophytic and ineptly named Macro-

SPORIUM parasticum Thum. has been presented to the Wisconsin

Herbarium by Dr. E. E. Honey. In this connection it should per¬

haps be noted that the allegedly parasitic Macrosporium porri

Ell., likewise on Allium cepa, was reported many years ago as

occurring in Wisconsin, but was not recorded in Davis’ notes.

NOTES ON WISCONSIN PARASITIC FUNGI. IX.

H. C. Greene

University of Wisconsin , Madison

These notes, unless it is otherwise specified, are concerned

with collections of fungi made during the season of 1945. The

type specimens of the new species described are deposited in the

University of Wisconsin Cryptogamic Herbarium.

Desmodium acuminatum ( D . grandiflorum of Gray's Manual,

7th ed.) in the vicinity of Madison in the summer of 1945 was

heavily parasitized by a powdery mildew, presumably MlCRQ-

SPHAERA diffusa Cke. & Pk., but perithecia were not formed

since the mildew was in turn parasitized by Cicinnobolus.

Mature perithecia of Sphaerotheca humuli var. fuliginea

(Schl.) Salm. occur on the lower portions of plants of Veronica

serpyllifolia collected at Madison, May 12. This is very good

specimen material accompanied by conidia, indubitably the cur¬

rent season's, so it seems reasonable to assume that the perithecia

also developed in 1945. A rather long, abnormal warm spell in

April may account for this, for Wisconsin, phenomenally early

development of perithecia. On the same plants is found Ramu-

laria veronicas FckL, represented in the herbarium on this

host by a single previous specimen taken at Madison thirty

years ago.

Apothecia of Sclerotinia seaveri Rehm were collected April

8 at Madison on overwintered fruits of Prunus serotina. Speci¬

mens of the Monilia stage from Wisconsin are numerous, but

none of the perfect stage were in the herbarium. A week later

apothecia of Sclerotinia fructicola (Wint.) Rehm were found

on mummified fruits of wild plum, Prunus americana. This also

was not represented in the herbarium.

The ascigerous stage of Claviceps purpurea (Fr.) Tul. oc¬

curred in some quantity on fallen grains of Bromus inermis at

the University Farms in late June. Of the many previous collec-

285

236 Wisconsin Academy of Sciences , Arts and Letters

tions of this fungus in the herbarium there are none of the per¬

fect stage from Wisconsin or elsewhere in America, but there

are several specimens of European origin.

The growth known as Ophiodothis haydeni (B. & C.) Sacc.

is common on certain species of Aster in marshes in Wisconsin.

Fungi Columbiani 3042 is a good representation of this. Whether

it is a single organism or a group of organisms is question¬

able for the developments so far as observed, have been uniform¬

ly sterile. The original description of 0. haydeni is utterly

inadequate.

Previous collections of Entyloma aster-sericeanum Zund.

have all been made from mature host plants late in the season.

A specimen taken near Madison on April 12, however, shows

the fungus in good maturity on the basal leaves of developing

shoots of Aster sericeus. Evidently the fungus overwinters in

the crown of the host plant and develops concurrently with the

earliest leaves which dry up and fall away later in the season.

All previous specimens have been on the small upper stem leaves.

Puccinia extensicola Plowr. I on Hieracium longipilum

occurred in considerable abundance at Madison in May and

June. A poor specimen from Blue River, Grant Co., collected in

1935 by Davis and me provides the only previous Wisconsin

record on this host, nor does Dr. Cummins know of collections

from elsewhere.

The uredinoid aecia of Puccinia hieracii (Schum.) Mart,

were found on Hieracium canadense at Madison, May 29. There

do not appear to be any other Wisconsin specimens of this stage

of the rust.

Puccinia asteris Duby occurred in great abundance on

Aster oblongifolius var. angustatus Shinners on a hillside “prai¬

rie” in Dane Co., near Sauk City, August 1. A 1940 collection,

reported as being on Aster oblongifolius, appears likewise to be

on var. angustatus.

Plants of Aristida tuberculosa, heavily infected with telia of

Uromyces peckianus Farl., were gathered at Arena, Iowa Co.,

October 1, 1944. This material was overwintered out of doors,

and in the spring of 1945 an attempt was made to infect plants

of Plantago rugelii growing in the open. The plants were heavily

mulched with the infected grass in the latter part of April. No

infection had occurred by the first week in June, despite weather

• Greene— Notes on Wisconsin Parasitic Fungi . IX 237

conditions which were seemingly very favorable for such devel¬

opment. Apparently no cultures have been made with teliospores

from A. tuberculosa . According to Arthur, evidence has been

obtained of strongly marked racial characters, partly geographic

and partly dependent on host. Perhaps this has some bearing on

the present case.

A fungus which may be identical with Phoma sepinicola

(Kickx) Sacc. has been found on living stems of Rosa sp. The

pycnidia are gregarious, the conidia greenish-hyaline, mostly

about 6 X 8/a. The Phoma is on dark purplish areas, but on the

same stems there are conspicuous ellipsoid arid lesions with

many small pycnidia which contain microconidia, 4-5 X 2/a. It

seems doubtful that the latter are connected with the Phoma.

Perhaps they are the precursors of Diaporthe or some similar

form.

Phyllosticta discincta J. J. Davis was described on Uvu~

laria grandiflora (Trans. Wis. Acad. Sci. 16: 747, 1910). In

Davis' “Parasitic Fungi of Wisconsin,” p. 63, it is erroneously

stated that Oakesia sessilifolia was the host. The original packet

contains indubitable leaves of Uvularia. Davis later expressed

doubt as to the validity of Ph. discincta, but I have recently

made a collection of identical material on the same host. The

spores are small and do not resemble those of the Ph. cruenta

complex, so it seems quite probable that this is a good species.

What is taken to be Macrophoma tiliacea Peck is prevalent

on dead branches and twigs of specimens of Tilia americana on

the University of Wisconsin campus. This may be a weak para¬

site. Many of the large, widely ostiolate pycnidia are v/ithout

contents, but when found, the spores are uniformly hyaline and

without any suggestion of gradation into Sphaeropsis.

Diplopia taxi (Sow.) DeNot occurred on languishing foliage

of Taxus sp. in a nursery at Madison, March 22. Probably, but

not certainly, saprophytic. (On leaves of the same plants, but

so far as observed not associated with the Diplodia, there oc¬

curred an Ascomycete which is doubtfully referred to Phacid-

ium. The fructifications are almost, if not quite aparaphysate,

the large asci about 105-110 X 20-25/a, and the hyaline, oblong-

ellipsoidal ascospores 30-33 X 12-14/a. Even if. this properly

belongs in Phacidium, on the basis of dimensions it cannot be

Phacxdium taxi Fr. or P. taxigola Dearn. & House. Davis has

reported the former from Wisconsin.)

238 Wisconsin Academy of Sciences , Arts and Letters

In an earlier paper (Farlowia 1: 575, 1944) mention was

made of Stagonospora on Andropogon furcatus and other grasses

in Wisconsin. It was stated that the spores were about of the

order of 30-35 X 10-12/x, several septate and markedly guttu-

late. In an additional collection, made August 4, 1945, on Andro¬

pogon furcatus of what is plainly the same thing, the mature

conidia are from about 40-55 X 8-13^. The pycnidia are sub¬

cuticular and can be easily lifted from the leaf with a needle.

This fungus evidently occurs consistently on A furcatus and

perhaps is distinct from similar forms on Sorghastrum and

Panicum.

A Septoria of the S. solidaginicola-fumosa-davisii complex

on Aster tradescanti from Madison has the characters which

are usually assigned to S. fumosa Peck. However, this species

has seemingly not been reported on Aster before.

A specimen of Septoria atropurpurea Peck on Aster macro-

phyllus from Devil’s Lake, Sauk Co., has exceedingly large mul-

tiseptate sporules, up to 120 X 3/x. They are extruded in eirrhi.

In a collection of Marsonia delastrei (Delacr.) Saac. on

Silene stellata, made near Mazomanie, Dane Co., many of the

conidia are biseptate and 30/x or more long.

A doubtfully parasitic fungus which has been determined as

Pestalotia funerea Desm. is abundant on blighted needles of

cuttings of Pinus strobus from a nursery at Wisconsin Rapids.

Coll. A. J. Riker, June 6, 1944. Det. M. P. Backus.

Limber and Cash (Mycologia 37: 129, 1945) state that the

fungus which has been known as Leptothyrium dryinum Sacc.,

and which occurs on oaks in Wisconsin, is not properly a mem¬

ber of the Leptostromataceae. Following Gilman and Archer

they consider that the proper name of the organism is Actino-

plete dryina (Sacc.) Hoehn.

In my experience Hyphomycetes sometimes, probably com¬

monly, overwinter by means of the formation of sclerotia which

frequently resemble immature perithecia, but which, instead of

forming asci, merely give rise to large numbers of conidia, fol¬

lowing spring rains. In the spring of 1945 at Madison leaves

of Pentstemon digitalis bearing these “pseudoperithecia” on old

spots of Cercospora pentstemonis Ell. & Kell., and leaves of

Zizia aurea bearing similar structures on old spots of Cerco¬

spora cordatae Chupp & Greene, were found to bear numerous

tufts of conidia produced on short conidiophores from the sur-

Greene — Notes on Wisconsin Parasitic Fungi. IX 239

face of the sclerotial elements. A similar condition was noted

the year previously for Cercoseptoria leptosperma (Pk.) Petr.

Cercosporella saxifeagae Rostr. on Saxifraga Pennsyl¬

vania was found in abundant development in the Wisconsin

River bottoms near Sauk City, May 30, and also at Madison,

August 9. The only earlier collection was made by Davis at

Portage, Columbia Co., in 1927.

Additional Hosts

Not previously recorded as bearing the fungi mentioned in

Wisconsin.

Peronospora parasitica (Pers.) Fr. on Cardamine parvi-

flora var. arenicola. Iowa Co., Arena, May 23. Apparently the

first report of the fungus on this host.

Sphaerotheca humuli (DC.) Burr, on Humulus ameri-

canus. Dane Co., Madison, July 9. Perithecia apparently not

hitherto recorded from Wisconsin on this host, although conidia

are often developed.

Claviceps purpurea (Fr.) Tul. on Calamagrostis neglecta.

Sclerotia were observed on a specimen of C. neglecta in the phan¬

erogamic section of the University of Wisconsin Herbarium. The

specimen was collected by J. R. Heddle at Madison, July 1909.

Coleosporium solidaginis (Schw.) Thlim. II, III on Solidago

rigida. Dane Co., Madison, July 9.

PUCCINIA EXTENSICOLA Plowr. 0 on Aster azureus. Sauk Co.,

Cactus Bluff, Town of Prairie du Sac, July 7. The lesions had

apparently reached the limits of development and a number of

leaves from different plants over a wide area showed no mature

aecia, although the arnphigenous pycnia were well formed. On

Aster laevis. Columbia Co., Black Hawk's Lookout, near Prairie

du Sac, June 17. The North American Flora lists this on A. laevis

from Colorado, Nebraska and New Mexico, so the present col¬

lection represents a decided eastward extension of range insofar

as the particular host is concerned. On Aster lucidulus. Dane

Co., Madison, June 19. This listing is made since A. lucidulus

is now set out from A. puniceus, although Davis reports the rust

on A. puniceus . (Mature aecia are present on A. laevis and A.

lucidulus) .

240 Wisconsin Academy of Sciences , Arts and Letters

Puccinia eleocharidis Arth. I on Eupatorium maculatum.

Dane Co., Madison, June. The host is the common species which

occurs in marshes in Wisconsin and was not formerly differen¬

tiated from the woodland species, E . purpureum. Other para¬

sites which, in my personal experience, occur on E. maculatum

in Wisconsin are Erysiphe cichoracearum DC. and Cerco-

spora perfoliata Ell. & Ev.

Puccinia rubigo-vera (DC.) Wint. II, III on Elymus glaucus.

Waushara Co., Hancock, August 1944. Coll. Etlar Nielsen; II

on Agropyron intermedium . Dane Co., Madison, July 1944. Coll.

Etlar Nielsen. Both these hosts were cultivated.

Phyllosticta decidua Ell. & Kell, on Verbena stricta. Col¬

umbia Co., Black Hawk’s Lookout near Prairie du Sac, June 17 ;

On Epilobium sp. Dane Co., Madison, July 26. Not previously

reported on any species of Epilobium in Wisconsin. As Davis

noted for a collection of this fungus on Agrimonia striata (Trans.

Wis. Acad. Sci. 19(2): 678, 1919), the conidia are decidedly

brownish, and it is a question whether the fungus is properly

placed under Phyllosticta.

Phyllosticta liatridis J. J. Davis on Liatris spheroidea.

Dane Co., Madison, August 4. As in an earlier collection on

Liatris spheroidea f. benkei (the white-flowered form of L.

spheroidea) , the conidia are somewhat smaller than those of the

original collection on Liatris spicata . In type of lesion and in

other respects there is great similarity however.

For the sake of a record a microconidial fungus, so-called

Phyllosticta angelicae Sacc., which occurs commonly in the

fall on Angelica atropurpurea in the vicinity of Madison, is listed

here. This is connected with Fusicladium depressum (B. & Br.)

Sacc., and both are supposedly stages of Phyllachora angel¬

icae (Fr.) Fckl. The latter, however, is included in Saccardo’s

enumeration of dubious Phyllachorae whose perfect stages are

not known, and so far as I am aware the existence of such a

stage has never been verified for Ph. angelicae. In an earlier

note (Trans. Wis. Acad. Sci. 3^: 85, 1942) I reported the lack of

further development, following overwintering, on several oc¬

casions, of such material. At that time I had overlooked the

existence of the name Phyllosticta angelicae, so failed to

mention it.

Greene — Notes on Wisconsin Parasitic Fungi . IX 241

Stagonospora bromi Smith & Ramsb. on Bromus kalmii.

Sauk Co., Cactus Bluff, Town of Prairie du Sac, July 7. A scanty

collection, but entirely characteristic. On Bromus ciliatus . Dane

Co., Madison, July 19. Collected in quantity in the Lake Wingra

marsh. On Bromus purgans. Sauk Co., Devil's Lake, July 27.

Although typical leaf lesions are numerous, the pycnidia are very

sparingly produced in this specimen.

Septoria gommonsii Ell. & Ev. on Girsium altissimum. Sauk

Co., Cactus Bluff, Town of Prairie du Sac, July 7. This is the

host from which Ellis and Everhart described S. COMMONSII.

That this species is really distinct from Septoria cirsii Niessl

is questionable.

Septoria angularis Dearn. & Barth, on Solidago latifolia.

Columbia Co., Gibraltar Rock near Okee, July 1. The sporules

are 26-88 X 1.5/*. As the authors indicated, this is probably very

close to, if not identical with, Septoria fumosa Peck. The spots

are distinctly larger and squarer than those which I have been

accustomed to associate with S. fumosa.

Septoria cacaliae Ell. & Ev. on Gacalia tuberosa. Dane Co.,

Madison, August 9. Of the species of Cacalia on which this has

been collected in Wisconsin ( C. atriplicifolia, C . reniformis , C.

suaveolens, C. tuberosa) only C. atriplicifolia shows well-marked

lesions with a distinct, narrow purple border and arid center.

While on the other hosts the fungus usually occurs on definite

lesions, they are uniformly dark and lack differentiation between

center and margin. However, a specimen in the herbarium from

Kansas on C . tuberosa shows lesions of a type intermediate be¬

tween the extremes described.

Septoria lactucicola Ell. & Mart, on Lactuca spicata . Dane

Co., Madison, June 24. Davis reports Septoria unicolor Wint.

on this host, but in the present material I am unable to see that

the organism is other than S. lactucicola. It appears probable

that these species are one and the same, and since the Ellis and

Martin name is the earlier, it seems best to use it.

Marsonia potentillae (Desm.) Fisch, var. tormentillae

Trail on Fragaria vesca var . amerieana . Dane Co., Pine Bluff,

June 29. Listed as the variety because of the relatively small

conidia which do not exceed 18/*. Also from Madison and from

Gibraltar Rock, Columbia Co.

242 Wisconsin Academy of Sciences, Arts and Letters

Septogloeum convolvuli Ell. & Ev. on Convolvulus arvensis .

Dane Co., Madison, June 21. Placed here with some doubt. It is

probable that Septogloeum convolvuli and Stagonospora con¬

volvuli Dearn. & House intergrade. Some of the conidia in the

present specimen are up to 35 X 5/x and the walls of the fruiting

body are rather well formed.

Ramularia virgaureae Thiim. on Aster laevis. Columbia

Co., Black Hawk's Lookout near Prairie du Sac, July 18. In

general Davis' remarks (Trans. Wis. Acad. Sci. 2U: 274, 1929)

concerning a collection of this variable organism on Aster sagit-

tifolius apply to the specimen on A. laevis. The conidia here are

markedly catenulate with a considerable range in length. On

Solidago juncea, Dane Co., Madison, July 30. Definitely a Ramu¬

laria with no suggestion of Cercosporella.

Cercospora parvimaculans J. J. Davis on Solidago juncea .

Dane Co., Madison, July 30.

Tuberculina persicina (Ditm.) Sacc. on Puccinia panici

Diet. I on Euphorbia corollata. Dane Co., Madison, July 31. On

Uromyces acuminatus Arth. I on Phlox pilosa . Dane Co., near

Sauk City, May 30. The host is U. acuminatus polemonii (Pk.)

J. J. Davis of Arthur's Manual. Davis collected this on U. ACUMI¬

NATUS steironematis (Arth.) J. J. Davis where the phanero¬

gamic host was Steironema ciliatum.

Additional Species

The following species have not before been listed in these

or in Davis' notes as occurring in Wisconsin.

Sporodinia grandis Link on Boletus castaneus. Dane Co.,

Madison, July 25. Coll. E. J. Backus. Det. M. P. Backus. On

Strobilomyces strobilaceus. Dane Co., 6 mi. west of Black Earth,

August 12. Coll. & det. M. P. Backus.

Chamberlain and Allison (Phytopath. 85: 241, 1945) report

studies of Pyrenophora bromi (Died.) Drechsler, the perfect

stage of Helminthosporium bromi Died., on the basis of ma¬

terial on Bromus inermis from the grass nurseries at the Wis¬

consin Agricultural Experiment station in Madison.

Phomopsis excelsior Tapy on twigs of Quercus velutina.

Sauk Co., T12N, R5E S9 NW/SE, Highway 33, June 5. Coll. &

det. E. E. Honey.

Greene — Notes on Wisconsin Parasitic Fungi. IX 243

Coniothyrium concentricum (Desm.) Sacc. on Yucca fila~

mentosa. Dane Co., Madison, June 19. Although this occurred

on the still green leaves it is perhaps but doubtfully parasitic.

The host plant is growing on a roadside bank in the University

of Wisconsin Arboretum and is one of the southern species

which are hardy in this region.

Phyllosticta umbrino-fumosa sp. nov.

Maculis plerumque orbicularibus, 3-8 mm. diam., fuligineis,

immarginatis ; pycnidiis flavido-brunneis tinctis fumosis, 100-

. 125/a diam., applanatis, immersis, gregatim, ostiolatis; conidiis

hyalinis, oblongatis vel ellipsoideis, plerumque 2-guttulatis, 4-7

X 2.5-3/a.

Spots mostly orbicular, 3-8 mm. diam., fuliginous, immar-

ginate ; pycnidia sooty yellowish brown, 100-125 diam., flattened,

immersed, closely clustered, ostiolate; conidia hyaline, oblong

or ellipsoidal, mostly 2-guttulate, 4-7 X 2.5-3/a.

On leaves of Eupatorium urticae folium. Madison, Dane Co.,

Wisconsin, U. S. A., August 7, 1945.

The inconspicuous pycnidia are apparent only by transmitted

light, when they appear as translucent areas in the smoky spots.

Three other species of Phyllosticta which have been de¬

scribed on Eupatorium, and some of the principal characters in

which they differ from Ph. umbrino-fumosa are as follows:

Ph. eupatoriicola Kab. & Bub. Spots sordid, obscure brown,

immarginate; conidia 3-4 X 1-1. 5/a.

Ph. eupatorina Thum. Spots grayish with wide, dark purple

margin; conidia 6 X 2.5-3/a. The conidia here are similar.

Ph. eupatorii Allesch. Spots dark gray, with obscure purple

margin; conidia 10-15 X 3-4/a.

Phomopsis diachenii Sacc. on Pastinaca sativa. Dane Co.,

Madison, July and August. On fruit and living leaves. In gen¬

eral, on the fruit the fusoid conidia run 6-12 X 3/a, instead of

the 8-10 X 3/a of the description. The scolecospores are nar¬

rowly tapered at one end, frequently strongly curved at the

same end, and are from about 14-20 X 1.5-2/a. On leaves, both

pycnidia and their contents run somewhat larger than on the

dry papery earpels, but otherwise the microscopic characters

are practically identical. Although the description indicates that

244 Wisconsin Academy of Sciences , Arts and Letters

the fungus occurs on fruit only, in lack of evidence to the con¬

trary I assume that but one organism is involved since (1) the

host is the same (2) the microscopic aspect is the same (3)

the affected plants were identical or in close proximity and (4)

the infection occurred at the same time in late July and early

August. No spots occur on the fruits, which bear many pycnidia

evenly dispersed and which differ from non-infected fruits

principally in being somewhat paler. On leaves, however, the

spots are striking, variable, mostly somewhat ellipsoidal, usually

about 2-5 mm. long diam., although sometimes irregular and

considerably larger by confluence. The margins are deep purple,

of varying width, and contrast strongly with the pale to reddish-

brown centers. There are from one to several spots per leaflet,

and they are most conspicuous. Pycnidia are produced sparingly

on the leaves, in contrast to their abundant development on the

fruit. P. diachenii has also been found on fruits of the closely

related (to Pastinaca) Heracleum lanatum in a collection made

by Davis at Mineral Point, Iowa Co., August 10, 1927. This was

labeled Gloeosporium acheniicolum Rostr., but there can be no

doubt that it is a characteristic Phomopsis. Therefore, the rec¬

ord of Gloeosporium acheniicolum for Wisconsin must be

deleted.

Stagonospora brachyelytri sp. nov.

Maculis pallido-brunneis, marginibus angustis fuscis, ple-

rumque elongatis, 5-15 X 1.5 mm., venis sequentibus; pycnidiis

brunneis, subglobosis, 100-150/* diam., plerumque 100-110/*, ob¬

scure ostiolatis; conidiis hyalinis, granulosis, subciavatis vel

cylindraceis, saepe curvatis leniter, distincte 1-3-septatis, 13-

33 X 5-7/*.

Spots pale brown with narrow dark brown margins, mostly

elongate, 5-15 X 1.5 mm., following venation; pycnidia brown,

subglobose, 100-150/* diam., mostly 100-110/*, indistinctly ostio-

late; conidia hyaline, granular, subclavate or cylindrical, often

slightly curved, distinctly 1-3-septate, 13-33 X 5-7/*.

On leaves of Brachyelytrum erectum. Devil’s Lake, Sauk Co.,

Wisconsin, U. S. A., July 27, 1945.

Many conidia were measured and, while the extremes are

as given, those conidia which have less than 3 septa and are

shorter than 22-23/* are exceptional.

Greene— Notes on Wisconsin Parasitic Fungi . IX 245

Stagonospora cryptotaeniae sp. nov.

Maculis conspicuis, orbicularibus, centris fuscis, marginibus

flavidis, latis, ca. .5-1 cm. diam. ; pycnidiis hypophyllis, fumosis,

muris exilibus, magnis, frequenter 250/* diam. distincte ostio-

latis; conidiophoris fere obsoletis vel obsoletis; conidiis 0-3-

septatis, plerumque 1-2-septatis, hyalinis granulosis, oblongatis

vel irregularibus, 25-55 X 14-16/*.

Spots conspicuous, orbicular, with dark brown center and

wide yellowish border, about .5-1 cm. diam., pycnidia hypophyl-

lous, sooty, thin-walled, large, frequently 250/* diam., or some¬

what larger, ostiole well-defined; conidiophores almost or quite

obsolete ; conidia 0-3~septate, mostly 1- and 2-septate, hy¬

aline, granular, oblong or somewhat irregularly shaped, 25-

55 X 14-16/*.

On leaves of Cryptotaenia canadensis. Madison, Dane Co.,

Wisconsin, U.S.A., June 24, 1945.

When fresh, the large pycnidia are noticeably erumpent.

They tend to collapse, however, on drying and pressing, as

would be expected where the walls are so thin and the host

substrate of such delicate texture. There is seemingly a com¬

plete absence of reports of Stagonospora on Umbelliferae in

America, and in Europe, but there is no doubt that this is a

representative species of the genus.

In 1929 Davis (Trans. Wis. Acad. Sci. 2 A: 290, 1929) men¬

tioned, with descriptive notes, a species of Septoria which he

had found at Portage on Lithospermum canescens. What is

plainly the same fungus, on the same host, has recently been

found in some quantity at Madison, so a formal description

seems justified. Davis filed his specimen as Septoria litho-

spermi nom. herb. It seems proper to employ this name for the

new species which is described as Septoria lithospermi Davis

& Greene.

Septoria lithospermi Davis & Greene sp. nov.

Maculis parvis, irregularibus, orbicularibus vel angularibus,

fuscis ; pycnidiis epiphyllis, subepidermidibus, supra imperfectis,

globosis, brunneis, 60-70/* diam.; conidiis hyalinis, obscure sep-

tatis, plerumque curvatis, 35-65 X ,2-3/*.

Spots small, irregular, orbicular or angled, dark brown;

pycnidia epiphyllous, subepidermal, imperfect above, globose,

brown, 60-70/* diam.; conidia hyaline, obscurely septate, usually

curved, 35-65 X 2-3/*.

246 Wisconsin Academy of Sciences , Arts and Letters

On leaves of Lithospermum canescens. Madison, Dane Co.,

Wisconsin, U. S. A., July 25, 1945.

The pycnidia are most inconspicuous and can be detected

only in sections. As Davis stated, Lithospermum in Wisconsin

is usually notable for lack of parasites.

Gloeosporium hepaticae Peck on Hepatica acutiloba. Dane

Co., Madison, May 12. Dearness & House (Report of State

Botanist of New York, 1919, p. 35) described Gloeosporium

acutiloba which is said to differ from G. hepaticae in having

larger, chiefly hypophyllous acervuli and much smaller spores,

mostly 7 X 2/a, as opposed to 15-25 X 6.5-7.5/a for G. HEPATICAE.

In my Notes III (Trans. Wis. Acad. Sci. 35: 118, 1943) I

mentioned, with descriptive comments, what appeared to be a

species of Colletotrichum with curved, appendaged conidia, oc¬

curring on Andropogon furcatus and Sorghastrum nutans . Dr.

D. A. Preston has pointed out that this is probably Ellisiella

caudata Sacc., and so it seems to be, although whether Ellisiella

can really be generically distinct from Colletotrichum seems

doubtful to me.

Ramularia ovularioides sp. nov.

Maculis irregularibus, plerumque elongatis, supra fusco-brun-

neis, infra pallidioribus, frequenter totis segmentis foliis co-

opertis ; conidiophoris hyalinis, floccis 8-12, divergentibus, 20-30

X 3-3.5/a, nascentibus ab basibus gibbosis parvis, brunneis,

substomatibus ; conidiis hyalinis, ellipsoideis vel cylindraceis,

10-25 (plerumque 10-15) X 3.5-4.5/a, 0-1-septatis.

Spots irregular, mostly elongate, dark brown above, paler

below, frequently involving entire segments of leaf; conidio-

phores hyaline, in tufts of 8-12, widely spreading, 20-30 X

3-3.5/a, produced from small, brown, substomatal, tuberculate

bases; conidia hyaline, ellipsoid or cylindrical, 10-25 (mostly

10-15) X 3.5-4.5/a, 0-1-septate.

On leaves of Anemone cylindrical Madison, Dane Co., Wis¬

consin, U. S. A., July 31, 1945.

This species is very destructive to the host plant. All plants

in a group of 25 or 30 were seriously blighted. There is little

evidence of catenulation. Most of the conidia are short and non-

septate, reminiscent of Ovularia. A small collection of this

species was made in 1942 and discussed in an earlier note

(Trans. Wis. Acad. Sci. 35: 119, 1943).

Greene— Notes on Wisconsin Parasitic Fungi . IX 247

Mycogone roseola Pound & Clements on Helvetia elastica.

Columbia Co., Gibraltar Rock near Okee, July 1. This has the

appearance of a vigorous parasite.

Helminthosporium avenae Eidam on Arena sativa. Dane

Co., Madison, July 5. This common species seems not to have

been previously mentioned in Davis’ notes or in my current

series, so is therefore included for the sake of a complete record.

Professor J. G. Dickson has presented the herbarium with a

specimen of the perfect stage of this fungus, Pyrenophora

avenae Ito, on oat straw, Monticello, Green Co., June 12. (Al¬

though the propriety and legality of continued usage of the

name of an imperfect stage after demonstration of the con¬

nected perfect stage may be questioned, I feel that in many

cases such a name is more useful than that of the little known

and rarely encountered perfect stage, especially where parasites

are concerned).

A species of Cercospora which was found on leaves of Tri-

osteum perfoliatum was sent to Professor Chupp for inspection

and determination. He regards it as a new species so it is

described jointly with him as Cercospora triostei Chupp &

Greene sp. nov.

Cercospora triostei Chupp & Greene sp. nov.

Maculis subcirculis vel irregularibus, 2-4 mm. diam., con-

stanter umbrinis vel cum cingulis fuscis; fructificationibus am-

phigenis; floccis minutis, nigris; stromatibus parvis, 20/* diam.

vel minus, cellis fuscis compositis; fasciis cum 2-12 cauliculis

divergentibus ; conidiophoris olivaceo-brunneis pallidis vel medi-

is, interdum angustatis apicibus, parce septatis non ramosis,

G-4-geniculatis, rectis vel curvatis vel tortuosis, apicibus ob-

tuso-rotundatis vel subtruncatis, 3-5 X 30-120/*; conidiis oliva-

ceis pallidis vel mediis, obclavato-cylindraceis, rectis vel leniter

curvis, 1-7-septatis, basibus obconico-truncatis, apicibus brevi-

bus, conicis, 3-5.5 X 30-70/*.

Spots subcircular or irregular, 2-4 mm. diam., uniformly

dull brown or with a narrow dark margin; fruiting amphige-

nous; pustules minute, black; stromata small, up to 20/* diam.,

made up of dark brown cells; fascicles 2-12 divergent stalks;

coriidiophores pale to medium olivaceous brown, fairly uniform

in color, sometimes more narrow near the tip, sparingly sep¬

tate, not branched, 0-4-geniculate, straight to curved or tor¬

tuous, bluntly rounded to subtruncate tip, 3-5 X 30-120/*;

248 Wisconsin Academy of Sciences, Arts and Letters

conidia pale to medium olivaceous, obclavate-cylindric, straight

to mildly curved, 1-7-septate, base obconically truncate, tip

short, conic, 3-5.5 X 30-70 /x.

On leaves of Triosteum perfoliatum. Pine Bluff, Dane Co.,

Wisconsin, U.S.A., June 29, 1945.

Cercospora lycii Ell. & Halst. on Lycium halimifolium. Dane

Co., Madison, August 12.

Plakidas (Phytopath. 35: 185, 1945), following Chupp, con¬

cludes that the fungus on Juniperus communis var. depressa

(and also on Juniperus virginiana) which has been called Cerco¬

spora SEQUOIAE var. juniperi Ell. & Ev., common in Wisconsin

and originally described from Wisconsin material, is not prop¬

erly a Cercospora because the conidia are echinulate and because

they are produced from a thick, more or less convex sclerotial

stroma, considered to be a sporodochium. It appears that this

is probably Exosporium deflectens Karst., although the conidia

show a much greater range in length than the 14-20/x of Kar-

sten’s description. The specimens in the herbarium have been

relabeled as Exosporium deflectens, and the name Cercospora

sequoiae var. juniperi must be deleted.

THE WISCONSIN SPECIES OF PELTIGERA

Papers on Wisconsin Lichens , No . 2,

John W. Thomson, Jr.

Department of Botany , University of Wisconsin

In the recent revival of interest in lichens in the United

States, need has been felt for bringing up to date the taxo¬

nomic treatments of the North American lichens. Among those

groups in need of revision is the genus Peltigera. A series of

papers by V. Gyelnik of Budapest, in preparation toward a

monograph of the Peltigeras of the world, necessitates a criti¬

cal re-examination of our own species.

In the determination of Peltigera specimens, the usual spore

and apothecial characters by which other groups of lichens

are identified are in most cases of less importance than the

thallus characters. The character of the upper surface, the type

of venation of the under surface, and the nature of the rhizinae

are of prime importance. The rhizinae as classified by Gyelnik

(16) are: 1. simple, e.g. P . canina or P. polydactyla var. doli-

chorrhiza; 2. pappose, simple but with a tuft of hyphae at the

tip, e.g. P. canina var. spuria f. sorediata; 3. fasciculate, with

a number of more or less parallel strong rhizinae bound in a

bundle, e.g. P. polydactyla; 4. fibrillose, with a number of

lesser hyphae extending at right angles to the main rhizoid and

sometimes repeatedly branched and forming a spongy mat, e.g.

P. canina var. spongiosa . The presence or absence of soredia

and isidia also provide characters of some value. The Pelti¬

geras are a plastic group, much modified by the environment,

and variance of these characters sometimes makes it difficult

to place certain specimens. It would seem that transplantation

experiments would go far toward helping understand the spe¬

cies' limits in this genus. Field studies of this type are much

needed.

249

250 Wisconsin Academy of Sciences , Arts and Letters

Almost no mention of Wisconsin Peltigeras has been made

in the literature. I. A. Lapham (19) lists P. rufescens ( P .

canina var. rufescens) from Milwaukee. This specimen is now

in the Tuckerman Collection at the Farlow Herbarium, Harvard

University. Bruce Fink (4) lists Peltigera pulverulenta from

a collection made by L. H. Pammel at La Crosse. This speci¬

men, in the Herbarium of the University of Michigan, however,

is not this species but P. polydactyla var. crassoides.

The principal collectors of Peltigera material in Wisconsin

are L. S. Cheney, R. H. Denniston, and the writer. Dr. Ben-

niston’s private herbarium has now been given to the Uni¬

versity of Wisconsin and is being incorporated in the Univer¬

sity Herbarium. I am indebted for additional collections to

Dr. G. S. Bryan, Dr. R. I. Evans, Dr. L. H. Shinners, Robert

Ellarson, and Mrs. F. E. Lund, Jr. Unless otherwise stated,

the collections cited are by the writer. The full set of speci¬

mens is in the Herbarium of the University of Wisconsin.

Duplicates of many are in the private herbarium of the writer

and have also been used in exchanges. In as many cases as

possible the nomenclature has been checked back to the origi¬

nal publications. Spore measurements were made in water

mounts.

Grateful acknowledgment is made to Dr. David H. Linder

of the Farlow Herbarium, Harvard University, for making

fully available the facilities of that herbarium during the prog¬

ress of this study.

This work was supported in part by a grant from the Spe¬

cial Research Fund of the University of Wisconsin.

Key to the Wisconsin Peltigeras

1. Algae of thallus Chlorophyceae ; warty cephalodia containing Nostoc

on the upper or under surface of the thallus. Sect. Phlebia Wallr.

2. Upper surface of the thallus with warty cephalodia; under surface

with scattered, fasciculate rhizinae; thallus and lobes large, up to

6 cm. across; apothecia vertical.

3. Underside lacking veins, covered with an even tomentum, bluish

or brownish black at the center, pale white or brownish at the

margin, la. P. apkthosa var. typica.

3. Underside with brown veins, interspaces pale, center blackish,

margins paler.

4. Margins not crisped. ... lb. P. apkthosa var. variolosa

4. Margins crisped. . . . 1c. P. apkthosa var. variolosa f. crispa

Thomson— The Wisconsin Species of Peltigera 251

2. Lower surface of the thallus with small black cephalodia on the

veins; under surface attached centrally or at one side by a group

of rhizinae ; thallus small, up to 2 cm, across, apothecia horizon¬

tal. ... 2 . P. venom

1. Algae of thallus Myxophyceae; thallus lacking cephalodia. Sect. Em-

prostea (Ach.) Vainio.

5. Thallus upperside smooth cr verruculose-scabrid, lacking tomentum.

6. Upper surface smooth, shining, lacking scattered cylindrical isidia

but sometimes with flat, squamulose, isidia; underside dark, vein¬

less or with broad flat veins which are not strongly raised as in

P. canina.

7. Underside with broad, flat veins.

8. Apothecia horizontal.

9. Lacking isidia. . . . 8. P. horizontalis

9. With squamulose, flat isidia ... 3a. P. horizontalis f.

zopfi.

8. Apothecia vertical.

10. Rhizinae short, mostly fasciculate. . . . 4a. P. poly-

dactyla var. typica

11. Apothecia over 2 mm. long, lobes more than a cm.

across.

12. Lacking isidia.

13. Thallus lobes with margins raised, more or

less undulate but not crisped. ... 4a. P.

poly dactyla var. typica

18. Thallus lobes with dilacerate or dilacerate-

crisped margins. . . . 4d. P. polydactyla f.

lophyra

12. With flat isidia, especially along the margins

and cracks in the thallus ... 4c. P. polydacty¬

la f. microphylla

11. Apothecia about 2 mm. long, plant small, lobes less

than 1 cm. across. . . . 4b. P. polydactyla f. mi¬

cro car pa

10. Rhizinae long, over 5 mm., simple. 4e. P. polydactyla

■ var. dolichorrhiza

7. Underside lacking veins; plants small, thick, pusilloid; apo¬

thecia vertical; rhizinae short, fasciculate . . . 4f. P. poly¬

dactyla var. crassoides

6. Upper surface verruculose-scabrid, with scattered cylindrical

isidia; underside pale, veins narrow, strongly raised . . . 5. P.

evansiana

5. Thallus with a layer of tomentum on the upper surface, at

least toward the margins.

252 Wisconsin Academy of Sciences , Arts and Letters

14, Thallus large, lobes usually longer than 8 cm., prostrate,

with only the margins ascending; rhizinae simple or fibrillose.

15. Rhizinae fibrillose-confluent . . . 6g. P. canina var. spon-

giosa

15. Rhizinae simple or fibrillose but not confluent or forming

a spongy mat below.

16. Veins white, rhizinae white; plant usually of shady

places and bogs, pliable; grayish or brownish green in

color . . . 6a. P. canma var. albescens

16. Veins brown, often white at the margins, rhizinae

brown.

17. With small, squamulose isidia along the edge of

the thallus or on the upper surface, especially

along cracks. . . . 6d. P. canma var. rufescens f.

innovans

17. Lacking isidia

18. Lobes long and broad, over 1.5 cm. across,

veins brown to margin, plant pliable, growing

in shady places, tomentum of upperside usual¬

ly arachnoid. ... 6b. P. canina var. ulor-

rhiza

18. Lobes short and narrow, usually less than 1.5

cm. across, veins usually white near the mar¬

gin, brown to the center, plant brittle, often

splitting when pressed, growing in sunny places,

tomentum of upperside usually fairly dense.

... 6c. P. canina var. rufescens

14. Thallus small, lobes up to 3 cm. long, ascending, often

cochleate; veins raised, narrow, white at the margins, brown

to the center; rhizinae simple or pappose (sometimes fibril¬

lose in f. sorediata)

19. Thallus lacking soredia . . . 6e. P. canina var.

spuria

19. Thallus with coarse gray soredia in rounded spots

on the upper surface . . . 6f. P. canina var.

spuria f . sorediata

(1) Peltigera aphthosa (L.) WilldL, Flora Berolinensis. 347,

1787

Lichen aphthosus L., Sp. PL 1148, 1753.

Lichen caninus var. aphthosus Weis., Plant. Cryptog.

Flora Goettingens. 80, 1770.

Lichen verrucifer Gmelin., Syst. Natur. II, Pars. II, 1373,

1770.

Plate I. — 1. P. aphthosa, upperside; 2. P. aphthosa , underside; 3. P.

aphthosa var. variolosa , underside; 4. P. aphthosa var. variolosa f. crispa.

All Figures same scale as Plate 3.

Plate II.- — 1. P. polydactyla, underside; 2. P. polydactyla , upperside;

3. P. polydactyla var. dolichorrhiza (from a specimen from Maine det. by

Gyelnik) ; 4. P. polydactyla f. microcarpa; 5. P. polydactyla f. lophyra;

6. P. polydactyla f. microphylla. Figs. 5 and 6 enlarged to scale on Plate 5,

other Figures on same scale as Plate 3.

Plate III.— 1. P. horizontalis, underside; 2. P. horizontalis, upperside;

3. Three thalli of P. canina var. spuria; 4. Four pieces of the same thallus

of P. canina var. rufescens; 5. P. venosa, 4 thalli, uppersides, above;

lowersides below.

Plate IV. — 1. P. canina, upperside; 2. P. canina var. spongiosa, under¬

side; 3. P. canina var. ulorrhiza, underside; 4. P. canina var. albescens ,

underside. All to same scale as Plate 3.

i

Thomson— The Wisconsin Species of Peltigera 253

Lichen verrucosus Weber, Spicil. Flor. Goettingens. 273,

1778.

Peltigera amplissima Hoffm., Deutschl. Flora. 106, 1796.

Peltidta aphthosa Ach., Method. Lich. 287, 1803.

Peltidea aphthosa var. verrucosa Ach., Lich. Univ. 515,

1810.

Peltidea verrucosa Rohling, Deutschl. Flora III, 117, 1813.

Peltigera aphthosa var. phymatodes Wallr., Flora Cryptog.

German., Ill, 560, 1831.

Peltigera aphthosa f. verrucosa Dietrich, Lichenogr. Ger¬

man., 27, 1832-1837.

Thailus large, lobes broad, up to 10 cm. long and 6 cm. broad,

edges ascending; upper surface smooth erect tomentose toward

tips of lobes, with warty cephalodia scattered upon it, pale to

apple green when moist, leaden green when dry; underside pale

at the margins, blackening toward the center, veinless or with

white or dark veins, the veinless variety with an even, close, nap

of tomentum ; rhizinae scattered, fasciculate ; apothecia large, up

to 16 mm. across, on extended lobules, the sides reflexed, the mar¬

gins becoming crenate, the disk reddish-brown to blackish-brown ;

the hypothecium brown ; the hymenium hyaline ; paraphyses sim¬

ple, thickened and brown at the tips; asci cylindrico-clavate ;

spores acicular, 3-5 sepate, 49-53 X 4-5/x, in Wisconsin material.

On earth, rotting logs, humus, rocks.

la. Peltigera aphthosa var. typica.

Adams: Cold Water Canyon, 1894, L. S. Cheney; Douglas:

Amnicon Falls, 1942, Mrs . F. E. Lund , Jr.; upper Brule Bog

near highway P, 1943; west side of ravine of Black River,

Pattison State Park, 1942; Marathon: Wausau, 1894, L. S.

Cheney; Marinette: Pike River Falls, Amberg, 1939; Oconto:

Mountain, 1915, J. J. Davis; Polk: St. Croix Falls, 1927, W. F .

Typical Peltigera aphthosa is confined in Wisconsin to white

cedar bogs and cold ravines and is not as widespread nor as

abundant as var. variolosa. The typical variety is covered with

a soft, even brown or bluish-black tomentum below. It may

reach a larger size than the veined variety.

lb. Peltigera aphthosa var. variolosa (Mass.) Thomson,

Comb. Nov.

254 Wisconsin Academy of Sciences , Arts and Letters

Peltigera aphthosa f. variolosa Mass., Schedul. Critic. Ill,

64, 1856.

Peltigera aphthosa f. minor Tuck., Synops. N. A. Lich. I,

106, 1882.

Peltigera leucophlebia f. variolosa Gyel., Magy. Bot. Lapok

23: 79, 1925.

(non Peltigera aphthosa var. leucophlebia Nyl. (24) vide

Nilsson (23)).

Peltigera leucophlebia Gyel., Magy. Bot. Lapok 23: 79, 1925.

Peltigera aphthosa f. angustiloba Fokin et Nikolsk, Trudy

Vjastkago gosudarst Musej., 1926.

Peltigera variolosa Gyel., Magy. Bot. Lapok 25: 252, 1927.

Peltigera variolosa f. dactylodes Gyel., Nyt. Magazin for

Naturvid. 68: 269, 1930.

Adams: Cold Water Canyon, 1894, L. S. Cheney ; Ashland:

Bad River Gorge, Mellen, 1927, L. R . Wilson; Burnett : no lo¬

cality, 1927, L. R. Wilson ; Door: Washington Island, 1931,

J . J. Davis ; Douglas: Falls of Copper Creek, Superior, 1942;

ravine of Black River, Pattison State Park, 1942; Iron: west

of mouth of Montreal River, 1896; L. S . Cheney; Juneau: Wis¬

consin Dells, 1936; Sauk: Rocky Arbor State Park, Wisconsin

Dells, 1945; Vilas: Conover, 1897, L. S. Cheney.

Variety variolosa is characterized by the brown to black

venation of the underside of the thallus; otherwise the char¬

acters are as in the typical plant. This variety is more southern

and widespread in the United States than the typical plant

and is usually in warmer and drier habitats.

1c. Peltigera aphthosa var. variolosa f. crispa (Vainio) A

Zahlbr.

Peltidea aphthosa f. crispa Vainio, Meddel. Soc. Fauna et

Flora Fennica 3 : 99, 1878.

Peltigera aphthosa f. crispa A. Zahlbr., Cat. Lich. Univ.

Ill, 451, 1925.

Peltigera variolosa f. crispa Gyel. apud Erichs., Verhandl.

des Bot. Verein der Prov. Brandenb. 60: 223, 1928.

Marinette: Thunder River, Thunder Mt., 1937.

This form differs from the variety only in that the thallus

and margins are crisped.

Thomson^— The Wisconsin Species of Peltigera 255

2. Peltigera venosa (L.) Baumg., Flora Lipsiens. 581, 1790.

Lichen venosus L., Sp. PL 1148, 1753.

Peliidea venosa Ach., Method. Lich. 282, 1803.

Thai I us small, up to 2 cm. across, rounded, single, fan¬

shaped lobes, sometimes somewhat divided ; upper surface

smooth, shining, apple-green when wet, grayish- or brownish-

green when dry; underside with strong, dark brown or black

veins and white interspaces ; attached by a single group of rhi-

zoids at one side ; cephalodia on the veins, dark green to black ;

apothecia marginal, round, horizontal, up to about 5 mm. in

diameter, the margin crenate, the disk reddish to blackish

brown ; hypothecium brown ; hymenium hyaline ; spores hyaline

to brownish, (1- ) 3 septate, 24-37 X 6-8 p in Wisconsin material.

Among mosses on wet cliff faces.

Rare and thus far collected only on the cliffs on the west

side of the ravine of the Black River at Pattison State Park,

Douglas Co., 1942 and by J. W. Thomson, Jr. and R. I. Evans,

1944.

3. Peltigera horizontalis (Huds.) Baumg., Flora Lipsiens.

562, 1790.

Lichen horizontalis Huds., Flora Anglica. 543, 1762.

Lichen horizontalis var. nebulosus Vill., Hist. Plant. Dau¬

phin III, 960, 1789.

Peltidea horizontalis Ach., Method. Lich. 288, 1803.

Peltigera canina var. horizontalis March apud Hall, Vrolik

et Mulder, Bijdrag t. d. Natuurk. Wetensch 5: 198, 1830.

Peltigera rufescens var. horizontalis Sprengl., Flora Halens.,

ed. 2: 543, 1832.

Peltigera horizontalis f. lobatoides Gyel., Magy. Bot. Lapok

28: 61, 1929.

Thallus large, the rounded lobes up to about 3 cm. broad

and several cm. long; upperside smooth, shining, bluish-gray

or greenish gray when moist, lead gray or brownish gray or

brown when dry, lacking tomentum or cylindrical isidia but

sometimes producing tiny flat squamules or “isidia” when re¬

generating following injury; underside reticulate with broad

flat veins forming an almost confluent tomentum, the veins

brown to black, the interspaces white; rhizinae black, sparse,

fasciculate ; apothecia brown to chestnut-brown, round, horizon-

256 Wisconsin Academy of Sciences , Arts and Letters

tal, the margins crenate; hypotheciiim brown; hymenium hya¬

line; spores hyaline, fusiform, 3 septate, 25-45 X 3.5-6 /* in

Wisconsin material.

On soil, fallen logs, and mossy rocks in moist woods.

Ashland: Mellen, 1927, L. R. Wilson; Columbia: 5 miles

southwest of Portage, 1945 ; Douglas : upper Brule River,

1942; Pattison State Park; 1942; Winneboujou, 1942; Falls of

Copper Creek, Superior, 1942; road to Wisconsin Point, Su¬

perior, 1942; Forest: east of Wapigon Lake, 1945; Iron: shore

of Lake Superior west of Montreal River, 1896, L. S. Cheney ;

Marinette: Pike River Falls, Amberg, 1939; Oneida: Rhine¬

lander, 1893, L. S. Cheney; Sauk: Rocky Arbor State Park,

1945.

When the apothecia are not present, P. horizontalis is in¬

distinguishable from P. polydactyla. Many herbarium speci¬

mens thus cannot be satisfactorily placed in either species.

Specimens showing regeneration in the form of small flat

“isidia” or squamules have been named Peltigera Zopfi by

Gyelnik but I am doubtful that these should be segregated as a

distinct species. Following the practice in the Cladoniae (e. g.

Cladonia rangiferina f. prolifera, C. pyxidata var. neglecta f.

peritheta., C. caroliniana f. prolifera) the species of Gyelnik

becomes :

3a. Peltigera horizontalis f. zopfi (Gyel.) Thomson Comb.

Nov.

Peltigera Zopfi Gyel., Bot. Kozlem. 24: 134, 1927.

Wisconsin specimens which may fall into this category or

into the analogous form, Peltigera polydactyla f. microphylla

Anders, were collected at Pattison State Park, Douglas Co.,

1942. The specimens lack apothecia and cannot be placed in

either species with certainty.

4. Peltigera polydactyla (Neck.) Hoffm., Descript, et

Adumbr. Plant. Lich I, 19, 1790.

Lichen polydactilon Neck., Method. Muscor. 85, 1771.

Lichen caninus var. polydactylon Lightf., Flora Scotica II,

846, 1777.

Lichen caninus var. pellucidus Weber, Spicil. Flora Goet-

tingens. 270, 1778.

Thomson — The Wisconsin Species of Peltigera 257

Peltigera polydactylon Hoffm., Descript, et Adumbr. Plant.

Lich I, 19, 1790.

Peltidea polydactyla Ach., Method. Lich. 286, 1808.

Peltidea polydactyla var. pellucida Ach., Method. Lich. 287,

1803.

Peltidea hymenina Ach., Method. Lich. 284, 1803.

Peltidea glauca Pers. apud Ach., Lich. Univ. 518, 1810.

Peltidea pellucida S. Gray. A Natur. Arrang. Brit. Plants I,

429, 1821.

Peltidea horizontalis var. hymenina Ach., Synops. Lich. 238,

1814.

Peltigera canina var. attenuata March apud Hall, Vrolik et

Mulder, Bijdrag t. d. Natuurk. Wetensch. 5: 198, 1830.

Peltigera hymenina Del. apud Duby, Botanic. Gallic. II. 597,

1830.

Peltigera polydactyla f. pellucida Dietrich, Lichenogr. Ger¬

man. 27, 1832-37.

Peltigera rufescens var. polydactyla Torss., Enumer. Lich.

et Byssac. Scand. 8, 1843.

Peltigera polydactyla var. vulgaris Korb., Syst. Lich. Ger¬

man. 61, 1855.

Peltigera canina var. coriacea f. polydactyla Krmphbr.,

Denkschrift Kgl. bayer. Bot. Ges. 4; 2 abth: 124, 1861.

Peltigera canina var. polydactyla Branth. et Rostr., Bot.

Tidsskrift 3: 175, 1869.

Peltigera magyarica Gyel., Magy. Bot. Lapok 28 : 61, 1929.

Thallus variable, pusilloid (single erect lobes) to large and

imbricate, the lobes up to 4 cm. broad and several cm. long;

upperside dark greenish-blue when moist, slaty or bluish or

greenish gray or brownish when dry, the margins ascending,

some forms crisped or proliferate; upperside smooth, shining;

underside with broad, flat, white to more usually brown or

black, reticulated veins with very small white interspaces; rhi-

zoids sparse, short and fasciculate or over 5 mm. long and

simple, dark brown to black; apothecia 2 to 5 mm. across on

erect narrow lobules at the ascendent tips of the lobes, the

sides usually reflexed, the disk reddish- or chestnut-brown to

black, the margin crenate; hypothecium brown; hymenium

hyaline; spores hyaline, 3-9 septate, acicular, slightly curved

48-71 X 3-4.5 n in Wisconsin material.

258 Wisconsin Academy of Sciences, Arts and Letters

On soil, moss, logs, rocks and bases of trees in moist woods.

4a. Peltigera polydactyla var. typica.

Ashland: Mellen, 1927, L. R. Wilson ; Douglas: Pattison

State Park, 1942; at mouth of Dutchman’s Creek, Superior,

1942; Blueberry, 1907, coll, probably J. J. Davis; Ranger Sta¬

tion, Brule River, 1943; Stone’s Bridge, Brule River, 1942;

Upper Brule River, 1942; Grant: Glen Haven, 1921, Chas. E.

Allen; Iowa: Hollandale, 1925, R . H. Denniston; Tower Hill

State Park, 1945; Iron: Lake Superior Shore near Montreal

River, 1896, L. S. Cheney; Juneau: Upham Woods, Wisconsin

Dells, 1944; Lincoln: Grandfather Bull Falls, 1893, L. S.

Cheney; Marathon: Granite Heights, 1894, L. S. Cheney;

Marinette: Dunbar, 1945; Vilas: Sayner, 1941.

The species is very variable in character and a number of

varieties and forms have been split from it. Those which have

been distinguished from Wisconsin are:

4b. Peltigera polydactyla f. microcarpa (Ach.) Merat, Nouv.

Flor. Lich. ed. 2, I: 199, 1821.

Peltidea polydactyla var. microcarpa Ach., Lich. Univ. 520,

1810.

Peltigera polydactyla var. hymenina f. microcarpa Flotow,

28 Jahresber. Schlesisch. vaterland. Kultur. 125, 1850.

Peltigera polydactyla var. microcarpa f. cephalodigera Miill.

Arg., Flora 72: 144, 1889.

Peltigera polydactyla var. nervosa Gyel., Osterr. Bot. Zeit.

77: 225, 1928.

Peltigera polydactyla var. subnervosa Gyel., Magy. Bot. La-

pok 28: 61, 1929.

Thallus and apothecia small, the former about one cm. or

less across the lobes; the apothecia about two mm. across.

Bayfield : on a boulder in woods, Pigeon Lake, Drummond,

1945.

4c. Peltigera polydactyla f. microphylla Anders. Lotos 76:

320, 1928.

Peltigera perfida Gyel. apud Anders., Die Strauch-und Laub-

flecht. Mitteleurop. 49, 1928.

Peltigera microphylla Gyel., Bryologist 34: 18 & 19, 1931.

Similar to P. polydactyla, but with tiny proliferations or

“isidia” on the margins and upper surface, especially where ob¬

viously injured.

Thomson— The Wisconsin Species of Peltigera 259

Douglas: north of Brule, 1943.

4d. Peltigera polydactyla f. lophyra (Ach.) NyL, Lich.

Scand. 90, 1861.

Peltidea horizontalis var. lophyra Ach., Lich. Univ. 516,

1810.

Peltidea canina var. glabra Ach., Synops. Lich. 239, 1814.

Peltidea rufescens var. collina Ach. Meth. 285, 1803.

Peltigera polydactyla f. collina NyL, Synop. Lich I. 327,

1860.

As redefined by Gy elnik (17) the description becomes: sim¬

ilar to the typical plant but with the margins dilacerate to di-

lacerate-crisped.

Iowa: hill near Tower Hill Park, 1939.

4e. Peltigera polydactyla var. dolichorrhiza Nyl., Synops.

Lich. I., 327, 1860.

Peltigera dolichorrhiza Nyl., Lich. Nov. Zeland. 43, 1888.

Similar to the species but the thallus tends to be thinner.

The rhizinae are more than 5 mm. long, dark brown or black

and simple. The underside is light brownish white to brown.

Douglas: upper Brule River bog, 1943; Rusk: no locality,

1921, R . H . Denniston.

4f. Peltigera polydactyla var. crassoides Gyel. Magy. Bot.

Lapok 29: 51, 1930.

Peltigera polydactyla var. hymenina Auct. (Vide Gyelnik

Lc.)

Thallus small, dark colored, thick, rigid instead of flexible,

underside with the veins confluent and almost no interspaces,

rhizinae short, fasciculate. Otherwise as in the species.

This variety resembles P. malacea very much but is smooth

and shining above, lacking the erect tomentum of that species.

Dane: West of Middleton, 1936, R. H. Denniston; La Crosse,

no date, L. H. Pammel (specimen in Herb. Univ. Mich.) ; VER¬

NON: Coon Valley, 1945.

5. Peltigera evansiana Gyel., Bryologist 34: 16, 1931.

Thallus small, lobes usually a cm. or less across; upper

side pale gray, brownish-gray or brown, lacking tomentum but

scabrid, not smooth or shining, with cylindrical, dark brown,

sometimes branched, club-shaped isidia scattered over the sur¬

face, the isidia not associated with cracks or breaks in the

260 Wisconsin Academy of Sciences , Arts and Letters

thallus; underside pale brownish white to brown, with a net¬

work of light brown to brown, raised, narrow veins and large

interspaces; the rhizinae simple, up to 4 mm. long. Sterile in

Wisconsin material as well as in the type.

Adams : Coldwater Canyon, north of Wisconsin Dells, 1945 ;

Bayfield: Pigeon Lake, Drummond, 1945; Douglas: Upper

Brule River above Stone's Bridge, 1942; Winneboujou, 1942;

Brule River Ranger Station, 1943; Pattison State Park, 1944;

Mouth of Nebagamon Creek, 1943; Forest: East of Wapigon

Lake, 1945; east of Crandon, 1945; Grant: Glen Haven, 1921,

Chas. E. Allen; Iowa: Hollandale, 1925, R. H. Denniston; Tow¬

er Hill State Park, 1945; Juneau: Upham Woods, Wisconsin

Dells, 1944; Lafayette: Darlington, 1946, P. Nelson; Marin¬

ette : Wausaukee, 1945 ; Dunbar, 1945 ; Sauk : Rocky Arbor State

Park, 1945; Devil's Lake, 1924, R. H. Denniston; Waukesha:

Martin’s Woods, Big Bend, 1945.

6. Peltigera canina (L.) Willd., Flora Berolinens., Prodrom.

347, 1787.

Lichen caninus L., Sp. PI. 1149, 1753.

Lichen caninus var. cinereus Weis., Flor. Goetting. 78, 1770.

Lichen terrestris Lam., Flore. Frang., I. 84, 1788.

Lichen venosus Gilib., Exercit. Phytol. 601, 1792 (non

Linn.)

Peltidea canina Ach., Method. Lich. 283, 1803.

Dermatodea canina St. Hil., Expos. Famil. Natur. 20, 1805.

Peltigera canina var. palmata Del. apud Duby. Bot. Gallic

II. 598, 1830.

Peltigera canina var. vulgaris Duby, Bot. Gallic II. 598,

1830.

Peltigera canina f. digitata Hazsl., Birod. Zuzmo-Flor. 56,

1884.

Peltigera canina var. membranacea f. palmata Oliv., Expos.

Lich. Quest. France I. 156, 1897.

Peltigera canina var. canina Boist., Nouv. Flore. Lich. 2. 80,

1903.

Peltigera Plittii var. macrolobata Gyel., Fedde Repert. 29:

9, 1931.

Peltigera Plittii var. sandwicensis Gyel., Fedde Repert. 29:

9, 1931.

Plate

V. — P. canina var. spuria f. sorediata, upper with the ‘ spuria ’ type

thallus, lower with ‘hazslinszkyi’ type thallus.

Plate VI. — 1. P. canina var. rufescens f. innovans; 2. P. evansiana. Scale

came as Plate 5.

Thomson — The Wisconsin Species of Peltigera 261

Peltigera Plittii f. suffusa Gyel., Lilloa 3 : 64, 1938.

Peltigera canina f. palmata A. Zahlbr., Cat. Lich. Univ. III.

464, 1925.

Thallus very variable; size up to 30 cm. or more across,

lobes up to 30 mm. across, slate gray, greenish brown or brown

above with an arachnoid or dense tomentum toward the mar¬

gins, dull or shining above toward the older thallus parts; in

one form with coarse gray soredia in rounded spots on the up¬

per surface of the thallus ; below with strong network of raised

white to brown veins, with the white or brown rhizinae, simple,

fasciculate, or fibrillose, sometimes forming a spongy nap be¬

low; apothecia vertical, the sides reflexed, the disk light brown

to reddish or chestnut-brown or black, the margin crenate;

hypothecium brown, hymenium hyaline, spores 3-9 septate,

acicular, hyaline to yellowish, 23-67 X 3-6.5 ^ in Wisconsin

material.

A number of species have been separated from Peltigera

canina by various authors, but I find myself unable to accord

most such specimens anything more than the rank of variety.

P. canina is exceedingly variable and in the field the segregated

'species' do not seem to hold good. For example, thalli observed

on rock outcrops in Northern Wisconsin have been indubitable

P . rufescens in the center and P. canina var. ulorrhiza or var.

albescens at the border. Regeneration of the thallus following

injury by snails, insects, or larger animals brings about the

formation of tiny platelets or “isidia” along the borders of the

injury and along cracks in the thallus, thus producing the 'spe¬

cies' P. praetextata and similar segregates. Scholander has al¬

ready shown (26) that P. leptoderma is but an etomentose

variant of P. erumpens. It does not seem to me that the pres¬

ence or absence of soredia in the case of P. erumpens is a valid

character for species differentiation. As C. F. E. Ericksen

points out (3) P. leptoderma , P. Hazslinskyi, and P. erumpens

are doubtfully separable from P. spuria. As far as I am able

to make out in field studies, P. Hazslinskyi is but the juvenile

stage of growth of P. spuria. The young pusilloid condition

grades imperceptably into the larger canina-like, or really

spuria-like, thalli of 'P. erumpens ' in the same colony. I also

can not see that these are more than sorediate forms of the

spuria type of thallus as gradations with all the appearance of

P. spuria and esorediate or with from one fleck of soredia to

262 Wisconsin Academy of Sciences, Arts and Letters

a heavy flecking with the spots of soredia exist in the same

colony. On rock outcrops in the cutover areas of Northern

Wisconsin it is common to find the P. rufescens type of thallus

breaking up into the P. spuria type toward the margins where

the lichen intermixes with mosses. It is this type of observa¬

tion which leads me to reduce P. rufescens and P. spuria to

the rank of varieties rather than retaining them as species.

It is obvious that a plastic species such as P. canina would be

excellent material for a study of transplants and development

of individual thalli. Upon such a study depends the final plac¬

ing of the nomenclatorial units.

6a. Peltigera canina var. albescens (Wahlbg.) Thomson,

Comb. Nov.

Peltidea leucorrhiza Flk. Deutsch. Lich. No. 153, 8 Lief. 10,

1821.

Peltidea canina var. albescens Wahlb. Flora Suec. II. 842,

1826.

Peltigera canina var. leucorrhiza Flotow. 28 Jahresber,

Schlesisch Gesellsch fur vaterland Kultur. 124, 1850.

Peltigera canina var. membranacea f. genuina Krmphbr.,

Denkschrift. Kgl. Bayer. Bot. Gesell. 4, 2 abth. 124, 1861.

Peltigera canina f. leucorrhiza Arn. Flora 67 : 233, 1884.

Peltigera canina var. membranacea f. leucorrhiza Oliv., Ex¬

pos. Lich. Guest. France I. 156, 1897.

This variety is the typical material of Peltigera canina with

white or brown veins and interspaces and light rhizoids below.

According to international rules, Wahlenberg’s earliest usage

of varietal rank must be taken up.

Douglas : Pattison State Park, 1942 ; 1935, R . H. Denniston;

Brule, 1944; N. P. Johnson Bridge North of Brule, 1944; Ly¬

man Lake, 1942, Olive S. Thomson ; east of Superior, 1942;

falls of Copper Creek, south of Superior, 1942 ; Forest : east of

Wapigon Lake, 1945; Grant: Close Creek, Glen Haven, 1921,

C. E. Allen; Glen Haven, 1921, C. E. Allen; Iowa: Hollandale,

1925, R. H. Denniston; Oneida: Monico, 1945; Portage: nine

miles north of Stevens Point, 1938; Sauk: Rocky Arbor State

Park, Wisconsin Dells, 1945; Vilas: Sayner, 1941; Allequash

Lake near Trout Lake, 1938; Lac Vieux Desert, 1893, L. S.

Cheney .

Thomson— The Wisconsin Species of Peltigera

263

6b. Peltigera canina var. ulorrhiza (Flk.) Schaer. Enumer.

Critic. Lich. Europ. 20, 1850.

Peltidea ulorrhiza Flk. Deutsch. Lich. no. 154, 11, 1821.

Peltigera canina var. coriacea f. campestris Krmphbr., Denk-

schrift. Kgl. Bayer. Bot. Gesellsch. 4, 2 abth. 125, 1861.

Peltigera canina f. ulorrhiza Oliv., Memoir. Soc. Nation.

Scienc. Natur. Cherbourg. 36: 218, 1907.

The underside of this variety has the veins and rhizoids

dark brown ; the interspaces creamy or yellowish to pale brown.

It seems to represent a transition between typical canina and

var. rufescens.

Dane: Lake Wingra, 1893, Heald and Buell ; Door: Penin¬

sula State Park, 1928, R. H. Denniston ; Douglas : White cedar

swamp at head of Brule River, 1944, J. W. Thomson, Jr. and

R. /. Evans; falls of Copper Creek, south of Superior, 1942;

east of Superior, 1942; Fond du Lac: Wolf Lake, Marshfield,

1938, L. H . Shinners; Lincoln: Gleason, 1945; Marathon:

Wausau, 1938; Sauk: Rocky Arbor State Park, Wisconsin

Dells, 1945; Devil’s Lake, 1935; 1945; St. Croix: Hudson, 1945.

6c. Peltigera canina var. rufescens (Weis.) Mudd., Manual

Brit. Lich. 82, 1861.

Lichen caninus var. rufescens Weis., Plant. Cryptog. Flor.

Goettingens. 79, 1770.

Lichen rufescens Neck., Method. Muscor. 79, 1771.

Lichen rufus GmeJin., System. Natur. II, Pars. II. 1373,

1791.

Peltigera rufescens Humb., Flor. Friburg. Specim. 2, 1793.

Peltidea rufescens Ach., Method. Lich. 285, 1803.

Peltidea canina var. crispa Ach., Lich. Univ. 519, 1810.

Peltidea canina var. inflexa Ach., Lich. Univ. 518, 1810.

Peltidea malacea f. inflexa Ach., Synops. Lich. 240, 1814.

Peltidea canina var. rufescens Hook., Flora Scotica II. 60,

1821.

Peltigera canina var. crispa Merat., Nouv. Flor. Envir. Par¬

is ed. 2, I. 199, 1821.

Peltigera canina var. inflexa Duby. Bot. Gallic. II. 598, 1830.

Peltigera canina var. crispata Kickx., Flora Cryptog. Lou¬

vain. 91, 1835.

264 Wisconsin Academy of Sciences , Arts and Letters

Peltigera rufescens var. pachyphylla f. crispa Flotow, 28

Jahresberich. Schlesisch. Gesellsch. Vaterland Kultur.

124, 1850.

Peltigera rufescens f. incusa Korb., Syst. Lich. Germ. 59,

1855.

Peltigera canina var. coriacea f. genuina Krmphbr., Denk-

schrift. Kgl. Bayer. Bot. Gesellsch. 4, 2 abth. 124, 1861.

Peltigera canina var. coriacea f. incusa Krmphbr., Denk-

schrift. Kgl. Bayer. Bot. Gesellsch. 4, 2 abth. 125, 1861.

Peltigera canina var. membranacea f. rufa Krmphbr., Denk-

schrift. Kgl. Bayer. Bot. Gesellsch. 4, 2 abth. 124, 1861.

Peltigera malacea var. crispa Th. Fr., Nova Acta Reg. Soc.

Sclent. Upsal. Ser. 3. 3: 144, 1861.

Peltigera rufescens var. inflexa Kicks., Flore Cryptog. Flan-

dres I. 215, 1867.

Peltigera canina var. ulorrhiza f. crispa Hepp, Flecht.

Europ. No. 850, 1867.

Peltigera canina f. rufa. Rabh., Cryptog. Flora von Sach¬

sen, 2 abth. 309, 1870.

Peltigera canina f. crispata Koltz, Recueil Memoir, et Trav.

Soc. Botan. Luxemburg, 13: 172, 1897.

Peltigera canina f. rufa Koltz, Recueil Memoir et Trav. Soc.

Bot. Luxemburg 13: 172, 1897.

Peltigera canina subsp. rufescens var. inflexa Boist., Nouv.

Flore. Lich. part 2: 81, 1903.

Peltigera rufescens f. inflexa Zahlbr., Cat. Lich. Univ. Ill,

487, 1925.

Peltigera aloisii Gyel., Oestr. Bot. Zeitschr. 77 : 222, 1928.

Peltigera suomensis Gyel., Magy. Bot. Lapok, 28: 60, 1929.

Peltigera rufescens f. albidula Gyel., Magy. Bot. Lapok 28:

61, 1929.

Peltigera canina var. suomensis Ras., Annal. Mo. Bot. Gard.

20: 16, 1933.

Peltigera plittii f. ornata Gyel., Lilloa 3: 64, 1938.

Peltigera plittii f. suffusa Gyel., Lilloa 3 : 64, 1938.

Peltigera plittii f . subsuffusa Gyel., Lilloa 3 : 64, 1938.

As Gyelnik points out in Bot. Koz. 24, 1927 the differences

between typical P. canina and var. rufescens are slight. The

latter is smaller, more brittle, generally reddish, with cell walls

of the upper layer slightly thicker (2-3 instead of 1-2 /* thick)

Thomson — The Wisconsin Species of Peltigera 265

and grows in drier, more sunny habitats than the typical spe¬

cies, vars. albescens and ulorrhiza. The difference in the habi¬

tats suggests that var. rufescens is but an ecological variant of

P . canina. Not only in the identification of specimens but also

in the literature there has been considerable confusion of the

two “species.” In any large number of collections, all inter¬

grades between the two may be found and it does not seem to

me that they should be maintained as separate species. On

very dry, sunny, rock outcrops the plant is typically crisped

and narrow lobed with the lobes inrolled at the margins; but

as the same thallus may have broader, flatter lobes toward

the circumference where more moisture may be available

trickling down the rock, I have not recognized f. or var. in-

flexa. Such crisped, narrow-lobed plants with a very thick,

instead of an arachnoid, tomentum on the upper surface are

called P. plittii f. ornata by Gyelnik. The var. rufescens is the

most abundant variety of P. canina in this state as well as in

North America.

Bayfield: Brule Barrens, T46N, R.9W., Sec. 30, 1943;

Columbia : 5 miles southwest of Portage, 1945 ; Dane : Beeches’

Prairie near Sauk City, 1945, M. B. Berseng and Joan Wright;

Door: Washington Island, 1931, J. J. Davis; Douglas: Amnicon

Falls, 1942; Brule, 1943; South of Superior, 1942; 1944; Dewey,

1942, Sugar Camp Hill, north of Brule, 1943; Fond du Lac:

Oakfield, 1936; Forest: Crandon, 1945; Iowa: Blue Mounds,

1921, R. H. Denniston; Hollandale, 1925, R. H. Denniston;

Juneau: Germantown, 1894, L. S. Cheney; Marathon: Rib Hill,

1945; Marinette: Dunbar, 1945; Wausaukee, 1945; Oneida:

Monico, 1945; 2 miles west of Monico, 1945; Portage: Stevens

Point, 1894, L. S. Cheney; east of Stevens Point, 1938; Racine:

Wind Lake, 1945; St. Croix: By Lake St. Croix, Hudson, 1945

(Specimen is a transition between this and var. spuria) ; Sauk:

Spring Green, 1939; Parfrey’s Glen, 1924, R. H. Denniston;

Devil’s Lake, 1907, 1916, J. R. Heddle; Devil’s Lake, 1922, R. H.

Denniston; Baxter’s Hollow, Baraboo Range, 1939 ; Vilas : Con¬

over, 1893, L. S. Cheney; Vernon : Coon Valley, 1945.

6d. Peltigera canina var. rufescens f. innovans (Korb)

Thomson, Comb. Nov.

Peltidea ulorrhiza var. praetextata Flk. apud Sommerf..

Suppl. Flor. Lappon. 123, 1826.

266 Wisconsin Academy of Sciences , Arts and Letters

Peltigera canina var. membranacea f. sorediata Schaer.,

Enumer. Critic. Lich. Europ. 20, 1850. (non Peltigera

canina var. spuria f. sorediata 1. c.)

Peltigera rufescens f. innovans Korb., Syst. Lich. Germ. 60,

1855.

Peltigera rufescens var. praetextata Nyl., Synops. Lich. I.

324, 1860.

Peltigera canina var. coriacea f. sorediata Krmphbr., Denk-

schrift. Kgl. Bayer. Bot. Gesellsch. 4, 2 abth. 125, 1861.

Peltigera rufescens var. innovans Kickx, Flore. Crypt. Flan-

dres II. 215, 1867.

Peltigera canina var. crispa Flagey, Memoir. Soc. d’Emulat.

Doubs. 420, 1882. (Non Ach.)

Peltigera canina var. limbata Mudd, Manual Brit. Lich. 83,

1861.

Peltigera canina f. undulata Arn., Flora 67 : 234, 1884.

Peltigera canina f. vivipara Hazsl., Magy. Birod. Zuzmo-

Flor. 56, 1884.

Peltigera polydactyla var. undulata Hue, Bull. Soc. Bot.

France 41: 177, 1894.

Peltigera praetextata Wainio, Termeszetr. Fiizetek, 22: 306,

1899.

Peltigera canina var. praetextata Hue. Nouv. Archiv. du

Museum, Ser. 4, II. 95, 1900.

Peltigera canina subsp. rufescens var. praetextata Boist.,

Nouv. Flore. Lich. part 2. 81, 1903.

Peltigera praetextata var. subglabra Gyel. apud Erichs, in.

Verhandl. Bot. Verein Prov. Brand. 70: 221, 1928.

Peltigera praetextata f. lapponica Gyel., Magy. Bot. Lapok

28: 61, 1929.

Peltigera praetextata var. ilseana K. S. K., Fedde Repert.

28: 203, 1930.

Peltigera praetextata f. subglabra Gyel., Lilloa 3: 64, 1938.

Peltigera praetextata f. vivipara Gyel., Lilloa 3: 64, 1938.

This form resembles P. canina var. rufescens but has small

flat platelets or “isidia” growing from the edges and upperside

especially along cracks and injured spots. It represents a regen¬

eration phenomenon rather than a distinct species. Specimens

have been described with cylindrical, probably true isidia, but

I have not seen these. In accordance with the international

Thomson — The Wisconsin Species of Peltigera 267

rules, the earliest acceptable name in the formal rank is taken

up.

Columbia: Black Hawk Lookout opposite Prairie Du Sac,

1941, Olive Thomson; 5 miles southwest of Portage, 1945; JU¬

NEAU: Upham Woods, Wisconsin Dells, 1944; Sauk: Devil's

Lake, 1922, R . H. Denniston; Vernon : southeast of Coon Valley,

1946; Walworth: Whitewater, 1940; Waukesha: Martin's

Woods, Big Bend, 1945.

6e. Peltigera canina var. spuria (Ach.) Schaer., Lich. Helvet.

Spicil. Sect. 6. 265, 1833.

Lichen spurius Ach., Lichenogr. Suec. Prodrom. 159, 1798.

Peltidea spuria Ach., Method. Lich. 283, 1803.

Peltigera spuria Lam. et D. C., Flore Frang, ed. 3, II. 406,

1805.

Peltidea canina var. spuria Ach., Lich. Univers. 518, 1810.

Peltidea polydactyla var. spuria Schlecht., Flora Berolinens,

II. 84, 1824.

Peltigera polydactyla var. spuria Mann, Lich. in Bohemia

Observ. Dispos. 71, 1825.

Peltigera venosa var. digitata March apud. Hall, Vrolik et

Mulder, Bijdrag. t.d. Natuurk. Wetensch. 5: 198, 1830.

Peltigera canina var. pusilla E. Fries., Lich. Europ. Ref. 45,

1831.

Peltigera rufescens var. pachyphylla f. spuria Flotow, 28

Jahresber. Schlesisch. Gesellsch. Vaterland Kultur. 124,

1850.

Peltigera rufescens f. spuria Korb., Syst. Lich. Germ. 59,

1855.

Peltigera pusilla Korb., Syst. Lich. Germ. 59, 1855.

Peltigera rufescens Hook., Handbook New Zealand Flora.

566, 1867. (non alior)

Peltigera spuriella Wainio, Acta Societatis pro Fauna et

Flora Fennica 7 : 180, 1890.

Peltigera behringiana Gyel., Oester. Bot. Zeitschr. 77 : 222,

1928.

The thallus of this variety is quite similar to that of var.

rufescens with which it intergrades but the lobes are short,

more or less erect, separated from each other and usually are

gray or bluish-green rather than brownish in color. The apo-

268 Wisconsin Academy of Sciences , Arts and Letters

thecia are smaller, darker, and more strongly reflexed on the

sides than in var. rufescens.

Adams: Plainfield, 1935; Dans: Northwest of Evansville,

1944, Geo . S. Bryan; Douglas: Pattison State Park, 1942;

Winneboujou, Brule River, 1942; Langlade: Parrish, 1945;

Oneida: 3 miles west of Monico, 1945; Doherty Lake, 1893,

L. S. Cheney; Vilas : Sayner, 1941 ; south of Sayner, 1941;

Lost Creek, Sayner, 1938; Star Lake, 1946.

6f. Peltigera canina var. spuria f. sorediata Schaer. Enu-

mer. Critic. Lich. Europ. 20, 1850.

Peltidea erumpens Tayl. In Hook. London Jour, of Bot.,

6: 184, 1847.

Peltigera canina var. sorediifera Schaer. Lich. Helvet. Spicil.

Sect. 6, 265, 1833.

Peltigera canina f. sorediifera Zahlbr. Cat. Lich. Univ. III.

464, 1925.

Peltigera canina var. soreumatica Fiotow, 28 Jahresber.

Schlesisch. Gesellsch. fur. Vaterland Kultur. 124, 1850.

Peltigera leptoderma Nyl. Synops. Lich. I. 325, 1860.

Peltigera canina var. notata Th. Fr., Kgl. Svensk. Vetensk.

Akad. Handl. 7: 15, 1867.

Peltigera canina var. extenuata Nyl. apud. Wainio, Meddel.

Soc. Fauna et Flora Fennic. 5 : 49, 1873.

Peltigera ulcerata Miill. Arg., Flora 63 : 261, 1880.

Peltigera extenuata Wainio, Meddel. Soc. Fauna et Flora

Fennic. 6: 129, 1881.

Peltigera rufescens f. sorediata Oliv., Flore Lich. Orne, I.

92, 1882,

Peltigera rufescens var. vulnerata Mull. Arg., Flora 65 : 305,

1882.

Peltigera pusilla var. vulnerata Mull. Arg., Flora 69: 133,

1888.

Peltigera erumpens Wainio, Etud. Lich. Bresil. 182, 1890.

Peltigera spuria var. erumpens Harm., Bull. Soc. Scienc.

Nancy, Ser. 2, 31 : 248, 1897.

Peltigera canina var. erumpens Hue, Nouv. Archiv. du Mu¬

seum, Ser. 4, 2 : 96, 1900.

Peltigera canina f. cyatheum Norm, apud. Lynge, Bergens

Museums Aarborg 9, 109, 1910,

Thomson — The Wisconsin Species of Peltigera 269

Peltigera sorediata Fink apud. Corringt., Ohio State U.

Bull. 25: 356, 1921.

Peltigera Hazslinszkyi Gyel. apud. Anders. Die Strauch und

Laub Flecht. Mitteleurop. 44, 1928.

Peltigera erumpens f. glabrescens Gyel. apud. Erichs., Ver-

handl. Bot. Verein. Prov. Brandenbourg 70: 219, 1928.

Peltigera leptoderma var. brasiliensis Gyel., Oester. Bot.

Zeitschs. 77: 224, 1928.

Peltigera erumpens f. densa Gyel., Magy. Bot. Lapok, 28:

61, 1929.

Peltigera erumpens f . scabrida Gyel., Magy. Bot. Lapok, 28 :

61, 1929.

Peltigerq, vainioi Gyel., Magy. Bot. Lapok, 28: 61, 1929.

Peltigera erumpens f. leptoderma Scholander, Nyt. Magazin

for. Naturvid. 73: 52, 1933.

This form is distinguished by the rounded spots bearing

coarse gray soredia on the upper surface of the thallus. The

amount of tomentum on the upper surface of the thallus varies

from lacking or almost lacking to a rather dense layer at the

margins. The upper surface may be smooth to whitish pruinose

scabrid in spots. The thalli vary from very tiny, cochleate, sep¬

arate lobes to larger spuria-like lobes. The rhizinae on the un¬

derside vary from simple or pappose to a fairly dense mat of

fibrillose rhizinae. Numerous forms, varieties, and species have

been made of these variations but the intergradation observed

in the field is such as to lead one to believe that they are but

normal variation of a very plastic plant and not entitled to

nomenclatorial recognition. The smaller forms are more com¬

mon on newly invaded soil such as ditchbanks and abandoned

sandy fields; the larger plants tending toward the spuria and

rufescens types are on less disturbed habitats.

Adams: 10 miles east of Friendship, 1935; Bayfield: Pi¬

geon Lake, Drummond, 1945; Columbia: Black Hawk Lookout

opposite Prairie du Sac, 1935; Dane: Black Earth, 1921, E. J.

Kraus; Picture Rock, 1938, L. H. Skinners; Douglas: Upper

Brule River Valley, 1943, 1946; east of Superior, 1942; Dewey,

1942; Pattison State Park, 1944; J. W. Thomson , Jr. and R. I.

Evans; Superior, 1942; south of Superior, 1942; Dunn: Eau

Galle, 1927, R. H. Denniston; Jackson : 8 miles south of Black

River Falls, 1945; Marinette: Dunbar, 1945 (with parasites) ;

270 Wisconsin Academy of Sciences, Arts and Letters

Wausaukee, 1945; Monroe: 9 miles west of Tomah, 1935;

Oneida: 2 miles west of Monico, 1945; Sauk: Ableman, 1940;

Washburn: Sarona, 1944, R. I. Evans; Vilas: Conover, 1893,

L. S. Cheney; Star Lake, 1946.

6g. Peltigera canina var. spongiosa Tuck., Genera. Lich. 38,

1872.

Peltigera canina f. spongiosa Tuck., Synops. North. Amer.

Lich. I, 109, 1882.

Peltigera canina subvar. spongiosa Boist., Nouv. Flore Lich.,

part 2, 80, 1903.

Peltigera canina subsp. spongiosa Fink., Contrib. U. S. Nat*l.

Herb. 14: 163, 1910.

Similar to var. albescens with light colored veins and under¬

side, but the rhizinae on the underside are strongly fibrillose

and form a dense mat.

Douglas: Falls of Copper Creek, south of Superior, 1942;

Little Joe Falls, Brule River, 1943; Vernon: Coon Valley, 1945,

Bob Ellarson, this specimen approaches this variety but the mat

of rhizinae is not very heavy; Vilas: Found Lake near Sayner,

1941.

References

1. Anders, Jos. 1928. Die Strauch-und Laub-flechten Mitteleuropas. 40-

49.

2. Bitter, G. 1904. Peltigera-studien. Bericht. Deutsch. Botan. Gesellsch.

22 : 242-254.

- 1909. ibid. 186-195.

3. Erichsen, C. F. E. 1928. Die Flechten des Moranengebiets von Ost-

schleswig. Verhandl. Bot. Verein. Prov. Brandenburg. 70 : 128-223.

4. Fink, Bruce. 1899. Notes on lichen distribution in the upper Missis¬

sippi Valley, Mem. Torrey Bot. Club. 6: 285-307.

5. — - 1935. The Lichen Flora of the United States. U. Mich. Press.

6. Gyelnik, V. 1934. Additamenta lichenologica 1, Hedwigia 7 : 219-222.

7. — - 1932. Clavis et enumeratio specierum generis Peltigerae. Revue

Bryol. et Lichenol. 5 : 61-73.

8. • — - 1930. Eine neue Peltigera aus Finnland. Magy. Bot. Lapok

29: 34.

9. - 1938. Fragmenta Lichenologica I, Lilloa 3: 49-80.

10. - 1930. Lichenes Nonnulli Novi Criticique. Nyt. Magazin. for

Naturv. 68: 269-270.

- 1929. Lichenologiai Kozlemenyek 4-7. Magy. Bot. Lapok 27 :

91-93. f

11.

Thomson — The Wisconsin Species of Peltigera

271

12. - 1929. Lichenologiai Kozlemenyek 8-19 (nos. 8-10, 14-16). Magy.

Bot. Lapok. 28: 57-61.

13. - 1930. Lichenologiai Kozlemenyek 20-45 (no. 20.) Magy. Bot.

Lapok 29 : 23-24.

14. — - - 1927. Nehany Peltigera-adat Japanbol. Magy. Bot. Lapok 25:

252-254.

15. - 1928. Peltigera-Daten, Hedwigia 68: 1-4.

16. — - 1927. Peltigera-tanulmanyok. Botan. Kozlemenyek 24 : 122-140.

17. - — 1930. Revisio Peltigerarum Herbarii Achariam. Magy. Bot.

Lapok 29 : 49-58.

18. - 1925. Neber eine neue Flechte nebst kritische Bemerkungen

uber Peltigera aphthosa. Magy. Bot. Lapok.

19. Lapham, I. A. 1858-1859. Additions to the flora of Wisconsin-Lichenes.

Trans. Wis. State Agric. Soc. 5 : 424.

20. Lynge, B. and P. F. Scholander. 1932. Lichens from Northeast Green¬

land. Skrifter om Svalbard og. Ishavet No. 41.

21. Migula, W. 1926. Kryptogamen Flora von Deutschland, Osterreich und

der Schweiz. 378-385.

22. Moreau, F. et M. Recherches Sur les Lichens de la famille des Pelti-

geracees. Annal. Scienc. Nat. Bot. Ser. 10, 1 : 29-138.

23. Nilsson, G. 1932. Zur Flechten flora von Angermanland. Archiv. for

Botanik 24A, No. 3 : 1-122.

24. Nylander, Wm. 1858-60. Synopsis Methodica Lichenum I : 1-430.

25. Rasanen, Veli J. P. B. 1933. Contribution to the lichen flora of North

America. Ann. Mo. Bot. Gard. 20 : 7-21.

26. Scholander, P. F. 1933. Notes on Peltigera erumpens (Tayl.) Vain.

s.l. Nyt. Magazin for Naturvidenskaberne. 73 : 21-54.

27. Smith, A. L. 1918. A monograph of the British Lichens (2d ed.) I :

90-100.

28. Strato, Clemens. 1922. Uber Wachstum und Regeneration des Thallus

von Peltigera canina. Hedwigia 63 : 11-42.

29. Zahlbruckner, A. 1925. Catalogus Lichenum Universalis. Ill : 443-

494.

1932. ibid VIII : 317-329.

HOST-PARASITE RELATIONSHIPS AND GEOGRAPHICAL

DISTRIBUTION OF THE PHYSALOPTERINAE

(NEMATODA)

Banner Bill Morgan

Department of Veterinary Science, University of Wisconsin

INTRODUCTION

The host-parasite relationships of the subfamily Physalop-

terinae, a group of nematode worms varying in size from *4 to

4 inches in length which lives in the stomach of many animals,

are not very well known. This is due in part because the

Physalopteridae present more taxonomic difficulty than other

spirurids since they tend toward polydelphy.

Positive identification of species can be made only on char¬

acters possessed by both male and female specimens. The

characters of value for species determination in the Physalop-

terinae are as follows, in the order of their importance : (1)

dentition (generic value) , (2) number of uteri, (3) mode of

origin of uteri, (4) number of male ventral papillae, (5) ar¬

rangement of male ventral papillae, (6) shape of spicules , (7)

length of spicules, and (8) position of vulva, (figs. 1-16)

Certain characters cannot be used for specific determination

because of the wide variation within species : (1) posterior

sheath, (2) size and height of teeth, (3) length of esophagus,

(4) position of excretory pore and cervical papillae, (5) size

of eggs, (6) shape of bursa, (7) bursal markings, and (8)

shape of seminal receptacle.

Ortlepp (1922, 1937) divided the genus Physaloptera into

four groups on the basis of uterine numbers: Didelphys (2),

Tridelphys (3), Tetradelphys (4) and Polydelphys (more than

4) . Also the mode of origin of the uteri were taken into con¬

sideration. At the present time the origin of the uteri has

been divided into various groups: 2 A (2 uteri with common

278

274 Wisconsin Academy of Sciences , Arts and Letters

CHARACTERS USED FOR PHYSALOPTERINAE CLASSIFICATION

MALE BURSA— VENTRAL VIEW

M organ — Phy Salop te ra Host-Parasite Relationships 275

Explanation of Plate

All figures diagrammatic

Fig. 1. Anterior end, enface view of Skrjabinoptera.

Fig. 2. Anterior end, enface view of Physaloptera.

Fig. 3. Anterior end, enface view of Abbreviata.

Fig. 4. Anterior end, lateral view of Skrjabinoptera.

Fig. 5. Anterior end, lateral view' of Physaloptera.

Fig. 6. Anterior end, lateral view of Abbreviata.

Fig. 7. 2A type uteri.

Fig. 8. 2B type uteri.

Fig. 9. 2C type uteri.

Fig. 10. 3 type uteri.

Fig. 11. 4D type uteri.

Fig. 12. 4E type uteri.

Fig. 13. 4F type uteri.

Fig. 14. 5-1 5 G type uteri.

Fig. 15. 7-15H type uteri.

Fig. 16. Male bursa, ventral view of typical Physalopterinae.

276 Wisconsin Academy of Sciences , Arts and Letters

trunk), 2B (2 uteri without common trunk), 2C (2 uteri with¬

out common trunk, branches from the lateral sides), 3 (3 uteri

without common trunk), 4D (4 uteri with common trunk), 4E

(4 uteri with bifurcation of common trunk), 4F (4 uteri with¬

out common trunk), 5-15G (5-15 uteri with common trunk),

and 7-15H (7-15 uteri without common trunk). (Figs. 7-15)

When Schulz (1927) divided the genus Physaloptera into

several genera according to pseudolabial dentition the family

Physalopteridae was broken into two subfamilies, namely,

Physalopterinae and Proleptinae. The latter contains five gen¬

era: Proleptus, Thubunaea, Heliconema, Ortleppina, and Physa -

lopteroides. Morgan (1943) placed the genus Physalopteriodes

Wu and Liu (1940) in the subfamily Physalopterinae, but fur¬

ther evidence showed closer relationship of Physalopteroides to

Thubunaea, and consequently has been placed in the subfamily

Proleptinae. The subfamily Physalopterinae has since been

grouped into four genera by Schulz (1927) and Baylis (1934).

The genera include Physaloptera Rudolphi, 1819; Abbreviata

(Travassos, 1920) Schulz, 1927 ; Skrjabinoptera Schulz, 1927 ;

and Pseudophysaloptera Baylis, 1934.

LIFE CYCLE

The life cycles of the Physalopterinae are still unknown.

On the basis of findings in allied genera of the spirurids, this

group of nematodes will probably require certain arthropods

as intermediate hosts. Alicata (1937) found that the embryo-

nated eggs of Physaloptera turgida after ingestion by the cock¬

roach ( Blatella germanica) developed to the 3rd stage larvae.

Hobmaier (1941) reported similar findings for P. maxillaris.

Cram (1931) and Boughton (1937) reported the finding of

immature Physaloptera encysted in the breast muscles of bob-

white quail ( Colinus v. virginiania) and the ruffed grouse

( Bonasa umbellus) . As soon as some of the life cycles of the

more common species of Physaloptera have been determined

the host-parasite relationships of the group will become more

easily understood. This would enable one to conduct animal

experiments on host-specificity.

The knowledge of host specificity and speciation is yet too

meager for final analysis and no doubt many synonyms exist.

Considerable more experimental evidence is needed to justify

the naming of new species created entirely on host occurrence.

Physaloptera

Morgan— Physaloptera Host-Parasite Relationships

UJ Ul

< <

a o

a d

<n < ~

3 H O

J o m

o flc

o o

CO

LU

5

<

_J

<

2

<

2

3 3

i 5

o >•

S Sfc

UI

1 2

g E

< -j

Z <

K H

IU

" UJ

1 § I

OC

EC

Ul

>

2 uj

§ s

Sis

Q.

277

HYAENIDAE

FELIDAE

GANIDAE

Chart 1.— Host spread of the Physaloptera arranged by Classes, Orders, and Families.

278 Wisconsin Academy of Sciences , Arts and Letters

i -

LACERTILIA

SGINCIDAE

IGUANIDAE

LACERTiDAE

TEIIDAE

Physaloptera

2-A UTERI

— REPTILIA —

OPHIDiA

COLUBRIDAE

CROTALIDAE

VIPER IOAE

FALCONIFORMES

ACCIPITRIDAE

FALCONIDAE

EDENTATA RODENTIA

MYRMECOPHAGIDAE CRICETIDAE

BR ADYPODIDAE MURIDAE

MARSUPIALIA

OIDELPHIDAE

RODENTIA

CRICETIDAE

SCIURIDAE

MURIDAE

FALCONIFORMES

ACCIPITRIDAE

FALCONIDAE

AVES

MAMMALIA

i

CARNIVORA

PROCYONIDAE

MUSTELIDAE

FELIDAE

2-B UTERI

REPTILIA

OPHIDIA

COLUBRIDAE

MAMMALIA

2-C UTERI

AVES

i

— - |

LORICATA

ALLIGATORIDAE

CORACIIFORMES

BUCEROTIDAE

INSECTIVORA

TALPIDAE

CHIROPTERA

RHINOLOPHIDAE

- , - -

CARNIVORA

PROCYONIDAE

MUSTELIDAE

FELIDAE

CANIDAE

INSECTIVORA

ERINACEIDAE

CUCULIFORMES

CUCULIDAE

MAMMALIA

CARNIVORA

VIVERRIDAE

HYAENIDAE

CANIDAE

FELIDAE

GALL IFOR MES

PHASIANIDAE

3 UTERI — 4-D UTERI

MAMMALIA MAMMALIA

PRIMATA EDENTATA

GEfilDAE MYRMECOPHAGIDAE

7-15-H UTERI

- MAMMALIA -

4-F UTERI

MAMMALIA

PRIMATA

LASIOPYGIDAE

I

MARSUPIALIA

DIDELPHIDA^

RODENTIA

OASYPROCTIDAE

5-15-G UTERI

MAMMALIA

RODENTIA

CRICETIDAE

PRIMATA

CALLITRICHIDAE

LASIOPYGIDAE

CEBIOAE

Chart 2. — Different types of uterine origin in the Physaloptera arranged

by Classes, Orders, and Families of the hosts.

Morgan— Physaloptera Host-Parasite Relationships 279

Abbreviate!

AMPHIBIA

I

AMURA.

i

RANIDAE

BUFONIDAE

LAGERT1LIA

GHAIAAELEONTtOAE

GECKONIOAE

VARANIDAE

SGINCIDAE

• AGAMIOAE

ANGUIOAE

LAGERTIDAE

REPTILIA •

I

OPHSDIA

COLUBRIOAE

VIPERIDAE

ELAPIDAE

BOIDAE

CHELONIA

TESTUDINiDAE

ARTIODACTYLA

SUIDAE

MAMMALIA

RODENTIA

©GTODENTIDAE

SGIURIDAE

MURfDAE

CARNIVORA

FELIDAE

PRSMATA

HOMINIDAE

LASIOPYGIDAE

CALLITRIGHIDAE

Chart 3.— Host spread of the Abbreviates arranged by Classes, Orders,

and Families.

280 Wisconsin Academy of Sciences, Arts and Letters

Abbreviate

2 -A UTERI

REPTILIA

LACERTILIA

VA R AN I DAE

AGAMIDAE

I -

LACERTILIA

CHAMAELEONTIDAE

GECKONIDAE

LACERTIDAE

VARANIDAE

SCINCIDAE

AGAMIDAE

ANGUIDAE

RODENTIA

OCTODONTIDAE

SCIURIDAE

MURIDAE

4-D UTERI

- REPTILIA - - -n

' I

OPHIDIA CHELONIA

COLUBRIDAE TESTUDINIDAE

ELAPIDAE

BOIDAE

MAMMALIA - - ,

I I

CARNIVORA PRIMATA

FELIDAE LASIOPYGIDAE

HOMINIDAE

4“E UTERI

I REPTILIA —

LACERTILIA

CHAMAELEONTIDAE

VARANIDAE

AGAMIDAE

5-15-6 UTERI

~1

OPHIDIA

COLUBRIDAE

VIPERIDAE

MAMMALIA

Chart 4. — Different types of uterine origin in the Abbreviate, arranged by

Classes, Orders, and Families of the hosts.

Morgan — Physaloptera Host-Parasite Relationships 281

RELATIONSHIPS OF THE PHYSALOPTERA

Physaloptera is now characterized by 1 externolateral tooth

and 1 internal group of 3 teeth, and usually no denticles on

the margins of the pseudolabia. The genus contains approxi¬

mately 42 species. They are represented in the following hosts :

Amphibia ; Anura (Ranidae) [1 species]; Reptilia; Lacer-

tilia (Scincidae, Teiidae, Amphisbaenidae, Iguanidae), Ophi-

dia (Colubridae, Crotalidae, Boidae), Loricata (Alligatoridae)

[4 species]; Aves: Falconiformes (Accipitridae, Falconidae),

Cuculiformes (Cuculidae), Galliformes (Phasianidae, Tetra-

onidae), Strigiformes (Strigidae), Coraciiformes (Buceroti-

dae) , Ciconiiformes (Threskiornithoidae) , Sphenisciformes

(Spheniscidae) [12 species]; Mammalia: Carnivora (Canidae,

Felidae, Mustelidae, Procyonidae, Hyaenidae, Viverridae) [9

species] ; Rodentia (Muridae, Sciuridae, Cricetidae, Dasyproc-

tidae) [5 species] ; Chiroptera (Rhinolophidae) [1 species] ;

Edentata (Myrmecophagidae, Bradypodidae) [2 species] ; In-

sectivora (Erinaceidae, Talpidae) [4 species] ; Marsupialia

(Bidelphiidae) [2 species] ; Primata (Cebidae, Lasiopygidae,

Callitrichidae) [3 species].

The genus Physaloptera is widely distributed throughout

the animal kingdom embracing 42 species among 4 classes,

15 orders representing 33 families. The number of hosts totals

well over 250 different animals. Chart 1 shows in tabular form

the spread of hosts as arranged by orders. The Amphibia is

represented by 1 family, Reptilia by 8, Aves by 5, and the

Mammalia by 19. The order Carnivora has the largest number

of families, six being represented.

Of the 42 known valid species several are somewhat re¬

stricted in geographical distribution. This may be due to the

range of the various susceptible hosts; P. abjecta, P. torquata,

and P. limbata has been reported only from the United States ;

P. amphibia only from the Philippine Islands (Luzon) ; P. bra-

chycera from Angola ; P. crosi, P. galinieri, P. losseni, P. rapa-

cis, P. aduensis, P. canis, P. immerpani, P. bedfordi from

Africa; P. hieracideae from Australia; P. mexicana from Mex¬

ico; P. reevisi from China; P. mirandai, P. magnipapilla, P.

papillotruncata, P. torresi , P. anomala and P. bonnei , from

South America ; P. masoodi and P. tumefaciens from India. The

most cosmopolitan species would include P. praeputialis, P.

282 Wisconsin Academy of Sciences , Arts and Letters

maxillaris , P. turgida, P. getula , P. ram, P. a£a£a and P.

cauda.

From the examination of host-parasite lists, several species

of Physaloptera may occur in a similar host. For example, P.

praeputialis may occur in the cat, mountain lion, bobcat, dog,

or gray fox. Two different species of Physaloptera have never

been reported occurring in the same animal at the same time ; for

example, P. praeputialis and P. rara from the same dog. Super

parasitism is not known to occur in the Physalopterinae.

The genus Physaloptera is the only member of the subfamily

Physalopterinae reported from birds. So far as is known, species

occurring in birds have not been found in any other class of

vertebrates. It is probably safe to say that species of Physalop¬

tera found in birds are host specific for Aves. This is, of course,

discounting aberrant or pseudoparasites.

This group of nematode worms are found mainly in the phar¬

ynx, crop, esophagus, proventriculus, ventriculus and occasionally

in the intestine of various bird hosts. Rarely, immature forms

are found encysted in breast muscles of birds. There are two

cases on record of Physaloptera found in the orbital cavity of

birds; P. acuticauda from a South American black hawk and

Physaloptera sp. from a sacred ibis.

All of the species of Physaloptera found in birds have been

recorded from the order Falconiformes ; P. acuticauda has the

widest host range, having been found in Falconiformes, Galli-

formes, and Cuculiformes ; P. galinieri next with a host range

of two orders, Falconiformes and Coraciiformes. The remain¬

ing species from birds are restricted entirely to Falconiformes.

Many immature Physaloptera have been reported from birds

not in the order Falconiformes, thus suggesting accidental hosts.

The Physaloptera from the Reptilia have a spear-shaped left

spicule, a characteristic not found in the Physaloptera from the

Amphibia, Aves, or Mammalia.

The following parasite-host list of the Physaloptera has been

arranged by classes, orders, and families to facilitate a compre¬

hensive view of host parasite relationships. The geographical

distribution is also given. Morgan reported in detail the Physal¬

optera from certain host groups (1941, 1942, 1948, 1944).

Physaloptera papuensis Johnson and Mawson, 1940, prob¬

ably from a bandicoot from Papua; P. peramelis Johnson and

Mawson, 1939, P. parvicollaris Johnson and Mawson, 1940, from

Morgan — Physaloptera Host-Parasite Relationships 283

Parameles nasuta (long-nosed bandicoot) ; P. peragale Johnson

and Mawson, 1940, P. thalacomys Johnson and Mawson, 1940,

from Peragale minor (rabbit bandicoot or bilby) and P. sareoph-

ili Johnson and Mawson, 1940, from Sarcophilus harrisi (Tas¬

manian devil), all from various Australian marsupials, cannot

be evaluated because of inadequate descriptions. The species

position of these Physaloptera must remain doubtful until the

types are re-examined and the number and mode of origin of

the uteri determined. Also, the male spicules and papillae should

be accurately pictured. Physaloptera troughtoni Johnson and

Mawson, 1941, from Rattus assimilis may be a synonym of P.

getula while the position of P. banfieldi Johnson and Mawson,

1941, from Melomys banfieldi (Muridae) is not clear. The de¬

scription of P. hieracideae Johnson and Mawson, 1941, from

Hieracidea orientalis (brown hawk) is not complete for species

identification.

LIST OF SPECIES OF PHYSALOPTERA AND HOSTS

ACCORDING TO CLASSES, ORDERS, FAMILIES

AND GEOGRAPHICAL DISTRIBUTION

Amphibia

1. P. amphibia Linstow, 1899. Anura (Ranidae) Philippine

Islands, Luzon.

Reptilia

2. P. abjecta Leidy, 1856. Syn. P. variegata Reiber, Byrd, and

Parker, 1940. Ophidia (Colubridae) U. S. A. (Pennsyl¬

vania, Georgia, Florida, Mississippi, Wisconsin)

3. P. obtusissima Molin, 1860. Syn. P. monodens Molin, 1860 ;

P. squamatae Harwood, 1932. Ophidia (Colubridae, Cro-

talidae, Boidae). Lacertilia (Scincidae). Brazil, New

Britain, U. S. A. (California, Pennsylvania, Ohio, Wis¬

consin, Mississippi, Texas) .

4. P. retusa Rudolphi, 1819. Syn. Spiroptera retusa Dujardin,

1845, P. largarda Sprehn, 1932 ; P. mucronata of Leidy,

1856. Lacertilia (Teiidae, Amphisbaenidae, Iguanidae)

Loricata ( Alligatoridae) . Brazil, U. S. A. (New York,

Utah).

284 Wisconsin Academy of Sciences , Arts and Letters

Aves

5. P. acuticauda Molin, 1860. Syn. P. truncata Schneider, 1866 ;

P. quadridentata Walton, 1927. Falconiformes (Accipi-

tridae). Cuculiformes (Cuculidae). Galliformes (Pha-

sianidae) Brazil, Africa, Mexico, French Guiana, U. S. A.

(Florida, Wisconsin, Arizona).

6. P. alata Rudolphi, 1819. Syn. Spiroptera physalura Dujar-

din, 1845; P. megalostoma Creplin, 1839; P. alata chev-

reuxi Seurat, 1914; P. alata nouveli Seurat, 1915. Fal¬

coniformes ( Accipitridae) . Italy, Austria, Africa, Chi¬

na, Germany, Brazil, Japan, France, Australia, Canada,

U. S. A. (Colorado, Wisconsin).

7. P. brachycerca Kreis, 1938. Falconiformes (Accipitridae) .

Angola.

8. P. crosi Seurat, 1914. Falconiformes (Accipitridae). Af¬

rica.

9. P. galinieri Seurat, 1914. Falconiformes (Accipitridae).

Coraciiformes (Bucerotidae). Africa.

10. P. hieracideae Johnson and Mawson, 1941. Falconiformes

(Accipitridae) . Australia.

11. P. losseni Ortlepp, 1937. Falconiformes (Accipitridae).

Africa.

12. P. mexicana Caballero, 1937. Falconiformes (Accipitri¬

dae) . Mexico.

13. P. rapacis Monnig, 1926. Falconiformes (Accipitridae).

Africa.

14. P. reevisi Chu, 1931. Falconiformes (Accipitridae). Chi¬

na.

15. P. subalata Schneider, 1866. Falconiformes (Falconidae,

Accipitridae) . Brazil, Europe.

Mammalia

16. P. mirandai Lent and Freitas, 1937. Marsupialia (Didel-

phiidae) . Brazil.

17. P. turgida Rudolphi, 1819. Syn. P. ackerti Hill, 1939. Mar¬

supialia (Didelphiidae). Brazil, U. S. A., Panama.

18. P. magnipapilla Molin, 1860. Edentata (Myrmecophagi-

dae). Brazil.

Morgan — Physaloptera Host-Parasite Relationships 285

19. P. papillotruncata Molin, 1860. Syn. P. pyramidalis Lin-

stow, 1879. Edentata ( My rmecophagidae, Bradypodi-

dae) . Brazil.

20. P. aduensis Baylis, 1928. Rodentia (Muridae). Africa

(Nigeria).

21. P. getula Seurat, 1917. Syn. P. bispiculata Vaz and Pere¬

ira, 1935. Rodentia (Muridae, Cricetidae, Sciuridae).

Africa, Brazil. U. S. A. (Florida, Georgia, Texas, Lou¬

isiana, Colorado, Montana).

22. P. massino Schulz, 1926. Syn. P. spinicauda McLeod, 1933.

Rodentia (Muridae, Sciuridae). Russia, Canada, U. S.

A. (Minnesota, Wisconsin).

23. P. muris-brasiliensis Diesing, 1861. Syn. Spiroptera bilabi-

ata Molin, 1860 ; P. circularis Linstow, 1897 ; P. sciuri

Parona, 1898; P. ruwenzorii Parona, 1907; P. inermisl

Linstow, 1906. Rodentia (Muridae, Cricetidae). Bra¬

zil, Hawaiian Islands, U. S. A. (Iowa, Georgia).

24. P. torresi (Travassos, 1920). Rodentia (Dasyproctidae) .

Brazil.

25. P. maxillaris Molin, 1860. Syn. P. semilanceolata Molin,

1860 ; P. mephites Solanet, 1909 ; P. mydai Baylis, 1926.

Carnivora (Mustelidae, Procyonidae) . Brazil, Europe,

Argentina, Trinidad, British West Indies, Canada, Mex¬

ico, Borneo. U. S. A. (New York, Iowa, Illinois, Wis¬

consin, Louisiana, California, Nebraska, Montana).

26. P. rara Hall and Wigdor, 1918. Syn. P. cerdocyona Sprehn,

1932 ; P. felidis Ackert, 1936 ; P. clausa of Caballero and

Peregrina 1938; P. turgida of Leigh, 1940. Carnivora

(Canidae, Felidae). Mexico, Germany, U. S. A. (Wis¬

consin, Michigan, Tennessee, Minnesota, Iowa, Missis¬

sippi,' Nebraska, Kansas, North Dakota, South Dakota,

California, Virginia, Arizona, Illinois, Kansas).

27. P. praeputialis Linstow 1889. Syn. Chlamydonema feline -

um Hegt , 1910; C. praeputialis Travassos, 1917 ; C. prae-

putiale Yorke and Maplestone, 1926. Carnivora (Feli¬

dae, Canidae, Viverridae) . Brazil, Batavia, Belgium,

British Guiana, Federated Malay States, China, Ceylon,

Dutch Guiana, India, Eastern Russia, Southwest Rus¬

sia, Puerto Rico, Panama, Africa (Nigeria, Tanganyika,

Zanzibar) Union of South Africa, Mexico. U. S. A.

286 Wisconsin Academy of Sciences, Arts and Letters

(Wisconsin, Iowa, Indiana, West Virginia, California,

Oregon, Arizona, Virginia, New Hampshire, Nevada).

28. P. torquata Leidy, 1886. Syn. P. papillotruncata of Cana-

van, 1931. Carnivora (Mustelidae, Procyonidae) . U.

S. A. (Pennsylvania, Wisconsin, Illinois, Montana, Ari¬

zona, Iowa).

29. P. anomala Molin, 1860. CARNIVORA (Felidae). Brazil,

Dutch Guiana.

30. P. brevispiculum Linstow, 1906. Syn. P. malayensis Ort-

lepp, 1922 ; Chlamydonema fuelleborni Mirza and Narain,

1934. Carnivora (Felidae Hyaenidae) . Ceylon, Feder¬

ated Malay States, Nigeria, India.

31. P. canis Monnig, 1929. Carnivora (Canidae, Felidae).

South Africa.

32. P. masoodi (Mirza, 1934). Syn. Chlamydonema masoodi

Mirza, 1934. Carnivora (Felidae). India.

33. P. terdentata Molin, 1860. Syn. P. digitata Schneider 1866.

Carnivora (Felidae). Brazil, Sudan.

34. P. clausa Rudolphi, 1819. Syn. Spiroptera clausa Du jar-

din, 1845; P. dispar Linstow, 1904. Insectxvora (Eri-

naceidae). Russia, China, Europe, Africa (Nigeria,

Tunis, Tanganyika).

35. P. immerpani Ortlepp, 1937. Insectivora (Erinaceidae) .

South Africa.

36. P. seurati Issaistchikoff 1926. Insectivora (Erinaceidae).

Russia, Europe.

37. P. limbata Leidy, 1856. Insectivora (Talpidae). U. S. A.

(Wisconsin, Iowa, Illinois, Maryland, Kansas, Vermont) .

38. P. bedfordi Ortlepp, 1932. Chiroptera (Rhinolophidae) .

South Africa.

39. P. cebi Ortlepp, 1923. Primata (Cebidae). South Amer¬

ica.

40. P. dilatata Rudolphi, 1819. Syn. P. multiuteri Canavan,

1929. Primata. (Lasiopygidae, Cebidae, Callitrichidae) .

Brazil, Panama, Peru.

41. P. tumefaciens Henry and Blanc, 1912. Primata (Lasiopy¬

gidae), India.

42. P. bonnei ‘Ortlepp, 1922, Host unknown. Probably Rep-

tilia. Dutch Guiana.

Morgan — Physaloptera Host-Parasite Relationships 287

UTERINE TYPE RELATIONSHIPS

The genus Physaloptera has the widest uterine range in the

subfamily Physalopterinae. Eight different uterine origins and

numbers are represented. The 2-A type has the largest host

spread in the group including the Reptilia (8 families), Aves (3

families), and Mammalia (5 orders and 9 families). The 2B

type includes the Reptilia (1 family) and Mammalia (4 orders

and 9 families). The 2C group is represented by Aves (3 or¬

ders, 4 families) Mammalia (1 order, 4 families). The remain¬

ing uterine types of 3, 4-D, 4-E, 5-1 5 G and 7-15H are found

only in mammals as shown by Chart 2.

RELATIONSHIPS OF THE ABBREVIATA

Abbreviata is characterized by 1 externolateral tooth, 1 in¬

ternolateral tooth and 2 double submedian teeth on each pseu¬

dolabium; usually the entire margin of the pseudolabia is den-

tated (Fig. 1-16). This genus has been recorded from many

hosts, namely, Amphibia, chiefly Reptilia, and Mammalia.

Schulz (1927) placed 23 species in this genus although approx¬

imately 27 species are now listed. Of the 27 species, 17 are

parasites of the Reptilia: Lacertilia (Varanidae, Agamidae,

Lacertidae, Anguidae, Scincidae, Geckonidae, Chamaeleontidae) ,

Ophidia (Colubridae, Elapidae, Boidae, Viperidae), Chelonia

(Testudinidae) . One species is from the Amphibia: Anura

(Ranidae, Bufonidae). The remaining 9 species are recorded

from the Mammalia: Carnivora (Felidae) [1 species] ; Artio-

dactyla (Suidae) [1 species] ; Primata (Hominidae, Lasio-

pygidae, Callitrichidae) [3 species] ; Rodentia (Muridae, Sciu-

ridae, Octodontidae) [4 species]. Many of the Abbreviata are

found in Africa, the remainder in Europe, Asia, Australia, and

North America.

The genus Abbreviata is not as widely distributed among

hosts as the Physaloptera. However, the majority of species

are found in Reptiles. Three classes, 5 orders and 22 families

are represented among the hosts. Chart 3 shows the distri¬

bution of the genus arranged by host family relationships. The

Amphibia is represented by 2 families, Reptilia by 12 and the

Mammalia by 8. The lizards are by far the most prominent

host.

288 Wisconsin Academy of Sciences , Arts and Letters

Geographically, 15 species are found in Africa, 3 in the

United States, and 2 in Australia. The locality of the hosts

of three species is not known. No species is very widely dis¬

tributed; A. abbreviata, A. varani, and A. caucasica have the

widest geographical range.

The parasite host list of the Abbreviata has been arranged

similar to that of the Physaloptera for convenience in study.

This genus does not occur in birds, although there have

been two cases on record of accidental or pseudoparasites of

Abbreviata from Aves. Abbreviata originally described from

Reptilia was reported by Linstow (1883) from Aconia alba

(white stork) and A. gemina (Linstow, 1899) originally de¬

scribed from a cat was reported by Railliet (1915) from a

domestic chicken. Morgan (1945) reported in some detail on

the genus Abbreviata.

UTERINE TYPE RELATIONSHIPS

Only 5 uterine types are found in the Abbreviata in com¬

parison with 8 for the Physaloptera. The 2-A type uteri is

restricted to the Reptilia (2 families), type 4-D to Reptilia (12

families) and Mammalia (3 orders, 6 families), 4-E to Rep¬

tilia (5 families) ; type 5-15-G and 7-1 5-H are found only in

the Mammalia as shown in Chart 4.

LIST OF SPECIES OF ABBREVIATA AND HOSTS

ACCORDING TO CLASSES, ORDERS, FAMILIES,

AND GEOGRAPHICAL DISTRIBUTION

Amphibia

1. A. ranae (Wallon, 1931). Anura (Ranidae Bufonidae).

U. S. A. (Indiana, Louisiana, Wisconsin, Illinois, Okla¬

homa). [Larval form.]

Reptilia

2. A. gracilis (Ortlepp, 1922). Lacertilia. Uganda.

3. A. leptosoma (Gervais, 1848). Lacertilia (Varanidae,

Agamidae) . Algeria.

4. A. abbreviata (Rudolphi, 1819). Lacertilia (Lacertidae,

Agamidae, Anguidae). Ophidia (Colubridae) . Spain,

Algeria, Turkestan, British East Africa.

Morgan — Physaloptera Host-Parasite Relationships 289

5. A. amaniensis (Sandground, 1928). Lacertilia (Agami-

dae). Africa.

6. A. antarctica (Linstow, 1899). Syn. Physaloptera alba

Stossich, 1902. Ophidia (Elapidae, Boidae). Lacer¬

tilia (Scincidae, Varanidae). Australia.

7. A . bancrofti (Irwin-Smith, 1922) . Syn. Physaloptera natri-

cus Kreis, 1940; P. physignathi Baylis, 1924. Lacer¬

tilia (Geckonidae, Agamidae). Ophidia (Colubridae).

Australia, New Zealand.

8. A. heterocephala (Kreis, 1940). Lacertilia (Agamidae).

Location not given.

9. A. leidyi (Walton, 1927). Lacertilia (Varanidae). Loca¬

tion not given.

10. A. oligopapillata (Kreis, 1940). Lacertilia (Scincidae).

Location not given.

11. A. ortleppi (Sandground, 1928). Lacertilia (Chamaele-

ontidae) . Africa.

12. A. pallaryi (Seurat, 1917). Lacertilia (Agamidae). Mor¬

occo.

13. A. polydentata (Walton, 1932). Lacertilia (Geckonidae).

British East Africa.

14. A. varani (Parona, 1889). Syn. Physaloptera quadrovaria

Leiper, 1908. Lacertilia (Varanidae Iguaindae) . Ophi¬

dia (Colubridae). Ceylon, India, China, U. S. A. (Mary¬

land, Wisconsin, Illinois).

15. A. achari (Mirza, 1935). Lacertilia (Agamidae). India.

16. A. paradox a (Linstow, 1908). Syn. Physaloptera affinis

Gedoelst, 1916. Lacertilia (Varanidae). Ophidia (Vi-

peridae, Colubridae). South Africa, Algeria, Belgian

Congo, Sudan, Nigeria.

17. A. tasmania (Ortlepp, 1937). Lacertilia (Chamaeleonti-

dae). Rhodesia.

18. A. terrapenis (Hill, 1941). Chelonia (Testudinidae). U.

S. A. (Oklahoma).

Mammalia

19. A. joyeuxia (Gendre, 1928). Artiodactyla (Suidae). Af¬

rica.

20. A. africana (Monnig, 1923). Rodentia (Muridae, Octodon-

tidae, Sciuridae). South Africa.

290 Wisconsin Academy of Sciences , Arts and Letters

21. A. leiperi (Skrjabin, 1924). Rodentia (Sciuridae) . Russia.

22. A. capensis (Ortlepp, 1922). Rodentia (Sciuridae). South

Africa.

23. A. musculi (Th waite, 1927). Rodentia (Muridae). Ceylon.

24. A. vandenbrandeni (Gedoelst, 1924). Carnivora (Felidae).

Belgian Congo.

25. A. caucasica (Linstow, 1902). Syn. Physaloptera mordens

Leiper, 1908. Primata (Hominidae, Lasiopygidae) . Cau¬

casus, Uganda, Africa, Arabia, North East Africa.

26. A. poicilometra Sandground, 1936. Primata (Callitrichi-

dae) . East Africa.

27. A. multipapillata (Kreis, 1940). Primata (Lasiopygidae).

Location not given.

RELATIONSHIPS OF THE SKRJABIN OPT ERA AND

PSEUDOPHYSALOPTERA

Skrjabinoptera is characterized by a single internolateral

tooth on each pseudolabium and is restricted entirely to the Rep-

tilia* At the present time 4 species are included in this genus

and are found in the Ophidia (Colubridae), Lacertilia (Cha-

maeleontidae, Iguanidae) .

Skrjabinoptera colubri and S. simplicidens have been report¬

ed only from Australia ; S. chamaeleontis only from Africa (Bel¬

gian Congo ; S. phrynosoma is the only well-known representative

of the genus. The geographical distribution is limited to South¬

ern United States and Mexico, the range of the horned toad,

American chamaeleon, and tree lizards. This species is restricted

entirely to the family Iguanidae. The 2A and 4D types of uteri

are represented in this genus, 2 species in each group. Morgan

(1943) reported on the occurrence of this genus in North

America.

LIST OF SPECIES OF SKRJABINOPTERA AND HOSTS

ACCORDING TO CLASSES, ORDERS, FAMILIES, AND

GEOGRAPHICAL DISTRIBUTION

Reptilia

1. S. colubri (Rudolphi, 1819) Ophidia (Colubridae) Australia.

2. S. chamaeleontis (Gedoelst, 1916) Lacertilia (Chamaeleon-

didae) Belgian Congo, Africa.

3. S. simplicidens (Ortlepp, 1922) Lacertilia Australia.

Morgan - — Physaloptera Host-Parasite Relationships 291

4. S, phrynosoma (Ortlepp, 1922) Lacertilia (Iguanidae)

Mexico, U. S. A. (Texas, New Mexico, Oklahoma, Califor¬

nia, Arizona, Utah, Idaho, South Carolina, Louisiana, and

Florida) .

The genus Pseudophysaloptera has pseudolabia similar to

Physaloptera, however, the male is without caudal pedunculated

papillae and spicules. Only one species is known at the present

time; P. soricina from Crocidura sp. (shrew) from Tanganyika.

Chen (1927) reported this parasite from Suncus coerulus

(shrew) from China. Morgan (1941) also recovered this species

from Sorex p. personatus (masked shrew). The genus so far as

known is restricted to the Insectivora (Soricidae). The female

possesses the 2-A type uteri. The species, reported by Yokagawa

(1922) from Sorex sp. (P. formosana) cannot be grouped con¬

veniently because of inadequate descriptions.

Bibliography

Alicata, J. 1937. Larval development of the spirurid nematode, Physalop¬

tera turgida , in the cockroach, Blatella germanica. Papers on Helm. 30

yr. Jubileum, K. I. Skrjabin. 11-14.

Baylis, H. A. 1934. On a collection of cestodes and nematodes from small

animals in Tanganyika. Ann. Mag. Nat. Hist. 13:338-353.

Boughton, R. 1937. Endoparasitic infestations in grouse, their pathogen¬

icity and correlation with meteroro-topographical conditions. Minn.

Tech. Bui. 121:1-50.

Chen, H. T. 1937. Some parasitic nematodes from mammals of South

China. Parasitology. 29:419-434.

Cram, E. B. 1931. Recent findings in connection with parasites of game

birds. Trans. Amer. Game Confer. 18:243-247.

Hobmaier, M. 1941. Extramammalian phase of Physaloptera maxillaris.

Molin, 1860 (Nematoda) Jour. Parasitol. 27:233-235.

Johnson, T. H. and Mawson, P. M. 1939. Sundry nematodes from Eastern

Australian marsupials. Trans. Roy. Soc. South Australia. 63:204-209.

- 1940 a. Nematodes from South Australian marsupials. Ibid. 64:95-

100.

— - - 1940 b. New and known nematodes from Australian marsupials.

Proc. Linn. Soc. New South Wales. 65:468-476.

- — 1941 a. Some parasitic nematodes in the collection of the Australian

museum. Rec. Australian Mus. 21:11-12.

- - 1941 b. Some nematodes from Australian birds of prey. Trans.

Roy. Soc. South Australia. 65:80-35.

Lin STOW, O. 1883. Nematoden, Trematoden, and Acanthocephalen gesam-

melt von Prof. Fedtschenko in Turkestan. Arch. f. Nat. 49:274-314.

292 Wisconsin Academy of Sciences , Arts and Letters

* - - 1899. Nematoden Aus der Berliner Zoologischen Sammlung. Mitt.

Zool. Mus. 1:5-28.

Morgan, B. B. 1941 a. A summary of the Physalopterinae (Nematoda) of

North America. Proc. Helm. Sec. Wash. 8:28-30.

- 1941 b. Additional notes on North American Physalopterinae (Ne¬

matoda) . Ibid. 63-64.

— - 1942 a. The Physalopterinae (Nematoda) of North American Ver¬

tebrates. Sum. Doctoral Diss. Univ. Wis. 6:88-91.

- 1942 b. The nematode genus Skrjabinoptera Schulz 1927. Lloydia.

5:314-319.

- 1943 a. The Physalopterinae (Nematoda) of Aves. Trans. Amer.

Micro. Soc. 62:72-80.

- — 1943 b. The Physaloptera (Nematoda) of Rodents. Wassmann

Collector. 5:99-107.

- 1943 c. The Physaloptera (Nematoda) of reptiles. Le Naturaliste

Canadien 70:179-185.

- = — 1945 a. The Physaloptera (Nematoda) of carnivores. Trans. Wis¬

consin Acad. Sci. 36:375-388.

- 1945 b. The nematode genus Abbreviata (Tavassos, 1920) Schulz,

1927. Amer. Midi. Nat. 34:485-490.

- 1943 d. New host records of nematodes from Mustelidae (Carni¬

vora). Jour. Parasitol. 29:158-159.

Ortlepp, R. J. 1922. The nematode genus Physaloptera Rud. Proc. Zocl.

Soc. London. 999-1107.

- 1937. Some undescribed species of the nematode genus Physaloptera

together with a key to the sufficiently known forms. Onder. Jour. Vet.

Sci. 9:71-84.

Rudolphi, C. 1819. Entozoorum synopsis, cui accedunt mantissa duplexet

indices locupletissimi. Berolini.

Schulz, R. 1927. Die. Familie Physalopteridae Leiper, 1908. Nematodes

und die principen ihre Klassifikation. Samml. Helm, arb.- Prof. K. I.

Skrjabin. 287-312.

Travassos, L. 1920. Contribuicoes para canhecimento da fauna helminto-

lojica Brazileira. Mem. Inst. Oswaldo Cruz. 73-77.

Wu, H. and Liu, C. 1940. Helminthological notes. II. Senensia. 11:397-406.

YokAGAWA, S. 1922. On a new species of Physaloptera ( Physaloptera for-

mosana) and the tumour caused by this parasite. Trans. Jap. Path. Soc.

12:201-202.

HIGHEST ABANDONED BEACH RIDGES IN

NORTHERN DOOR COUNTY, WISCONSIN

0. L. Kowalke

I. Introduction

The abandoned beach ridges in Wisconsin along Lake Michi¬

gan and Green Bay have been studied by several investigators

and all of them reported the existence of ridges much above the

present level of the water. James W. Goldthwait in 1907, in Bul¬

letin XVII of the “Wisconsin Geological and Natural History

Survey,” described his own studies of the beach ridges in eastern

Wisconsin, together with a compilation of the observations of

others. It is generally agreed that the highest of these ridges

were formed during the existence of the post-glacial “Lake Al¬

gonquin.”

A topographic survey by the writer and E. F. Kowalke in

1935 at Ellison Bay, Wisconsin, disclosed abandoned beach

ridges, respectively at 588 feet, 604 feet, and 650 feet above

sea level, whereas the level of water in Green Bay stood at 578

feet. Consideration of the locations of those beach ridges at

the 650 foot level, or Algonquin stage, together with other top¬

ographical features, indicated that at one time there were

“islands” where now there is consolidated land.

What shapes did these “islands” have and where were they

located? Were there islands farther to the south of Ellison

Bay? The purpose of this study was to answer these questions.

II. Surveying Procedures

Except for swampy and very stony areas, the northern end

of Door County is for the most part under cultivation and is

provided liberally with roads. It was thus convenient to walk

from the water’s edge inland either north to south or east to

west until a ridge was encountered. That spot was then marked

293

294 Wisconsin Academy of Sciences, Arts and Letters

on a map of land ownership printed on the scale of 1.5 inches

to the mile.

The elevations of such discovered ridges were then deter¬

mined by leveling with a transit and a stadia rod, the bases for

the elevations being the bench marks set at intervals along

Wisconsin State Highway 42 by the U. S. Coast and Geodetic

Survey. From these bench marks the water level in Green Bay

or Lake Michigan was ascertained when the elevation of a

beach ridge was being measured. The water level did not

change significantly from day to day in July and August of a

given year, but it did vary from year to year; and this study

extended over four summers during July and August. After

the elevations of the highest ridges in a given area had been

ascertained, and after their locations were noted on the land

ownership map, leveling was continued to ascertain where the

elevations of the crests of the ridges would lie in such areas

where no ridges were found or could be discerned.

Reference is now made to Plate I. The highest beach ridges

found are represented there by broad solid lines and the eleva¬

tions in feet above sea level indicated by the numbers in the

lines. The locations of the levels of the crests of beach ridges

in areas where no ridges were found are represented by lines

made of dashes. In other words, the lines made with dashes

represent where the elevation of the crest of the ridge would

lie if it were continued. The combination of a solid line and a

dashed line showing a given elevation is called a contour and

it must close on itself and make a complete loop.

III. The Ancient “Islands” and “Water Ways”

Inspection of the map in Plate I reveals that the contours

of the beach ridges at the 650 to 636 foot levels form six

“islands.” The size of the “islands” seems to increase the far¬

ther south it is located. The elevation of the ridges at the north¬

ern end of the map is 650 feet above sea level, whereas 18

miles to the south in the latitude of Kangaroo Lake the eleva¬

tion is 636 feet. This difference in elevations has been ascribed

to a lifting of the land due to the disappearance of the ice

farther north. George M. Stanley, in Bui . of the Geological

Society of America, 47 ; 1933, and 48 ; 1665, tells of his studies

of abandoned beach ridges on Georgian Bay in Lake Huron

and of finding, for example, that the highest Algonquin ridges

Kowalke — Abandoned Beach Ridges in Wisconsin 295

lay at 794 to 875 feet above sea level. Robert R. Schrock, in

Trans . Wisconsin Acad, of Science, Arts and Letters, 32; 203,

on “Geology of Washington Island,” points out that “Lake

Algonquin” stood at levels of 650 feet and 630 feet above sea

level.

On the “islands” the slope of the land towards Green Ray

is rather steep and often quite abrupt; the slope toward Lake

Michigan is in general gentle and uniform. On the Green Bay

ends of the “islands” from Ellison Bay south to Egg Harbor

the elevation of the high land stands between 700 and 770

feet above sea level. Each of the “islands” has a core of Ni¬

agara limestone which is covered with earth varying in depth

from a few inches to several feet; in some places the rock

cover is a glacial deposit of sand and gravel in the form of

mounds or small drumlins.

Because of the steep and abrupt slopes of the land towards

Green Bay, the beach ridges are often absent; there was no

support for the material rolled along by the waves. For exam¬

ple, there are no beach ridges at the 650 foot level on the cliff

in Section 16, Town 32 North, Range 28 East near Ellison Bay,

nor in the Peninsula State Park near Ephraim in Sections 15

and 23 and in Sections 21 and 28, all in Town 31 North, Range

27 East, nor along the cliffs at Fish Creek in Section 32, Town

31 North, Range 27 East. On the other hand, on the more

gradual slopes of land towards Lake Michigan, the highest

ridges are found in nearly all cases where it was possible for

wave motion to form them. Where such ridges lie on cultivated

areas, they are often obliterated by the successive plowings

and draggings.

Some aspects of the smaller “islands” are worthy of com¬

ment. Beginning at the north end, consider the “island” lying

north of Ellison Bay in Sections 2, 10, and 11, of Town 32

North and in Section 35, Town 33 North, in Range 28 East.

The waterway between it and the “island” lying just east was

narrow and the slopes of the ground on both “islands” is steep

at the entrance from Garret Bay in Section 2. The creek now

empyting into the Mink River rises in the southeast corner

of Section 2.

Consider next the northeast portion of the “island” between

Sister Bay and Ephraim, lying just south of the village of

Sister Bay in Sections 5, 6, 7, 8, 16 and 17 of Town 31 North.

296 Wisconsin Academy of Sciences, Arts and Letters

It may originally have been a separate island for it has a core

of limestone and the high point of present ground is around

680 feet above sea level. Note how close the lobe of the con¬

tour extending southeasterly through Section 7 approaches the

lobe which extends northward along the west side of Section

17 ; the ends of the lobes are only a few hundred feet apart.

The crest of the divide between these lobes had an elevation

of about 660 feet which is only around 20 feet above the crest

of the beach ridge contour. The soil of that divide on the north

side of Highway 57, at the southwest corner of Section 8, is

loam on top of gravel and sand, whereas immediately to the

south of the highway at that point, the soil is red clay.

Another small but interesting “island” is the one located

just north of Baileys Harbor in Sections 7, 8, and 17 of Town

30 North, Range 28 East. This “island” also has a core of

solid limestone and the beach ridge on it is intersected twice

by Highway 57. One such intersection is on the section line

between Sections 7 and 8, Town 30 North, Range 28 East and

is shown in Plate II. The crest of the section of the beach

ridge at this place is about 10 to 12 feet above the adjacent

land. The ridge may be discerned in the upper left of the pic¬

ture winding its way toward the southeast.

A third small “island” is located south of Fish Creek in

Sections 4 and 9 of Town 30 North, Range 27 East. It has a

core of limestone and its upper surface is dome-shaped rising

to an elevation of around 660 feet. There are no evidences

of beach ridges at the 636 foot level.

Although it is not now an “island,” the area just west and

northwest of Kangaroo Lake, lying in Sections 10, 14, 15, 22,

23, 26, 27, 34 and 35 of Town 30 North and in Section 2 of

Town 29 North, all in Range 27 East, could have been an

“island” if the barrier across the northern half of Section 16,

Town 30 North had not been there. The elevation of the crest

of this barrier is around 660 feet above sea level and the top

loam soil is thin and is underlain with a gravel deposit. Be¬

cause the barrier was under cultivation, no shore lines were

discernible and no digging was done.

In the absence of the barrier across the northern half of Sec¬

tion 16, there could have been two arms of Lake Michigan ex¬

tending northwesterly to meet the arm extending southeasterly

Plate II. — East face of cut through beach ridge by Highway 57 on section

line between Sections 7 and 8, T30N, R28E.

Plate III. — Ancient beach plane showing rounded field stones. Section 16,

T31N, R28E.

Plate IV. — Mound of rounded beach stones. SE.1^ of SE.1^ of Section 9,

T31N, R28E.

Plate V. — Sharp-edged stones in fence.

Plate VI. — Rock fence along Highway 57 in Section 20, T31N, R28E.

Plate VII. — Rock fence showing shell fossils protruding from faces.

SW.% of SW.% of Section 6, T30N, R28E.

Plate VIII.— Silicified corals — fossils.

Kowalke — Abandoned Beach Ridges in Wisconsin 297

from Green Bay at Fish Creek. At present there are, in fact,

two creeks flowing into Lake Michigan where once arms of

water from Lake Michigan could have existed.

Cursory examination of the region lying between Egg

Harbor on Green Bay and Clarke’s Lake south of Jacksonport

on Lake Michigan suggests the existence of additional “islands”

and water ways. Circumstances have, for the present, pre¬

vented additional surveys in this area.

IV. The Stones of the Fields

The stones now lying on the ground tell something about

whether they had been subjected to attrition or to other

agencies which reduced their size.

Wherever the waters of ancient “Lake Algonquin” had cov¬

ered gently sloping beaches, the rocks now found are well

rounded. An example of such beach is the one extending north¬

westerly from North Bay on Lake Michigan to Sections 9 and

16 in Town 31 North, Range 28 East. Sections 9 and 16 are

almost entirely under cultivation and the rocks dislodged when

plowing have been gathered into piles either in the fields or

along the fence lines bordering the highway. In Plate III is

shown the ancient beach with its piles of stones in the SE *4

of NE. % of Section 16 and in Plate IV a pile of rounded

stones in the SE. % of the SE*4 of Section 9.

On the other hand, where the water had not covered the

land, the stones removed from the surface by plowing gener¬

ally had corners which were quite sharp.

The field stones along Highway 57 south of the village of

Sister Bay are fiat pieces which originated from a chip lime¬

stone formation. Two illustrations will be shown: the first,

Plate V, is a stone fence near the center of Section 18, Town

31 North, and the second, Plate VI, is another fence along

Highway 57 in Section 20.

The field stones northwest from Baileys Harbor along

County Trunk Highway F are also sharp-edged but somewhat

larger than those shown in Plate V above. A stone fence a

little east of C.T. Highway F in the SW. % of SW. % of Sec¬

tion 6, Town 30 North, Range 28 East is shown in Plate VII.

Attention is called to the faces of the rocks shown in the

picture. These faces have portions of shell fossils protruding

298 Wisconsin Academy of Sciences , Arts and Letters

and sometimes bits of coral. The shells and bits of coral are

high in silica content and have resisted dissolution by carbo¬

nated waters more than the limestone matrix.

Searches in fields and also among stones in fences whose

elevations are higher than 650 feet above sea level reveal some

interesting and beautiful coral fossils. Some of these are shown

in Plate VIII. At first it was puzzling to account for the fact

that such delicate forms escaped destruction or dissolution.

Chemical analysis of several varieties of the corals and of

some shells showed that the content of silica ranged from

around 78 percent to around 97 percent. The silica being in¬

soluble in water charged with carbon dioxide was not etched

while lying in leaf mold, whereas the limestone was dissolved.

Hence the presence of delicately constructed corals and the

presence of shell remains protruding from some limestones.

The author is grateful to his wife, Winifred Titus Kowalke,

for her assistance in the field and for her constructive sugges¬

tions; and also to his brother E. F. Kowalke for drawing the

map in Plate I.

ARTIFICIAL HYBRIDS BETWEEN MUSKELLUNGE

AND NORTHERN PIKE

John D. Black and Lyman 0. Williamson

Wisconsin Conservation Department

Introduction

The study on which this report is based was made as a

part of the research program of the biology division of the

Wisconsin Conservation Department in order to determine the

degree of fertility between the muskellunge, Esox masquinongy

masquinongy (Mitchill) and the northern pike, Esox Indus

(Linnaeus), the characteristics of such hybrids, and the desira¬

bility of this hybrid as a game fish. One of the principal long¬

time objectives of the study is to determine the value of this

hybrid as a large, voracious predator for introduction into

overpopulated waters.

The present report concerns itself with the characteristics

of the two parent forms and of the hybrid, with preliminary

observations on the fertility of the hybrids, and with observa¬

tions concerning the status of the “tiger” muskellunge.

The study was initiated by the junior author and was in¬

terrupted by his call into the armed forces. If has been con¬

tinued and somewhat expanded by the senior author for the

present report.

History

Since the early studies of Francis Day, notably those report¬

ing the development of the zebra trout and other trout hybrids

by artificial fertilization (1884-1885), there has been consider¬

able interest in the study of the characteristics of fish hybrids.

This interest has lately been intensified by a comprehensive

series of studies and reports by Dr. Carl L. Hubbs and his

associates. A very good bibliography and summary of the gen¬

eral field may be found in Hubbs, Hubbs & Johnson (1943),

299

300 Wisconsin Academy of Sciences, Arts and Letters

Embody (1918) appears to be the first American to con¬

cern himself with interspecific- hybrids in the genus Esox. He

reported the artificial hybridization of the northern pike, Esox

lucius , with the chain pickerel, Esox niger.1

The characteristics of the parent species and the hybrid

are discussed and well illustrated by Embody. He found the

hybrids to be intermediate in the matter of scale development

on the operculum, but found that the juvenile hybrids were

like the young of the northern pike rather than being inter¬

mediate or with a distinct color pattern of their own. The

adult color pattern of the hybrids was not discussed.

Underhill (1939) reported on a repeat of the experimental

crossing by Embody and found that the hybrids were essentially

intermediate but that “the longest, however, shows the typical

spotted color of the adult pike” (E. lucius ). The fish reported

upon were in their second winter.

Greeley and Bishop (1933) reported, without comment, on

the discovery of specimens in the Upper Hudson River water¬

shed which they classified as hybrids between Esox americanus

and Esox niger .

Eddy (1941) and Eddy and Surber (1943) reported upon

the first hybrids between the northern pike and the muskel-

lunge, and Eddy (1941) has published some excellent figures

of the typical adult muskellunge, northern pike and of the

artificially produced hybrids.

One other hybrid combination is known in the genus Esox,

that of Esox niger X Esox vermiculatus based on two speci¬

mens from Missouri in the University of Michigan collection

(Hubbs, unpublished).

The problem of hybridization between these two species

and the relationship of the so-called “tiger” muskellunge to

the hybrids in Wisconsin, arose in 1937 when Dr. Edw. Schne-

berger (unpublished data) observed a small male northern

pike taking part in the spawning act of a pair of muskellunge

in Island Lake, Rusk County, Wisconsin. Early in June of that

year he obtained a specimen about 20 inches (509mm.) long

from Island Lake that displayed both muskellunge and northern

pike characteristics and had some of the vertical bars of the

“tiger” muskellunge. Dr. Schneberger identified the fish as a

i Embody used the name Esox reticulatus for the chain pickerel. Terminology in the pres¬

ent paper follows that of Hubbs and Lagler (1941).

Black & Williamson — Muskellunge and Pike Hybrids 301

northern pike X muskellunge hybrid. This identification was

confirmed by Dr. Carl L. Hubbs, then Curator of Fishes at the

Museum of Zoology, University of Michigan. This specimen

is now in the University of Michigan^ Museum of Zoology,

and was re-examined during the present study.

In the spring of 1939 a group of muskellunge eggs were

fertilized with milt from a northern pike. The eggs were in¬

cubated in glass hatching jars at the Woodruff Hatchery, and

the fry were placed in rearing ponds where they reached a

length of 14 inches during the first growing season. This

growth was somewhat better than that attained by young of

either of the parent species during the same period. A similar

rapid early growth was reported by Eddy (1941). In 1940 six

lots of eggs were obtained and reciprocal muskellunge X north¬

ern pike crosses were made with fish of various color patterns.

Each lot of eggs was kept separate and the young reared in

separate rearing ponds. At the end of the first growing season

morphological and color characteristics were compared. There

were no noticeable variations in either morphology or color in

the six lots, nor did they differ from the 1939 specimens. No

special records were kept of the percent of hatch of these eggs,

but the hatch was good, probably in excess of 80%. When the

difficulties of handling small lots of eggs in the hatchery are

taken into account this figure is very satisfactory. A sat¬

isfactory hatch was likewise obtained from a lot of eggs hatched

during the 1945 season, also at the Woodruff Hatchery, and the

young hybrids again exhibited a more rapid growth than the

young of either of the parent species reared under identical

conditions.

The 1939 hybrids were planted out in six small, landlocked

lakes that were overpopulated with pan fishes. Other than a

small planting of 258 fish in 200-acre (81 hectares) Lake Win-

gra, Dane County, Wisconsin, the largest lake planted was

Trilby in Vilas county. This lake has an area of 150 acres (61

hectares) and was stocked on July 2, 1940 with 200 fingerling

hybrids. On September 13, 1940, 176 of these hybrids averag¬

ing 12 inches in length, were recovered by the use of fyke

nets.

The large adult hybrid material used in the present study

represents material collected from the 1939 plantings or fish

302 Wisconsin Academy of Sciences, Arts and Letters

that were retained in the Woodruff Hatchery ponds. Several

were recovered in the Minocqua Thorofare, a body of water

immediately below the hatchery which was planted with fish

from the 1939 and 1740 year classes.

Materials and Methods

With the exception of one young hybrid borrowed from the

University of Michigan Museum of Zoology and originating in

the experimental laboratory of the University of Minnesota, all

of the material for the present study came from Wisconsin. All

of the hybrids were produced in the Woodruff Hatchery, near

Woodruff, Oneida County, except the one from Minnesota and

the wild specimen from Island Lake. A total of 46 hybrids

were critically studied. Fourteen of these were between 86

and 100 mm. (3. 4-3. 9 inches) in total length, 9 between 101

and 200 mm. (4-7.9 inches), and the remaining 23 were over

200 mm. long, most of them being in excess of 20 inches. The

largest hybrid examined was an adult female 997 mm. (39.2

inches) long from Lake Wingra, Dane county. Most of these

were examined dead, either as fresh or preserved specimens.

Five adults were etherized, examined and returned alive to the

Woodruff ponds. Eight “tiger” muskellunge were examined by

the junior author while in the hands of fishermen, but critical

counts and measurements were not recorded from these.

Complete counts and measurements were made on 36 mus¬

kellunge, 26 being between 100 and 200 mm. (3.9-7.9 inches)

in total length and 10 being adults, the largest 920 mm. long

(36.2 inches) . All but two of these were examined dead. Other

examinations, notably pore counts, and development of cheek

scales were made on other adults, but these were live, un¬

anesthetized fish from spawning nets and certain measurements

and counts were not practical. The total number of muskel¬

lunge examined was 83.

Fifty northern pike were studied, three of which were un¬

der 100 mm. (3.9 inches) in total length, 22 were between 100

and 200 mm. (3.9-7.9 inches) and the remaining 25 were

adults, ranging up to 820 mm. (32.2 inches) in length.

Counts were made of the number of pores of the lateral line

system opening into the ventral surface of the mandible. These

are referred to as “mandibular pores.” Counts were made for

Black & Williamson — Muskellunge and Pike Hybrids 303

the separate half- jaws and were totalled for both jaws. These

are presented as Tables 1 and 2.

Counts were also made of the branchiostegal rays on both

sides. The analysis of this count apparently was more signifi¬

cant when totalled rather than when given for one side only,

and is presented in that manner in Table 3.

Measurements were made of various aspects of the head

and three are here presented as being of the most value. These

are the measurement of the snout into the length of the head;

the length of the maxillary into the length of the head; and

the width of the snout into its length. Measurements were

made by means of a divider and fractions, to the nearest tenth,

were estimated by visual projection in the customary manner.

These measurements are reported in Tables 4, 5, and 6.

The development of the scales on the cheek, long a stand¬

ard taxonomic criterion of the genus Esox, was evaluated by

estimating the degree of squamation to the nearest tenth.

These estimates are presented in Table 7.

Color notes and observations on the size of the mandibular

pores were made from living or freshly killed specimens, as

were the photographs for the figures.

Characteristics of Hybrids and Parent Forms

Mandibular Pores

Of the several characteristics now in use to distinguish the

muskellunge from the northern pike, that of the number of

mandibular pores is one of the simplest and probably the most

reliable single character. As may be seen from the counts in

Table 1, the northern pike almost always has five large pores

on each mandible, for a total count of 10, as indicated in Table

2. Individuals do show up where one pore has been doubled

and two pores are found to be occupying the site normally

occupied by one. One northern pike fingerling from Lake Ne-

bagamon, Douglas County, has only four pores on each mandi¬

ble, an abnormality apparently caused by the loss of the third

pore on each side.

Of 83 muskellunge on which the mandibular pores were

counted only two fish had as few as 11 pores. There were sev¬

eral 6-6 counts. The data are tabulated in Tables 1 and 2. In

304 Wisconsin Academy of Sciences •, Arts and Letters

TABLE 1

Mandibular Pores in Single Jaw

TABLE 2

Total Number of Mandibular Pores

addition to the number of pores present, the size of the pores

is a very reliable characteristic to distinguish northern pike

and muskellunge from one another. This character is one which

is not readily measurable nor reducible to tabular form. The

mandibular pores correspond in size to the head pores on each

species. The differences in size of head pores is well shown in

Figures 5 and 7, where the top of the head is pictured.

The hybrids closely approach the muskellunge in the size

and number of the head and mandibular pores. This is shown

in Tables 1 and 2 and in Figure 6. The average number of

pores in the paired mandibles of the hybrid is 12.67 contrasted

to a value of 13.86 for the muskellunge and the single pore

counts run consistently lower, but the figures are so close that

the identification of any single individual on this one character

alone is not possible. The mandibular and head pores do ap¬

pear to be very slightly larger on the hybrids than on muskel¬

lunge of comparable size, but the difference is not decided and

certainly the pores are not intermediate in size as might be

expected.

Black & Williamson — Muskellunge and Pike Hybrids 805

Branchiostegal Rays

The number of branchiostegal rays, like the mandibular

pores, is a character of great value in distinguishing northern

pike from muskellunge. The values given by Hubbs and Lagler

(1941) hold very well for Wisconsin specimens. The figures

for the total number of rays per fish are presented in Table

3. It will be noted that there is no overlap in number between

TABLE 3

Total Number of Branchiostegal Rays

the northern pike and the muskellunge. It was felt that this

should be a valuable character for the identification of hybrids,

but the counts revealed an almost identical count with that of

the muskellunge. The counts for the hybrids, in fact, averaged

slightly higher than for the muskellunge, the average total

number being 35.50 in the hybrid and 35.30 in the muskellunge.

It was noticed there was a considerable tendency among the

hybrids to develop some unusually small weak rays at the an¬

terior end of the series. This seems to indicate some disturbance

in the ray formation and although the number would indicate

complete dominance of this character, the splint-like nature of

certain of these rays, especially in the specimens with the

higher counts, is indicative of some hereditary tendency to re¬

duce the rays in strength if not in number.

Measurements

Observation indicated that there was a decided difference

in the development of the snout and the caudal peduncle on the

northern pike as contrasted to the muskellunge, and it appeared

that the hybrid specimens were intermediate in these character¬

istics. Preliminary studies indicated, however, that the differ-

306 Wisconsin Academy of Sciences , Arts and Letters

ences in the caudal peduncle were more apparent than real,

and this measurement was dropped. The values for the width

of the snout measured into its length and the length of the

TABLE 4

Length of Snout Measured into Head Length

TABLE 5

Width of Snout Measured into its Length

TABLE 6

Length of the Maxillary Measured into the Head

Black & Williamson — Muskellunge and Pike Hybrids 307

maxillary measured into the length of the head were continued

through as many specimens as were available. The differences

did not prove to be as significant as it was thought they might,

and there is a decided change in proportions dependent on the

size of the fish, but there remains a consistent difference in the

three forms and the hybrids do appear to be intermediate.

The break in development apparently occurs around the 200

mm. total length stage and consequently the tabulation of these

measurements has been made for fish above and below that

value. Comparisons should be made between fish of the same

size group. The tabulation of these measurements is presented

in Tables 4, 5, and 6. Figure 1 also shows to good advantage

the longer snout of the young muskellunge when contrasted

with that of the younger northern pike. The differences are

most pronounced in the young and become obliterated with in¬

creasing size.

Cheek Scales

Fishermen have long relied on the presence or absence of

cheek scales to distinguish muskellunge from northern pike. The

character has, however, been more or less relegated to a secondary

place in scientific determinations within recent years. Data were

collected on this character from 46 northern pike, 46 hybrids and

42 muskellunge, and tabulated on the degree of development of

cheek scales by tenths. The results are summarized on Table 7.

Only three northern pike were not fully scaled on the cheeks

and the development of scales on these individuals was marked.

TABLE 7

Development of Cheek Scales

308 Wisconsin Academy of Sciences , Arts and Letters

Only two muskellunge showed cheek squamation in excess of 0.5.

The adult hybrids appeared to be intermediate.

In the hybrids and the muskellunge the development of cheek

scales occurs rather late in the development of the fish. The dif¬

ferences in average values of specimens over and under 200 mm.

shows this very well. The scales appear first as minute embedded

particles, quite difficult to distinguish. Only four of 16 hybrids

under 100 mm. in length showed any development of cheek scales,

and all individuals above 200 mm. had at least a part of the cheek

covered with scales.

Coloration and Color Pattern

The color differences between the young of the three forms

under consideration is well brought out by the photographs

(Figures 1 to 4). In the first few weeks the muskellunge and

hybrid fingerlings have similar color patterns. There is, how¬

ever, a tendency for the markings on the hybrids to be more

sharply defined. This can be seen in Figure 1 by comparing

muskellunge and hybrid fingerlings of the same age. As the

fingerlings become larger, however, (about 70 mm.) the simi¬

larity is lost and the hybrids take on a very distinct pattern

and are readily distinguishable from true muskellunge. The

characteristic “tiger” markings are now developed in the form

of vertical bars, separated by white stripes. Figure 2 shows

this characteristic pattern in comparison to the muskellunge

and northern pike fingerling. In this photograph it is seen

that from the side view the dorsal region of the muskellunge

is a solid color, while the white stripes of the hybrid continue

into the dorsal area. These stripes manifest themselves into a

reticulation when the hybrid is viewed from the top (Figure 3) .

One strong and persistent character of smaller muskellunge

fingerlings is the presence of a white stripe down the dorsal

mid-line (Figure 4). This character gradually fades away and

by the end of the first growing season it has either completely

disappeared or is very faint. This stripe also tends to develop

on the hybrid, but does not normally continue beyond the nape,

as shown in Figure 3.

The adult muskellunge is famous for its color variations

but in most cases it can readily be distinguished from the

hybrid by the fact that the reticulations of the young hybrid

Fig. 2. — Muskellunge fingerling. Center: Muskellunge x northern pike.

Lower: Northern pike. All four-months-old fish.

Fig. 1. — Upper: Two muskellunge fingerlings three weeks old; five

weeks old. Lower: Two muskellunge x northern pike five weeks, and three

weeks old.

Fig. 4. — Dorsal view of muskellunge fingerling.

Fig, 5.— Head of northern pike, length 52 cm., showing size of head pores.

Fig. 6 —Head of muskellunge x northern pike, length 41 cm., showing size

of head pores.

Fig. 7, . Head of muskellunge, length 98 cm., showing size of head pores.

Fig. 8. — Dorsal view of hybrid x northern pike backcross. Five weeks old,

length 70 cm.

Fig. 9. — Lateral view of hybrid x northern pike backcross. Five weeks old,

length 70 cm.

Black & Williamson — Muskellunge and Pike Hybrids 309

persist throughout life and the adult fish when viewed from

above continues to show the light lines cutting through the

dark pattern of the back. This is but rarely the case in true

muskellunge. The adult pattern of the hybrid is well illustrated

by Eddy (1941). Certain individuals from Muskellunge Lake,

Vilas County, Wisconsin, very closely approach this hybrid

color pattern, but most true muskellunge can be readily identi¬

fied by the color pattern. The Muskellunge Lake fish are ap¬

parently pure muskellunge since there is no record of northern

pike from the lake, nor have there been any pike or hybrid

plantings there.

Comparison with Supposed Natural Hybrids

During the course of this investigation nine supposed natu¬

ral hybrids, generally described as “tiger” muskellunge, have

been examined. The characteristics of these fish compare fav¬

orably with the hybrids produced by artificial crosses. They

agree as to color pattern, size and number of head pores and

cheek squamation with the artificially produced hybrids. Ex¬

perienced muskellunge fishermen who have examined the large

adult hybrids, especially those from Lake Wingra (the largest),

declare them to be the same as the highly prized “tiger” of the

northern waters.

Additional evidence that the “tiger” muskellunge is in reality

a hybrid is found from the fact that so far no truly authentic

“tigers” have been reported from waters in which northern

pike were absent, but always from lakes and flowages where

both muskellunge and northern pike occur. It is apparent from

the account given by Eddy (1941) and Eddy and Surber (1943)

that the term “tiger” muskellunge is not applied to the same

fish in Wisconsin that it is in Minnesota, where the typical

muskellunge is considered a “tiger.” The striping of the arti¬

ficial hybrid, and of the wild so-called “tiger” in northern

Wisconsin is a great deal more distinct than that shown by

Eddy and Surber in their color plate. This plate is a very good

reproduction of the normal or typical muskellunge of this

region.

Fertility and Sex Ratio of Hybrids

It has been found that most of the hybrid fish produced

or discovered in North American freshwaters are sterile. It

810 Wisconsin Academy of Sciences, Arts and Letters

was originally hoped that this would be the case with the

hybrid muskellunge because it was planned to use the fish ex¬

tensively in the reduction of pan-fish overpopulations where the

permanent introduction of a large predator might upset the

balance of the lake and do more harm than the original over¬

population. Results within the past few months indicate that

the hybrid muskellunge is not altogether sterile, although it

undoubtedly has a greatly reduced fertility.

The largest hybrid muskellunge examined was a ripe female

seined from Lake Wingra on April 4, 1945. The ovaries were in

a hemorrhagic condition and were partially filled with normal¬

appearing eggs. Microscopic examination of sections from the

fresh ovary made and studied by Professor R. K. Meyer of the

University of Wisconsin Department of Zoology, revealed that

ovulation had occurred. A second large female from Lake

Wingra appeared to have likewise spawned at least part of her

eggs.

A large female of the 1939 hatch was examined at the Wood¬

ruff Hatchery in the spring of 1945, along with five males of

the 1940 hatch. Although northern pike had completed their

spawning run at the time and muskellunge were spawning,

these fish were all in the “hard” condition. The female and one

of the males were injected with a suspension of whole, acetone-

dried pituitary of carp, 100 milligrams of the dried carp pitu¬

itary being injected into the female and 50 milligrams into the

male. (Hasler, Meyer and Field (1940) induced muskellunge

to spawn with this dose.) Within five days the female yielded

eggs. These were back-crossed with both muskellunge and

northern pike. Only a very small quantity of eggs (“several

ounces”) was secured from the injected female. The eggs were

divided into equal lots and one fertilized with milt from a north¬

ern pike, the other with milt from a muskellunge.

Only a very few of the backcross combination of hybrid X

muskellunge hatched and all of these fry died in the first few

hours after hatching.

Both the hatch and the survival of the other backcross, i.e.,

hybrid X northern pike, were much better. In spite of food-

supply difficulties caused by the abnormal spring of 1945, it

was possible to bring eight fish through to the five-weeks stage.

Three were killed and preserved at this size. The remaining

five will be kept alive, if possible, until they become mature.

Black & Williamsonr—MuskellMnge and Pike Hybrids 311

In hatchery practice two or three quarts of eggs are hatched

in each jar. It is extremely difficult to handle and hatch a

small lot of eggs properly. For this reason no definite con¬

clusion can be reached regarding the fertility of the hybrid

eggs. A rather low yield of eggs and a low fertility is indi¬

cated but further study is needed on this point.

The injected male did not yield milt and no attempt to

force ripening of the male by repeated injections of the hor¬

mone was made. Sectional studies of testes from the largest

male, obtained from tissues collected at the peak of the spawn¬

ing season, tend to confirm our opinion, based on gross ana¬

tomical examination, that the males are incapable of producing

sperm. The testes appear to be in an arrested state of de¬

velopment in all of the males examined.

Of the 27 hybrid specimens large enough for an accurate deter¬

mination of sex, 18 were females and nine were males. There

appears to be only a relatively slight disturbance in the normal

sex ratio.

The three hybrid X northern pike backcross specimens

available for study are remarkably uniform in appearance.

They are about five weeks old. Dorsal and lateral views of one

of these specimens are shown in Figures 8 and 9.

At five weeks of age the color pattern in the genus Esox is

remarkably uniform for all the species in the genus and there is

little distinctive about the backcross individuals.

The light mid-dorsal streak is strong and continues from

the tip of the snout to the anterior base of the dorsal fin, be¬

ing interrupted on the back half of the head. In this character

the backcross fish more closely resemble the muskellunge than

either the hybrid or the northern pike.

The three specimens at hand are peculiar in that the light

mid-dorsal stripe, the tip of the snout, side of the head and

tip of the chin are pinkish due to the transparency of the outer

skin and the showing through of the blood in these unpigment-

ed areas.

The branchiostegal ray counts on the three specimens at

hand are: 18-19, 19-19, and 18-18.

The mandibular pore counts are: 6-5, 6-6, and 5-6.

The scales of the cheek are not yet developed. The snout-

head measures are not distinguishable from hybrid specimens

of the same size.

312 Wisconsin Academy of Sciences, Arts and Letters

Summary

Artificial hybrids between the northern pike, Esox Indus

(Linnaeus) and the muskellunge, Esox masquinongy masqui-

nongy (Mitchill) have been produced in the Woodruff Hatch¬

ery of the Wisconsin Conservation Department. These have

been studied at various stages in their growth and comparisons

made between the hybrids and both of the parent forms.

Fertility in the cross is high, equal to that of either of the

parent species.

Analyses of the mandibular pore number, size of the head

pores, number of branchiostegal rays, various measurements

of the snout, and estimates of the development of the cheek

scales were made.

The development of the head pores is to some degree in¬

termediate, but definitely tends more toward the muskellunge

than to the northern pike, averaging 4.95 in the northern pike

(for one jaw), 6.34 in the hybrid and 6.95 in the muskellunge

(Table 1).

The branchiostegal ray count of the hybrid is as high as

that of the muskellunge, and is probably due to the development

of small rays in the anterior end of the series indicating only

a partial suppression of the full development found in the

muskellunge. The average total number of rays in the northern

pike was 29.64; in the hybrid 35.50; and in the muskellunge

35.30.

The snout measurements indicate a definite difference in

snout length of the muskellunge, which has the longer snout,

and the northern pike, and indicates that the hybrid is inter¬

mediate in this respect. The relative differences in snout length

tend to become obliterated with age and are not significant

in the adults.

Examination has revealed that the differences in the devel¬

opment of the cheek scales is still a useful character in the

distinction of northern pike and muskellunge. The hybrids are

intermediate in this character.

The color pattern of the young hybrid musky is quite dis¬

tinctive, the reticulated marking on the back being diagnostic

when contrasted with the dark area in the young muskellunge,

which is not broken by cross stripes but which is marked with

a sharp, distinct, white mid-dorsal streak. This mid-dorsal

Black & Williamson — Muskellunge and Pike Hybrids 313

streak of the muskellunge disappears with increasing size, and

the normal muskellunge presents an unbroken dark area. The

reticulations and light lines across the back of the hybrid persist

throughout life and together with the greater development of

the cheek scales in the hybrid offer one of the best key char¬

acters to its identification.

The characteristics of the adult hatchery-produced hybrids

appear to be identical with those of the so-called “tiger” mus¬

kellunge, and there is increasing evidence that the “tiger”

muskellunge in reality is a natural hybrid between the muskel¬

lunge and the northern pike.

Although the fertility appears to be low, recent evidence

indicates that the female hybrids, at least, are capable of natu¬

ral reproduction, that their eggs are fertile and that the re¬

sultant young survive at least in the backcross with northern

pike. Hybrid X northern pike backcross fish have been studied

at the five-weeks-old stage and five others will be reared through

to adults if possible. None of the very few hybrid X muskel¬

lunge backcross fry which hatched out survived more than a

few hours. At five weeks of age, the hybrid northern pike

backcross fish more clearly resemble muskellunge than northern

pike in color. The pore count averages 5.7 and the branchio-

stegal ray count averages 18.5 in the three fingerlings exam¬

ined. Further studies are being made on this aspect of the

problem.

Present data indicate that the male hybrids are probably

sterile.

Acknowledgments

Dr. Edw. Schneberger and Mr. D. John O’Donnell, Super¬

intendent of Fisheries and Chief Biologist respectively of the

Wisconsin Conservation Department, have jointly supervised

the present project and made it possible for us to carry on this

study. Valuable suggestions were made concerning the meth¬

ods of study by Prof. Carl L. Hubbs of the Scripps Institute

of Oceanography, University of California. Mr. Arthur Oehm-

cke, Area Supervisor of the Northeast Fisheries Area, Wiscon¬

sin Conservation Department, has provided all the facilities

for the spawning and care of the hybrids, the collection of the

parent species material and aided the study in every way pos¬

sible. Mr. Warren Churchill, Northeastern Area Biologist, Wis-

314 Wisconsin Academy of Sciences , Arts and Letters

consin Conservation Department, injected the adult hybrids,

has cared for many of the other details of the project, and is

continuing with the care of the young backcross material. Dr.

Reeve M. Bailey, Associate Curator, Fish Division, Museum

of Zoology, University of Michigan, loaned us the hybrid ma¬

terial in his charge. We are indebted to all of these men for

their valuable assistance.

Literature Cited

Day, Francis. 1884. On races and hybrids among Salmonidae. Proc. Zool.

Soc. London, 1884: 17-40, 376-880, 581-593, Ibid. 1885: 241-243.

Eddy, Samuel. 1941. Muskellunge and musky hybrids. Conservation Vol¬

unteer.: 41-44.

Eddy, Samuel and Thaddeus Surber. 1943. Northern fishes. Univ. Minn.

Press, Minneapolis. 252 pp.

Embody, G. C. 1918. Artificial hybrids between pike and pickerel. Jour.

Heredity.: 253-256.

Greeley, J. R. and S. C. Bishop. 1933. Fishes of the Upper Hudson water¬

shed with annotated list. In: A Biological survey of the Upper Hudson

watershed, Suppl. to the 22nd Ann. Rep. State of N. Y. Cons. Dept.

(1932). 64-101.

Hasler, Arthur D., Roland K. Meyer and Howard M. Field. 1940. The

use of hormones for the Conservation of Muskellunge, Esox masquinongy

immaculatus Garrard. Copeia, 1940. (1) : 43-46.

Hubbs, Carl L. and Karl F. Lagler. 1941. Guide to the fishes of the Great

Lakes and tributary waters. Cranbrook Inst. Sci. Bull. 18: 1-100.

Hubbs, Carl L., Laura C. Hubbs and Raymond E. Johnson. 1943. Hybrid¬

ization in nature between species of suckers. Contrib. Lab. Vert. Biol.

Univ. Mich. 22: 1-76.

Underhill, A. Heaton. 1939. Cross between Esox niger and Esox lucius.

Copeia. 1939 (4) : 237.

FOX HYBRIDS1

Leon J. Cole and Richard M. Shackelford2

Department of Genetics, University of Wisconsin

There are many kinds of foxes, classified in different species

and genera, but the present report is chiefly concerned with

two of these which have recently been brought under domesti¬

cation. These are the common red fox (Figure 1), including

the various color-phases derived from it, such as standard sil¬

ver (Figure 2), platinum, pearl platinum, and others, and the

Arctic fox, which has two principal color-phases, white (Figure

3) and blue (Figure 4). Under the designation red fox we are

including all the geographical “species’" and subspecies of the

red fox group, as they are apparently capable of interbreeding

freely; and for convenience we are referring to them all as

Vulpes vulpes (the North American equivalent being Vulpes

fulva) . For the same reason we shall not differentiate between

the described “species” of the Arctic fox, which is circumpolar

in its distribution. The typical form, Alopex lagopus lagopus, is

“restricted to Arctic regions of Europe and Asia, but said to

reach Bering Island in Bering Sea” (Anthony, 1928). In Alaska

and Arctic North America to Labrador and Greenland are other

related forms, some of which have been dignified by specific

names, but practical experience shows that they all interbreed

readily and may therefore be lumped together as the “lagopus

group” without further quarrel as to terminology. Unless spe¬

cifically indicated otherwise, therefore, references to “Vulpes”

will indicate members of the red fox group and “Alopex” those

of the lagopus group.

Black, or silver, and cross foxes, both of which are color-

phases of the red, occur in the wild with considerable frequency

1 Joint contribution from the Department of Genetics (Paper No 865) and Veterinary

Science. Supported in part by Wisconsin Agricultural Experiment Station, Project 614,

Fur Farm Research.

2 At present Lieutenant (j.g.), U.S.N.R. [Since returned]

315

316 Wisconsin Academy of Sciences , Arts and Letters

and their pelts have long been articles of trade. Other color

variants also occasionally appear among wild foxes, but most of

the newer color-phases have arisen and been established in

captivity. Domestication of the silver fox, and the present large

industry in this country and Europe, grew slowly from a small

and inauspicious beginning on Prince Edward Island in 1887

or 1888 (Ashbrook, 1923; Raynes and Jones, 1912). It was

not until 1910 that its “secrets” became generally known and

breeding stock available. Early breeding concentrated on the

black (dark silver) fox, though later lighter silvers became

popular. Other color-phases cropped up as mutations from time

to time, but these were looked upon as indications of “impurity”

in the stock and were disposed of quickly and quietly. In 1940

a shipment of “platina” pelts, a color-phase started in Norway

a few years before, was sent to New York. These skins brought

phenomenal prices and stirred up great interest in the “color-

phase fox” in this country (Cole and Shackelford, 1943). As a

consequence all new types that have appeared have since been

eagerly sought and not only cross breeding but hybridization

between species has been resorted to in the attempt to produce

new colors or color combinations.

The Arctic fox is typically white in winter and sooty brown

in the summer. The blue color-phase is everywhere bluish drab

and does not change to white in winter. In much of its range

the white phase predominates, but in others, notably the Aleu¬

tian Islands and the Pribilofs, the blue phase is in the majority.

Some management of the Arctic fox population was attempted

in Alaska by the Russians, beginning early in the nineteenth

century, when they are said to have introduced blue foxes to

Kiska Island and to have encouraged the killing off of the

whites while protecting the blues. Efforts to raise blue foxes on

islands farther to the east and south began about 1885 (Ash¬

brook, 1925). The foxes run wild on the islands, the natural

food obtainable being supplemented by products, mostly fish,

from the surrounding waters. In the winter the animals are

trapped for their furs, there being some selection of those to

be retained as breeders. While this may be referred to as fur

farming, it can scarcely be considered domestication. Within

the last decade or so, however, fur breeding and management

of the Arctic fox, after the manner of the red fox color-phases,

Cole & Shackelford — Fox Hybrids

317

has become quite widespread. The blue fox has proven to be

very amenable to such conditions.

Vulpes-Alopex Hybrids

There appear to be few, if any, records of Vulpes-Alopex

hybrids prior to what may be called their period of domesti¬

cation. This cannot be due entirely to isolation since the north¬

ern range of Vulpes overlaps that of Alopex to a considerable

extent. More likely it is due to a natural antipathy of the two

species. Also the breeding season of the blue fox is normally

considerably later than that of the red.

Case 1. The earliest record of a hybrid between these spe¬

cies which we have been able to locate appeared in the American

Fur Breeder in 1929 (Anon., 1929). There it is stated:

“In December [1928?] a fox breeder from the northwest

sent a novel fox pelt into the New York market that cre¬

ated quite a sensation. The breeder claimed that the pelt

was the result of a mating between a silver fox and a blue

fox.

“The skin was described as a dark, smoky brown, with

silvery guard hairs, and lacking the reddish cast observed

in many blue foxes.

“The breeder stated that the possibilities of breeding

out a hybrid strain of uniform animals from mating blue

foxes with silvers is uncertain, as other pups in the same

litter were irregularly marked, one having the body pelt of

a silver fox with a blue fox’s tail.”

Although not so stated, it is to be presumed that this mating

was made with foxes in confinement and that the silver was

the male parent. The description of the pelt agrees well with

that of other known hybrids, although the variation said to

occur in other members of the litter is rather surprising. The

statement that one pup had a “blue fox’s tail” presumably im¬

plies that it did not have a white tip.

Case 2. The following year (Anon., 1930) the same journal

had another note of a hybrid litter, as follows:

“Mr. P. J. Haggard, owner of the Big Bend Fur Farms

at Coleharbor, North Dakota, reports that he has a litter

of puppies from a native red female, which was acciden¬

tally mated to a blue fox male.”

We have no further information of this alleged mating.

318 Wisconsin Academy of Sciences, Arts and Letters

Case 3. The next report, in point of time, was made by a

Russian worker (Starkov, 1940) and is of especial interest as

artificial insemination was employed to produce the cross. The

hope was not only to produce in this way a new color character¬

istic in the silver fox but to combine with it the high prolifi¬

cacy of the blue fox. Two silver foxes were each inseminated

on the first and second day of heat with sperm of the Arctic

fox, procured by masturbation. One of the females produced a

single hybrid pup. Its juvenile coat was gray and the post¬

juvenile coloring like that of the Arctic fox. Later it darkened

somewhat and assumed more the color of young silver foxes.

Growth rate was greater and the eyes opened earlier than in

either parental form; also, the body was longer than in silver

fox pups of corresponding age. In external appearance the

hybrid had resemblances to both parents. It had rounded ears

and not only made little jumps like the Arctic fox but growled

like one when held up. The legs were long, however, as in the

silver. The underfur was light slate color and dense as in the

Arctic, but the tail was white at the tip. It is not stated wheth¬

er the guard hairs were banded, nor is the sex of the pup men¬

tioned. This is mistakenly claimed to be the first demonstra¬

tion that it is possible to cross these two species.

Case 4. The American Fur Breeder of August, 1941, has

the following note (Anon., 1941) :

“Art Doyle of the All Star Ranch at Winnipeg reports

a successful mating between a pearl platinum male fox and

a Greenland blue female fox. A litter of nine pups resulted

consisting of six males and three females. Mr. Doyle says

the pups have white-tipped brushes and the ears of a stand¬

ard fox.”

Mr. Doyle has kindly furnished additional information

which differs slightly from the note above. It appears that of

the nine pups seven, three males and four females, survived,

so the complete litter was presumably five males and four fe¬

males. The surviving males were pelted and attempts made to

breed the females. Three of these were paired respectively

with pearl platinum silver, white-marked silver and standard

silver males, and one with a Greenland blue male. However, no

pups were produced by any of the matings and all the hybrid

females were pelted the following season. A pronounced pe-

Cole & Shackelford — Fox Hybrids 319

culiarity of these females was their fear during the mating

season of any male except a blue (Arctic).

The fur quality of the female hybrids was much better than

that of the males and more silvery, the silver being scattered

and spikey in appearance in the males. The guard hair was

shorter than in silvers; underfur blue.

Case 5. In October, 1941, the authors had opportunity to

examine on the ranch of Mr. Sanford Colpitts at Salisbury, N.

B., a specimen reputed to be from a mating, made by another

breeder, of a red or cross male to a female blue fox. Its gen¬

eral appearance, as well as certain specific genetic characters,

indicated its hybrid origin. It was from a litter of eight pups,

six of which were dark in appearance, similar to the one in

the possession of Mr. Colpitts. The color of the other two

pups was not stated; presumably they died very early.

In general appearance this hybrid, a female, was inter¬

mediate between the two parent species but resembled rather

more the blue fox. Size and ear length were intermediate ; fur

with the general texture of that of Alopex but of a silver-black

color which must have come from Vulpes. Further, she had a

striking white tail-tip, which is entirely foreign to Alopex.

Figure 6, a photograph taken at the time of our visit, shows

the intermediate appearance of the head, the predominantly

Alopex coat character and the white tail-tip, which last obvious¬

ly came from the male parent.

This vixen was three years old at that time. Attempts had

been made each year to mate her back to Vulpes males (sil¬

vers), but without success. She at no time showed evidence of

being in heat.

Case 6. According to a note in the American Fur Breeder

(Anon., 1941), John W. Green-Armytage, owner of the Crichton

Blue Fox and Fur Ranch at Clarkleigh, Manitoba, reported

that he had successfully crossed a white fox with a white-face

silver (see Figure 10). Mrs. Green-Armytage has kindly fur¬

nished us with additional information. The white male was

“Greenland type.” There were six pups in the litter, four

males and two females. Two of the pups had white faces, a

black stripe down the middle of the back, fading into pale red¬

dish on the sides, and a golden color on the belly. It was sug¬

gested from this color that they be called “golden platinums.”

The other four hybrids were much redder, but also had the

320 Wisconsin Academy of Sciences, Arts and Letters

black stripe down the back. The brush was well described as

“golden platinum” and ended in a white tip on all six pups.

Attempts were made to breed male and female hybrids to¬

gether and also to breed a male hybrid to white-face Vulpes,

but were unsuccessful.

Case 7. Silver foxes (Vulpes) were introduced into Iceland

at some time in the past and Kristin P. Briem writes in the

Black Fox Magazine (Briem, 1941),

“You might like to know if silver foxes have not got out

of the pens and are living wild in the mountains. Several

foxes have got away for a time, but they have all been

caught, and, mostly shot. This country has no dense woods

to give protection to the silver foxes as their countries of

origin had. I have only heard of one silver fox vixen that

has bred in the wild and had young ones. This spring a

vixen was discovered and traced to her lair. She had mated

to a white fox and had three pups. I am told two of the

pups are as silver foxes, but one is more like the white fox.”

This report would seem to indicate that crossing may oc¬

cur between the two species in the wild when an individual of

one species, within the range of the other, can find no repre¬

sentative of its own kind with which to mate.

Case 8 . In 1942 and 1943, Mr. Eugene Finlay of Jefferson,

Oregon, produced two litters from blue females bred to a sil¬

ver fox male. The first year this male bred a blue female that

had been running in the pen with him. The resulting litter

consisted of 10 pups. The following year a strange female

was put with the same male, but he had apparently overcome

the inter-species antipathy and “she could not bluff him.” This

female raised a litter of 12 hybrid pups. In 1944 Mr. Finlay

attempted to mate a male Cody platinum fox3 to a blue female.

They were placed in adjoining pens, with a six-inch space sepa¬

rating them, to enable them to become acquainted. When put

together, they exhibited sexual interest but successful mating

was not obtained. A female hybrid retained by Mr. Finlay

came in season late and did not breed.

Four of the Finlay hybrids, two males and two females,

were obtained by the Associated Fur Farms, Inc., of New

Holstein, Wisconsin. One of the females died early in 1943, but

in October of that year, the senior author, accompanied by Dr.

3 A light strain of platinum silver. See Cole and Shackelford (1943).

Fig. 1. — North American Red Fox (Vulpes) .

Photograph courtesy Wisconsin Conservation

Department.

Fig. 2. — Standard Silver Fox (Vulpes). Courtesy

American National Fur and Market Journal.

Fm 3.-— White phase Arctic Fox (Alopex) in winter pelage. Photo¬

graph from Wisconsin Conservation Department.

1<ig. 4. — Blue phase Arctic Fox (Alopex) in winter pelage

Fig. 5. — Blue phase Arctic Fox in summer pelage, Aleutian Islands.

Photograph courtesy of U. S. Fish and Wildlife Service.

Fig. 6. — Vulpes-Alopex hybrid, female. From ranch of Mr. Sanford

Colpitts, Salisbury, N. B. Photographed October, 1941.

Fig. 7. — Vulpes-Alopex hybrid, male. From Silver male and Blue

female parents. Photograph from Wisconsin Conservation Depart¬

ment.

Fig. 8. — Vulpes-Alopex hybrids, female at left, male at right. Same

parentage as male in Figure 7. Photograph from Wisconsin Con¬

servation Department.

Fig. 9. — Hybrid shown in Figure 7 at left, Blue Fox at right. Photo¬

graph from Wisconsin Conservation Department.

Fig. 10.- — White-face color phase of Silver Fox (Vulpes) . Photo¬

graph from Mr. Bruno Delsman, Martian cL Wisconsin.

Cole & Shackelford— Fox Hybrids

321

C. K. Whitehair, had opportunity to observe the remaining

three animals (Figures 7 & 8) at the Elcho, Wisconsin, ranch

of the Associated Fur Farms. The female that died had acted

as if in heat during the season but had not been seen to breed.

Two or three weeks later it was thought she might be preg¬

nant but examination when she died a short time later revealed

no embryos. Efforts to breed the other hybrids proving futile,

these three animals "were generously donated to the Wisconsin

State Conservation Department in September, 1944, and have

since been available for study on the State Game Farm at

Poynette.4

TABLE 1

Approximate Measurements, in Inches, of Vulpes, Alopex, and Hybrids

Taken on live animals at Wisconsin State Game Farm, July 24, 1945.

These individuals of Vulpes and Alopex are, of course, not the actual par¬

ents of the hybrids.

In general features the hybrids are intermediate between

the two parent species, but when particular characteristics are

considered, some can be seen to be derived from one parent

and others from the other. The size, as can be seen from the

measurements given in Table 1, approaches more nearly to

that of Vulpes. When all three forms are fully furred out, the

resemblance to Alopex is heightened by the shorter guard hair,

which, together with the heavy coat of underfur, gives Alopex

its characteristic “wooly” appearance. In summer, when the

winter coat is shed, Vulpes and Alopex do not appear so dif¬

ferent in form (compare Figure 5).

4 We are greatly indebted to Mr. Eugene Finlay, of Oregon, and to Mr. Edw. P. Langen-

feld, of the Associated Fur Farms, for information concerning the history of these hybrids. We

also wish to thank Mr. W. F. Grimmer, Superintendent of Game Management, who has been

most kind in offering us every facility ; Dr. G. R. Hartsough [now with Associated Fur Farms,

Inc.] and Mr. Thalacher, who aided in handling these not-so-gentle animals ; and Mr. Staber

Reese, also of the Conservation Department, who took the photographs of them used in this

article.

322 Wisconsin Academy of Sciences , Arts and Letters

Ear-length is intermediate. The face (Figure 9) appears

more to resemble Alopex, but this also is probably in part due

to the fur.

The hybrids have the dense wooly under-fur of Alopex, but

the guard hair is shorter and consequently the general appear¬

ance is smoother. It is also much more silvered than the blue

fox. The blue is often silvered in the face, and to some extent

over the body, but this is due to the interspersal of white hairs,

and not to white bars on the dark hairs. Small samples of fur

plucked in December from the side of one of the male hybrids

and from a blue female are compared with silver in the follow¬

ing analyses:

Blue fox —

72 percent of the fibers fine, crinkly under-fur, 25 mm.

to 35 mm. long; about half of these gray throughout

their length, the others with an average of 7 mm. of

black at tip;

19 percent medium length guard hairs, 50-60 mm. long,

basal two-thirds wavy and white, apical third black;

9 percent long guard hairs (60-80 mm.), very little wavy,

white basally, black apically.

Hybrid —

68 percent fine, crinkly, gray to light gray under-fur, 15-30

mm., the longer ones with dark tips;

20 percent about the same length but less crinkly and black,

lighter towards the base;

12 percent guard hairs up to 40 mm. in length, black at

base and distally but with a narrow white bar , averag¬

ing 4 or 5 mm., just below the middle.

Silver—

20 percent wavy under-fur, 35-40 mm. in length, gray

basally and with black tip;

80 percent guard hairs, 55-70 mm. long; of these about

one-fourth gray to white at base, but otherwise black

throughout, the other three-quarters similar but with

a white bar of 8 or 10 mm. some 10-15 mm. from the

tip.

These descriptions and measurements would of course vary

a good deal in samples from different animals and if taken at

different seasons or from different parts of the body. They do,

however, serve to emphasize at least two points. The fur of

the hybrid resembles Alopex in being 70 to 80 percent under¬

fur, and is like the silver in having a white band on the guard

Cole & Shackelford — Fox Hybrids 323

hair, a condition that does not seem to be found at all in the

blue fox. This silvering, due to the banding of the guard hairs,

shows plainly in the photographs. Judged as silvers, they would

be put in about the “half” or “three-quarters” class. One of

the males — the one with the female (at the right in Figure 8)

— is distinctly lighter than the other (Figure 7).

In disposition the hybrids are “all blue fox” ; they bark like

blue foxes and stand up to an intruder in the pen where a sil¬

ver would sneak away unless cornered. They however seem

stronger and are certainly more vicious to handle than either

parent, and show no tendency to become tame, as blue foxes

often do.

Discussion

Some interesting conclusions can be drawn from Table 2,

which summarizes the principal facts relative to all the, hybrids

between Vulpes and Alopex known to us. It is significant that

the cross can be made in either direction, that is, with either

species as the male parent. Whether this can be done with

equal facility is not known, but it would seem that in either

case special manipulation is necessary to overcome natural

antipathies and to insure simultaneous breeding seasons. Fur¬

thermore, it is noteworthy, although natural, that the litter size

tends to correspond to that of the female parent. Normal litter

size in Vulpes is about 4 to 8, while Alopex may have from 8

or 9 to 12 or more pups to a litter. It will be noticed in Cases

6 and 7, in which Vulpes is the mother, the litters are of 6 and

3, respectively. Case 3 was not a normal mating and the litter

size of the others is not known. The four known litters of

Alopex females (Cases 4, 5 and 8) range from 8 to 12 pups

each.

The sex ratio among the pups would be of considerable

interest, but unfortunately these data are very inadequate.

The only litters for which the records are complete are Cases

4 and 6; these total nine males to six females. The statement

on Case 8, that the sexes were “about equal,” was an impres¬

sion from memory. According to “Haldane’s law” (Haldane,

1922), an excess of females would be expected in a cross as

wide as this, and Craft’s (1933) investigation of the mule and

other mammalian hybrids seems to support this hypothesis.

324

Wisconsin Academy of Sciences , Arts and Letters

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The present data for the fox are just the reverse, but they are

too few to admit of any definite conclusion.

Except for those cases in which a white Arctic-fox was

involved the hybrids are rather consistently dark and silvered,

irrespective of the direction of the cross. This would seem to

indicate clearly that the barring of the guard hair in Yulpes

is dominant to the plain, non-barred hair of the blue fox. Where

the white Alopex is used, the evidence is not so clear. In Case

6 there is a further complication in that the mother was a

white-face silver. White-face is a dominant character which is

lethal in the homozygous condition, hence white-face animals

are always heterozygous and segregation is to be expected

when they are mated to plain-faced individuals (Cole and

Shackelford, 1943). Such segregation evidently occurred in

this case as it is mentioned that two of the pups had white

faces and it is to be assumed that the others were unmarked.

How Case 7 is to be explained is not so clear. There seems to

be some indication of “white” and “silver,” but the descriptions

are too meager to make speculation profitable.

The situation regarding the white tip at the end of the

“brush,” which is characteristic of Vulpes, is confusing. In

general it would appear to be dominant in the cross, but in

Case 1 it is stated that one of the litter had the “body pelt of

a silver fox with a blue fox's tail.” This would suggest it was

much like the hybrids in Case 8 and also that there was segre¬

gation in the litter. The basis of this segregation is not ap¬

parent.

The incentive breeders have had in making the silver by

blue-fox cross has been the hope that, by backcrossing, the size

of the blues might be increased and the higher productivity

of the blues might be introduced into the silvers. Also the range

of color in the blues might be extended by bringing into that

species the barring from the silver. There seems little prospect,

however, that this goal can be attained. All attempts thus far

reported to breed the hybrids, either together or back to one of

the parental species, have been unsuccessful. As Starkov (1940)

pointed out, there is a big discrepancy in the chromosome

complements of the two species. In this laboratory Wipf and

Shackelford (1942) have verified 34 as the diploid number

for Vulpes. Preliminary study of Alopex by Wipf (unpub¬

lished) agrees with Andres (1938), who reported 52 chromo-

326 Wisconsin Academy of Sciences, Arts and Letters

somes for Alopex. As might be anticipated, the hybrid comple¬

ment is apparenty 43. With this situation, infertility of the

hybrids is not surprising. Reasoning from analogy, if any

fertility is to be obtained, it would probably come from breeding

a female hybrid back to the species that was its female parent,

that is, a female from a silver father and blue mother to a blue

male or from a blue father and silver mother to a silver male.

Other Hybrids

Since breeders are interested in the possibility of introducing

variations into the domestic fox by hybridization, it may be

worth while to discuss briefly what is known about crosses

with other more or less closely related species. We can make

no pretense of having reviewed the literature on the subject

thoroughly, but will mention certain instances, known or as¬

sumed, that have come to our attention.

Red fox X Gray fox. The pelt of the gray fox ( Urocyon

cinereoargenteus, Figure 11) is described as “coarse pepper-

and-salt, with the warm buffy under-fur showing through”

(Seton, 1937). It has little or nothing to recommend it as a fur

that would warrant its introduction into the domestic fox,

even were such possible. But in spite of the fact that their

ranges overlap in a considerable part of the United States there

are few if any authentic records of hybrids between them. Bez-

dek (1944) has recently described a pelt which showed charac¬

teristics of both species and it is assumed to be the result of

such a cross-mating in the wild. It had typical red fox face

and tail, including the white tip; back, sides and belly typi¬

cal of normal gray fox. The white tail-tip is interesting since

the tail of the gray is black at the tip. This was said to be the

only pelt of its kind among many thousands examined. It was

considered “almost worthless” by the fur trade.

Bezdek concludes that “the presence of only an occasional

report of such a case, and then in widely separated areas, sug¬

gests that when (and if) hybridization occurs in these genera,

the offspring do not reproduce.”

Fox X Dog. Hybrids between red fox and domestic dog

have been reported from time to time, with what authenticity

is not altogether clear. Reuter (1924) has reviewed the situa¬

tion at some length and concludes that while dogs and foxes

Fig. 11. — Gray Fox (Urocyon cinereoa/rgenteus ) . Photograph from

Wisconsin Conservation Department.

'

.

Cole & Shackelford-— Fox Hybrids

327

of opposite sex may sometimes manifest sexual attraction dur¬

ing their periods of heat, they have never been known to

produce young in the wild. Seton (1937, p. 510) states that

he once saw a curious creature at the Cincinnati Zoo that was

supposed to be a cross between a fox and a dog, but that its

appearance rather suggested a coyote. He also states that Lord

Cranley, in England, had for some time a cross between a black-

and-tan bitch and a tame male fox and that it was intermediate

between the two in color and other external characters, “but

strange to say, the tail tip was black.”

Heck (1932), Director of the Zoological Park at Hellabrunn-

Munich, reports and pictures an apparently authentic hybrid

between a male German fox (presumably Vulpes vulpes) and

a small Spitz-like female dog, although it is not definitely

stated that the mating was controlled. The hybrid was reddish

in color, but not so deep red as the fox. It had the fine-boned

appearance of the fox, a similar “gait” and the restlessness of

the fox. The coat was more shaggy than that of the fox and

the long, fine hair showed the influence of the Spitz mother.

At the time of reporting the hybrid pup was not old enough

for its fertility to be tested, but Heck seems to be convinced

that occasional individuals of all species hybrids may be fer¬

tile in backcrosses and that specific differences may be trans¬

ferred in this way. As further evidence of the ability of hy¬

brids to breed he cites the mating of a supposed fox-dog hybrid

with a female “small American wolf,” which resulted in three

pups. Unfortunately, too many of the reported cases of hybrids

and of their fertility are based on inferences from incomplete

information; only carefully controlled experiments can give

decisive results. Even if hybrids between dog and fox may be

produced, general experience of hybridization would make it

appear extremely unlikely that the cross could be carried be¬

yond the first backcross generation at most. Even if possible

or desirable, the incorporation of characters of one species into

the other would be a difficult task.

Coyote X Fox or Dog . There appears to be no record of

the coyote ( Canis latrans) hybridizing with the fox although

they occur together over a considerable part of their range.

This is perhaps just as well since the coyote seems to have no

characters that would be of interest to the breeder of foxes.

According to Seton (1937, p. 401) the coyote crosses readily

328 Wisconsin Academy of Sciences, Arts and Letters

with the dog and he cites a number of instances. He reports,

furthermore, in connection with one case, that the hybrids

“were intermediate in character and continued to be inter-

fertile with either stock, at least for two or more generations.”

Further evidence should be forthcoming before this case is

accepted at face value. Dice (1942) has recently given the de¬

tailed history of an authentic dog-coyote family. Unfortunately

the pups did not live long enough for their fertility to be tested,

which would be desirable under strict supervision.

Wolf X Dog or Fox. The North American Gray or Timber

wolf ( Canis nubilus and related species), and apparently also

the European wolf ( Canis lupus), cross so readily with the dog

that the geneticist wonders whether they can rightfully be

considered as separate species. But in spite of the fact that

the wolf and dog are so closely related, there appear to be no

records of the wolf crossing with either the fox or the coyote,

although their ranges overlap extensively. That these crosses

would occur in the wild seems extremely improbable, and so

far as is known they have never been attempted with confined

animals.

We may conclude, therefore, that while the red fox or its

color-phases may under unusual circumstances cross With other

related species as well as with the Arctic fox, it is unlikely

that any of these crosses will be of practical use to the breeder,

first because it is unlikely that such crosses can be carried be¬

yond the first hybrid generation, and second, because anything

that might be introduced in this way is in most cases of doubt¬

ful value.

Summary

This paper treats primarily of hybrids between the red fox

group ( Vulpes vulpes and related forms, including various color

phases) and the white and blue phases of the Arctic fox ( Alo -

pex lag opus) . Eight cases, culled from the literature, supple¬

mented by reports directly from breeders, are summarized. It

appears that this cross may be made in either direction with

fair facility, but so far as they have been tested, the hybrids

have been infertile. In one case the hybridization was ac¬

complished by means of artificial insemination.

Cole & Shackelford— Fox Hybrids

329

The hybrid is in general intermediate and to some extent

a combination of the characters of the respective parent species.

Rate of growth and size are more like the red fox but the

general appearance of the coat resembles more the Arctic fox,

especially in amount and fineness of under-fur. Silvering in the

blue fox is caused by interspersal of wholly white guard hairs,

while silvering in the red fox group is due primarily to a

white band on the otherwise black guard hair. This character

of banding on the guard hairs is carried by the hybrids and

appears to be definitely a dominant. The white tail-tip of the

red fox is usually expressed in the hybrid, but some have plain

tails and there may be segregation even within a litter.

When the white phase of the Arctic fox is used in the cross,

the hybrid pups show considerable reddish on the sides. In this

they resemble the Arctic whites in summer pelage, but the red

is even more distinct.

Crosses of white-face silver (heterozygous) with blues or

whites result in apparently normal segregation of the white-

face character in the pups.

The behavior, disposition and voice of the hybrids resemble

the blue parent more closely.

The size of the hybrid litters, as might be expected, con¬

forms to that of the species to which the female parent belongs.

Other hybrids, and reported hybrids, between species of the

dog family (Canidae) are mentioned briefly. These include,

besides the species discussed above, the gray fox (Urocyon

cinereoargenteus) , domestic dog, coyote ( Canis latrans) , and

wolf ( Canis lupus , and species). It is pointed out that these

crosses hold little promise of producing anything of value for the

practical breeder of fur animals.

Addendum

Just after the manuscript of the foregoing paper was com¬

pleted there appeared in the Canadian Silver Fox and Fur of

July, 1945, an article on “Arctic and Greenland Blue Foxes,”

by J. K. Aylwin, in which it was stated incidentally that ex¬

periments had been made on the “crossing of blues and whites

with the silver fox and subsequently with the white-face.” Up¬

on inquiry, Mr. Aylwin replied that his detailed records were

330 Wisconsin Academy of Sciences, Arts and Letters

not at present available to him, but he kindly sent consider¬

able general information which serves to corroborate and sup¬

plement that already in hand.

In 1931 a blue (Alopex) male was mated to a silver (Vul-

pes) female, with results similar to those reported above. The

progeny were intermediate in size, with the short ear of the

blue. Color a blend “but inferior, as far as the fur trade is

concerned, to an average specimen of either a straight blue

or silver.” General appearance more like blues than silvers.

In 1932 and years following the cross of white fox with

silver was made both ways. The general makeup and size of

the pups were the same as in the previous case, but when the

white Alopex is used a very definite red appears on the sides

of all the pups ( cf . Case 6 above).

In 1935 and subsequent years white foxes were crossed

with white-face silvers, in both directions. General appear¬

ance of the pups was the same but, as reported by Green-

Armytage, there was segregation of the white face, some of the

pups having it and others not. The red color on the sides is

even more pronounced in these white-face individuals, and is

pale, giving the appearance of a platinum cross fox. Others

are more or less red all over and may be likened to what is

known in the fur trade as a “bastard red” (smoky red of War¬

wick and Hanson, as presented by Ashbrook in his chapter

on “The breeding of fur animals” in the U. S. Department of

Agriculture Yearbook for 1937). This reddish color always

shows up when the white Arctic is used in the cross, on some

more than on others, and in some cases is comparable to the

red of a red fox. In most cases it is more definite than the

reddish that appears on the sides of the Arctic fox in summer.

These reddish hybrids go through some seasonal color change,

but it is not nearly as marked as in the Arctic fox ; the reddish¬

ness is always present and shows up strongest in the prime

coat.

The white tail tip was inconstant among these pups, some

of them having it, and others not.

The hybrids seemed to mate normally but, as in all other

reports that have come to us, they failed to reproduce.

331

Cole & Shackelford — Fox Hybrids

Addendum (August 1947)

One of the hybrid females mentioned in Case 8 was not ob¬

served in heat in 1945 or 1946, but came in heat the past breed¬

ing season. An Alopex male was placed with her, and she ac¬

cepted service on the 24th, 25th and 26th of March; she did not

appear to be pregnant at any time and no pups were born. The

other female died in November 1945 and was never observed in

heat. The male, having been completely castrated in March to

obtain material for chromosome studies, was pelted in Decem¬

ber 1946; castration appeared to have no affect on the notably

vicious disposition of this individual.

Literature Cited

Andres, A. H. 1938. On the chromosome complex in several Canidae. Cyte-

logia. 9 : 35-37.

Anonymous. 1929. Silver-blue hybrid creates sensation. Amer. Fur Breed.

1 : 26.

- 1930. Red and blue fox produce hybrid litter. Amer. Fur Breed.

S: 23.

- - — 1941a. Successful cross-breeding of blue and platinum fox reported.

Amer. Fur Breed. 1J+: 19.

— - 1941b. Crosses white fox with white face. Amer. Fur Breed. lh: 39.

Anthony, H. E. 1928. Field Book of North American Mammals. G. P.

Putnam’s Sons, New York-London. 674 pp.

Ashbrook, Frank G. 1923. Silver-fox farming. Dept. Bull. No. 1151 , U.

S. Dept. Agric. 59 pp.

- 1925. Blue-fox farming in Alaska. Dept. Bull. No. 1350 , U. S. Dept.

Agric. 33 pp.

Bezdek, H. 1944. A red-gray fox hybrid. Jour. Mammal. 25: 90.

Briem, Kristin P. 1941. Iceland reports. Black Fox Mag. 25: 46, 47.

Cole, Leon J., and Richard M. Shackelford. 1943. White spotting in

the fox. Amer. Nat. 77: 289-321.

Craft, W. A. 1938. The sex ratio in mules and other hybrid mammals.

Quart. Rev. Biol. 13: 19-40.

Dice, Lee R. 1942. A family of dog-coyote hybrids. Jour. Mammal. 23:

186-192.

Haldane, J. B. S. 1922. Sex ratio and unisexual sterility in hybrid animals.

Jour. Genet. 12: 101-109.

Heck, H. 1932. Bastarde. Kosmos (Stuttgart). 29: 406-409.

332 Wisconsin Academy of Sciences, Arts and Letters

Rayner, Ren. I., and J. Walter Jones. 1912. Domestication of the fox.

Amer. Breed. Mag. S: 37-45.

Reuter, M. 1924. Zur Fuchs- und Hundekreuzung. Zeitsch. Forst.- u. Jagdw.

56: 624-627.

Seton, Ernest Thompson. 1937. Lives of game animals. Vol. I. Cats,

wolves, and foxes. Literary Guild of America, Inc. New York.

Starkov, I. D. 1940. [Fox — Arctic fox hybrid.] (In Russian.) Zovetsch.

Zooteh. 11/12: 119.

Wipf, Louise, and Richard M. Shackelford. 1942. Chromosomes of the

red fox. Proc. Nat. Acad. Sci. 28 : 265-268.

COPPER IN LAKE MUDS FROM LAKES OF

THE MADISON AREA

M. Starr Nichols, Theresa Henkel, and Dorothy McNall

Wisconsin State Laboratory of Hygiene,

University of Wisconsin

The use of copper sulfate for the control of algal growths

started about 1904 with the work of Moore and Kellerman1 who

canvassed a large number of waterworks superintendents to

ascertain the extent of algal growths in open reservoirs. The

replies indicated the widespread nuisance created by the odors

resulting from the growth of algae in impounded waters and

caused them to work out dosages of copper sulfate necessary

for the destruction or control of the offending type of algae.

Later, after doing considerable field work in various parts of

the United States, they published additional work in tabular

form.2 These results are widely used as dosage tables when

algae control is contemplated. The city of Madison, Wiscon¬

sin was probably the first to apply copper sulfate to open lake

water on a large scale for the control of air-borne odors due

to algal growths. In 1918 bags of copper sulfate were dragged

behind moving boats on Lake Monona. The purpose was to

distribute the chemical in an attempt to alleviate the serious

odor nuisance which began to appear rather regularly at times,

notably during the summer months. About five tons were

applied to Lake Monona in this manner during that first sum¬

mer, but only with limited success in the control of the odors.

In the Brittingham Bay area of that lake the treatment proved

quite effective. During the following year and thereafter up

to 1925 similar treatments were attempted but only in a

rather irregular manner. In 1925 Dr. B. D. Domogalla* started

systematic treatment of the entire lake and used 108,600 pounds

of copper sulfate during May to September, inclusive. During

the growing season he added about 30,000 pounds of copper

333

334 Wisconsin Academy of Sciences , Arts and Letters

sulfate per month. His calculation of distribution dosage was

based on the upper fifteen feet of water and he states “In

applying copper sulfate care was taken, especially in the bays

and shallow waters, to keep the total concentration below six

pounds per million gallons which Whipple states will kill fish.”

During the growing season of each of the next twenty years

copper sulfate was applied to the waters of Lake Monona; the

amounts and total quantity varied somewhat depending upon

the need. Application control and better distribution technique

have resulted in the saving of a considerable amount of chemi¬

cal without reducing the effect of treatment.

The amount of copper sulfate crystals (CuSO4.5H20) ap¬

plied annually to Lake Monona is shown in Table 1.

TABLE 1

Copper Sulfate Applied to Lake Monona by Years*

* These data were furnished by The Madison Rivers and Lakes Commission through

the courtesy of Dr. B. D. Domogalla.

The amounts of copper sulfate applied to Lakes Waubesa

and Kegonsa, as furnished by courtesy of the Dane County

Clerk, are given in Table 2.

TABLE 2

Copper Sulfate Applied to Lakes Waubesa and Kegonsa

Nichols , et al. — Copper in Lake Muds 335

The amounts of copper sulfate applied annually to these

three lakes may vary somewhat from year to year; the data

given, however, are for the total amounts purchased each year,

and include a reasonable stock which is maintained for un¬

expected emergencies.

An outline map of the geographic relationship of the Madi¬

son Lakes is shown in Figure 1, and the physical data in

Table 9.

The question of the final disposition of this copper prompt¬

ed this report. If all the applied copper was deposited uniform¬

ly over the entire bottom surface of the lake, 1.2 grams of

metallic copper in the form of compounds of copper would be

found for each square foot of area. The calculation is for Lake

Monona with an area of 5.44 square miles; it does not include

the copper sulfate applied intermittently from 1918 to 1925.

In a series of experiments conducted by Woodbury, Palmer,

and Walton4 it was found that when copper sulfate was added

to distilled water in a concentration of 1.5 parts per million,

large mouth black bass were killed, and it appeared that this

concentration in distilled water slightly exceeded the toxic limit

for this species of fish. However, when a duplicate experiment

was carried out with water from Lake Mendota, which has an

alkalinity of about 170 parts per million, due principally to

calcium and magnesium bicarbonates, the definite toxic limit

was found to be about 200 parts of applied copper sulfate per

million. The pH of Lake Mendota water, which was the water

used in these experiments, has an average value of 8.0. When 200

parts per million of copper sulfate crystals are added to Lake

Mendota water a bluish-white precipitate forms and the pH drops

to 6.8. If, however, 200 parts of copper sulfate crystals per mil¬

lion are added to distilled water the pH drops to 5.6. The

water of Lake Monona has an alkalinity approximately equal

to that of Lake Mendota since the principal flowage into Lake

Monona is water from Lake Mendota. From these and other

qualitative data produced by our experiments we are of the

opinion that much of the copper sulphate added to lake waters

of notable alkalinity is precipitated as a basic copper compound

of somewhat variable composition, depending upon the condi¬

tions prevailing at the time the copper was added.

336 Wisconsin Academy of Sciences , Arts and Letters

337

Nichols , et al.— Copper in Lake Muds

Distribution of Copper in Lake Muds

Plagues of algae which have continually visited the Madison

lakes area for the past seventy-five years and probably long

before Madison existed, notably during the summer months,

have become increasingly more frequent in occurrence and

greater in intensity as the area has grown in population and

in agricultural and industrial activity. Some of the reasons

for this increase in frequency and intensity of blooming are

described in reports by Dr. Clair N. Sawyer whose research on

these lakes was performed from 1942 to 1944 under the spon¬

sorship of the Governor’s Committee. These reports, “Inves¬

tigations of the Odor Nuisance Occurring in the Madison Lakes,

Particularly Lakes Monona, Waubesa, and Kegonsa,” are to

be found in the library of the University of Wisconsin. It was

during the second period, 1943-44, that Dr. Sawyer studied lake

muds; and it was from his samples that most of the work on

the copper content was done, notwithstanding that previous

to this time Drs. Herman, Woodbury and Bartsch collected

about 60 samples of muds from various locations in the Four

Lakes chain. There was a total of over 200 samples of bottom

muds collected from nearly that many locations. The locations

may be ascertained by referring to the maps of the various

bodies of water which are included in this report. All of the

samples, except core samples, were collected from the bottom

surface by means of a dredge constructed on the principle of

a “clam shell” loading shovel. The core samples were obtained

by use of a Jenkins sampler obtained through the courtesy of

Professor W. H. Twenhofel of the University of Wisconsin. The

samples collected varied in consistency and composition from

black top slurry to rather stiff, compacted mud. A very few

of the samples collected near the shore line were principally

sand, while others were mixed with gravel. In all cases a rep¬

resentative sample consisting of about six ounces wet weight

was transferred from the collection dredge to glass bottles

with bakelite screw caps. Each sample was evaporated to dry¬

ness on a steam bath and afterwards dried in an oven at 105°

C. to remove the last trace of moisture. The entire sample

was ground in a mortar to homogeneity. Gravel and sandy

samples were not reduced to a powder since naturally occur¬

ring copper was not sought.

338 Wisconsin Academy of Sciences , Arts and Letters

Copper Determination

Two sets of determinations were made. The first determina¬

tion was for total copper and was intended to include all copper

from the organic matter and that copper precipitated as a salt

by virtue of the alkalinity of the lake waters. The second de¬

termination was made in such a manner that little of the

organically combined copper would be recovered. One gram

of the dried pulverized sample was taken for analysis in each

case.

Determination of Total Copper

One gram of the finely divided dried mud powder was trans¬

ferred to a Kjeldahl flask, 20 ml. of concentrated sulfuric acid

added, and the mixture digested until nearly colorless. The

sample was cooled and 100 ml. of distilled water was added

and the insoluble residue filtered off. The filtrate was then

treated with 2 ml. of 30 percent hydrogen peroxide and again

digested until entirely colorless. The digest was then cooled,

about 50 ml. of distilled water added and then neutralized with

concentrated ammonium hydroxide using a piece of litmus

paper as an indicator. The neutralized sample was then filtered

to remove the precipitated iron. The prepared filtrate contain¬

ing the copper was transferred to a 125-ml. separatory funnel,

5 ml. of alkaline ammonium citrate solution and 5 ml. of sodium

diethyldithiocarbamate solution were added and the whole mixed

well. (The alkaline ammonium citrate solution consists of 80 ml.

of concentrated ammonium hydroxide and 420 ml. of distilled

water in which are dissolved 20 grams of ammonium citrate.

The sodium diethyldithiocarbamate solution contains 0.5 percent

of the reagent dissolved in distilled water.) After about 5 min¬

utes had elapsed for the development of the copper carbamate

color, 10 ml. of amyl acetate was added and the color extracted.

Standards containing varying amounts of copper (0, 0.05, 0.1,

0.3 and 0.5 mg.) were prepared in exactly the same manner

and a curve drawn from readings using a No. 430 filter and

a Leitz-Maas photo-colorimeter. To make readings of the cop¬

per content of the samples, the amyl acetate solution of the

color complex was placed in the photo-colorimeter and the re¬

agent blank used to obtain 100 percent transmission, after

which the colored sample containing copper complex was im¬

mediately read for percentage transmission. By reference to

Nichols , et al. — Copper in Lake Muds

339

the standard curve the results in milligrams were then ob¬

tained. Since our results are given in milligrams per kilogram,

the milligrams of copper per gram of sample were then multi¬

plied by 1000.

Soluble Copper

A homogenous one-gram powdered sample of dried mud

was transferred to a Kjeldahl flask, 100 ml. of 0.1 N hydro¬

chloric acid was added and the contents boiled for one minute.

The mixture was filtered while hot, 15 ml. of concentrated

sulfuric acid added to the filtrate, evaporated to remove the wa¬

ter and digested until colorless. The sample was cooled, 50 ml.

of distilled water added, cooled again and then neutralized

with concentrated ammonium hydroxide. The solution was fil¬

tered to remove the iron hydroxide, treated with the copper

reagents, extracted with amyl acetate and the copper was de¬

termined, the same as for total copper.

Figure 2 shows the first forty-four samples as extracted

with amyl acetate and gives an idea of the intensity of the

colors obtained.

In Tables 3, 4, 5, 6, and 7 are given the data for both

soluble and total copper calculated as metal. The accompany¬

ing maps Figures 3, 4, 5, and 6 show the locations of the

sampling points from which the samples were obtained. The

TABLE 3

Lake Mendota. Total and Soluble Copper Found in Muds from Bottom

of this Lake at Various Depths

(For location of sampling points consult the map of this lake)

340 Wisconsin Academy of Sciences , Arts and Letters

TABLE 4

Lake Monona. Total and Soluble Copper Found in Muds from Bottom

of this Lake for Various Depths

(For location of sampling points consult the map of this lake)

Nichols, et al. — Copper in Lake Muds

341

TABLE 5

Lake Wingra. Total and Soluble Copper Found in Muds from Bottom

OF THIS LAKE AT VARIOUS DEPTHS

(For location of sampling points consult the map of this lake)

TABLE 6

Lake Waubesa. Total and Soluble Copper Found in Muds from this

Lake at Various Depths

(For location of sampling points consult the map of this lake)

842 Wisconsin Academy of Sciences , Arts and Letters

TABLE 7

Lake Kegonsa. Total and Soluble Copper Found in Muds from this

Lake at Various Depths

(For location of sampling points consult the map of this lake)

numbers on the lines correspond to the sample numbers given

in the tables ; the numbers in the circles show the total copper

in milligrams per kilogram of dry mud for each sampling point.

The tables also show the depth of the water at the place each

sample was taken.

Discussion of Results

It will be noted from Tables 1 and 2 that Lake Monona has

received copper sulfate treatment for algal control for a much

longer period than have Lakes Waubesa and Kegonsa. Further¬

more, during the years of treatment of Lakes Waubesa and

Kegonsa, these lakes were not “dosed” as heavily as Lake Mo¬

nona. This is reflected in the amount of copper found in the

layers of mud from these three lakes. In Table 3 for Lake

Mendota we find a low result for copper in the mud except for

two locations near which some slight copper sulfate treatment

VAHARA

RIMER

LAKE

MONO Nh

MAO A HA

mvm

LAKE .V

M CN OOTA

ig . M i$.J£L-J£,

YAMAHA

RWR

*£ ' JH c

' mm L'\

immA

' mo m

ilAHt; R

LAKE

AMYL ACETATE EXTMCrS

* cams smnmm

Fig. 2. — Showing the intensities of color produced by copper diethyl-

dithiocarbamate in amyl acetate solution from some of the determinations

of copper in lake muds.

Nichols , et ah — Copper in Lake Muds 343

had been applied. Lake Mendota results for copper in lake

muds, except for the two locations mentioned, serve as a normal

result for copper in muds from untreated waters. That these

low results for Lake Mendota muds are in conformity with

other untreated lakes is shown by results obtained from Lake

Wingra which are shown in Table 5. Likewise, a sample fur¬

nished by Professor W. H. Twenhofel, of mud from Allequash

Lake, located in the northern section of Wisconsin, showed a

similar amount of copper, 25 milligrams per kilogram of dried

mud. These two lakes (Wingra, Allequash) had received no

copper treatment. Contrasting these findings for non-treated

lakes with those receiving treatment, we find in all cases a sub¬

stantial increase in the copper content of muds from treated

lakes with the most marked increase in muds from Lake Mo¬

nona. This is what would be expected since this lake has re¬

ceived more copper and for a longer time than have the others,

Lakes Waubesa and Kegonsa.

The distribution of the precipitated copper in the treated

lakes is of interest. A study of Figure 4 shows the greatest

concentration of copper in the deeper parts of the lake. Sam¬

ple No. 163 in Table 4 shows the highest copper content found

— 1093 mg. per kilogram. While the depth here is only 12

meters that sampling point is near a heavily sprayed area on

the northeast shore. Other samples from deep sampling points

in this area are also high in copper, while shallower parts in

adjacent areas show less. It seems that the natural grading

process tends to carry the precipitated copper compounds to

lower levels. Somewhat the same results for distribution of

copper are shown for both Lakes Waubesa and Kegonsa in

Figures 5 and 6.

Results for the core samples are also worthy of note. From

Table 8 it is seen that in Mendota muds there is no appreciable

change in the copper content as the core penetrated to 8 feet,

while in Monona a five-fold increase is shown down to a pene¬

tration of four feet into the bottom mud. In Lakes Waubesa

and Kegonsa the penetration of copper had not extended below

the one-foot level. Interpretation of the penetration of copper

could be made upon the assumption that four feet of mud had

deposited since the treatments were begun some 20 years ago,

but such a rapid filling of the lake does not seem probable.

Possibly there was considerable more filling at the point at

344 Wisconsin Academy of Sciences, Arts and Letters

which the core sample was taken. Action of burrowing fauna

may have caused a mixing of the upper layers to this depth.

Any calculation of the total amount of copper deposited in

a lake such as Lake Monona is fraught with many difficulties

and assumptions. The area of Lake Monona is 5.44 square miles

but in many places the muddy deposit does not extend entirely

to the shore. Furthermore, the grading process in this lake,

as in many lakes, has carried the fine material out to deeper

water. Thus we find that the deposit containing excess copper

TABLE 8

Core Samples of Lake Muds from the Various Lakes

(For location of sampling points consult the map of these lakes)

Total Copper as Cu in mg. /kg. (dried mud)

TABLE 9

Physical Data and Alkalinity of Water of Lakes Studied

Type of Data

Common name of lake

Length in miles .

Width in miles .

Area in square miles. . .

Maximum depth meters

Average depth meters. .

Average drop between

lakes in inches .

Volume water millions

of gallons .

Alkalinity as CaCCb in

parts per million. . . .

Nichols , et al — Cower in Lake Muds

345

LAKE MENDOTA

D NUMBERS IN CIRCLES = MG- of Cu per kg. dry mud

— NUMBER OF SAMPLE. MUD ON LINE (see. tables?

Fig. 3.— -Outline map of Lake Mendota showing location of sampling

points, sample numbers corresponding to data in tables at these points,

and amount of copper found at the several sampling points, (c) refers to

core samples.

346 Wisconsin Academy of Sciences , Arts and Letters

Fig. 4.- — Outline map of Lake Monona and Lake Wingra showing loca¬

tion of sampling points, sample numbers corresponding to data in tables

at these points, and amount of copper found at the several sampling points,

(c) refers to core samples.

Nichols, et aL— Copper in Lake Muds

347

O NUMBERS IN CIRCLES = mg. cfCu per kg. dry mud

— NUMBER OF SAMPLE MUD ON LINE CSEE TABLES)

Fig. 5. — Outline map of Lake Waubesa showing location of sampling

points, sample numbers corresponding to data in tables at these points,

and amount of copper found at the several sampling points, (c) refers to

core samples.

348 Wisconsin Academy of Sciences , Arts and Letters

O NUMBERS IN CIRCLES = mg. of Cu per kg. dry mud

— NUMBER OF SAMPLE MUD ON LINE. (SEE, tables)

Fig. 6. — Outline map of Lake Kegonsa showing location of sampling

points, sample numbers corresponding to data in the tables at these points,

and amount of copper found at the several sampling points, (c) refers to

core samples.

Nichols, et al — Copper in Lake Muds 349

is about three feet thick in the deep part of the lake, while at

the periphery the deposit thins out to nothing. We can offer a

calculation of the amount of mud in the lake which contains

excess copper by assuming that we have a cone of mud ap¬

proximately five square miles in area at the base and three

and one-half feet deep in center; this assumption may be far

from true, however, since no attempt was made to “cone sam¬

ple” the lake bottom systematically. On this basis, as a be¬

ginning, we have a cone of 140 million square feet at base with

a maximum height of about 3.5 feet containing deposited cop¬

per. Furthermore, there should be (1/3 h a = volume; h =

height and a = area of base) approximately 160 million cubic

feet of mud deposit in this lake, allowing for the 0.44 square

mile of shore area which has little mud deposit. From Dr. Saw¬

yer's report the mud is approximately 15 percent solid matter

by weight, and considering the solid matter to have a specific

gravity of 2.5, we arrive at an approximate figure of eight

pounds for the average weight of one cubic foot of mud, or, on

a dry basis, of 12 pounds per cubic foot. At 12 pounds per

cubic foot and 160 million cubic feet of deposit we arrive at

1,920 million pounds of deposited mud on a dry basis. The

arithmetical average of the 71 copper determinations made on

muds from Lake Monona is 420 milligrams of copper per kilo¬

gram, or 200 milligrams (0.2 g.) per pound, of dry mud. On

this basis there were found 384 million grams, or 840,000

pounds, of copper calculated as the metal. Since 1.5 million pounds

of copper sulfate as CuS04.5H20, or 375,000 pounds calcu¬

lated as copper, were put into the lake the calculation is obvi¬

ously in error. It can be assumed that most of the copper-bear¬

ing deposit is found in the deeper water and not over the entire

bottom; so, if the major part of the deposit of mud is located

on the floor of the center one-third part of the area of the lake,

its magnitude will be of the order of 50 million cubic feet or

600 million pounds. Thus we arrive at a total of 120 million

grams of copper calculated as metallic copper, or about 260,000

pounds, found deposited in the lake as compared to 375,000

pounds applied. Although the total amount of copper depos¬

ited in the lake muds must probably remain unknown, it ap¬

pears that by far the greatest amount of that copper applied

remains as a deposit in the mud of the lake.

350 Wisconsin Academy of Sciences, Arts and Letters

A study of the equilibrium of this deposited copper with

that in solution in the overlying- water has not been attempted

in this work. Preliminary experiments tend to confirm the the¬

ory that the total alkalinity and the pH of the overlying water

control this solubility to a large extent, and because of the mag¬

nitude of these factors the precipitated copper is of very low

solubility and this solubility probably does not permit re¬

solution of the copper in sufficient quantity to be of value in

algae control.

Literature Cited

1. Moore, George T. and Kellerman, Karl F. 1904. A method of destroy¬

ing or preventing the growth of algae and certain pathogenic bacteria

in water supplies. Bulletin 64:15-44, Bureau of Plant Industry, U. S.

Department of Agriculture.

2. Moore, George T. and Kellerman, Karl F. 1905. Copper as an algicide

and disinfectant in water supplies. Bulletin 76:19-55, Bureau of Plant

Industry, U. S. Department of Agriculture.

3. Domogalla, B. D. 1926. Treatment of algae and weeds in lakes at

Madison, Wisconsin. Engineering News-Record, 97:24, 950-954.

4. Woodbury, Lowell A., Palmer, Robert C., and Walton, Graham. 1938.

Tolerance of fish to copper sulfate. Unpublished report to Committee

for Chemical Treatment of Lakes and Streams, State of Wisconsin.

(Authors have a copy of this report on file.)

PROCEEDINGS OF THE ACADEMY

Seventy-Sixth Annual Meeting

The seventy-sixth annual meeting of the Academy was held

in the Biology Building at the University of Wisconsin, Madi¬

son, Wisconsin, on Friday and Saturday, April 12 and 13, 1946.

Three other organizations participated jointly in the meeting,

— Wisconsin Junior Academy of Science, Wisconsin Archeologi¬

cal Society, and the Wisconsin Section of the American Chem¬

ical Society. The Academy section met in Room 102 Biology

Building, the Wisconsin Archeological Society in Room 356

Biology Building and the Wisconsin Section of the American

Chemical Society in Room 301 Biology Building. The annual

business meeting and election of officers was held on Friday

afternoon, followed by a tea at the home of Mrs. E. B. Fred.

The following program of papers was presented.

Academy Section

Friday morning

Dr. E. B. Fred, University of Wisconsin, Address of Wel¬

come; Norris F. Hall, University of Wisconsin, A Wisconsin

Chemical Pioneer— The Scientific Work of Louis Kahlenberg;

C. A. Elvehjem and W. A. Krahl, University of Wisconsin, Pel¬

lagra and Corn; George Urdang, American Institute of the His¬

tory of Pharmacy, How Chemicals Entered the Official Phar¬

macopoeias; Aaron J. Ihde, University of Wisconsin, Who dis¬

covered sulphur monochloride?; A. D. Hasler and L. V. Whit¬

ney, University of Wisconsin, An “Electric Eye” for Detecting

Schools of Fish.

Academy Section

Friday afternoon

John H. Kolb, University of Wisconsin, Background and

Foreground of Wisconsin’s Rural Communities; Leonard A.

Salter, University of Wisconsin, Do We Need a New Land

Policy?; Scudder H. Mekeel, University of Wisconsin, Where

is Social Science?

351

352 Wisconsin Academy of Sciences, Arts and Letters

Academy Section

Saturday morning

H. E. Reed, Burgess-Manning Company (Introduced by B.

S. Reynolds), The Industrial Utilization of the Cattail Spike;

Wm. D. Lewis and H. A. Schuette, University of Wisconsin,

Typha Seed Oil — preliminary report; E. A. Birge, Wisconsin

Geological and Natural History Survey, The Lakes of Vilas

County, Wisconsin; Karl U. Smith, University of Wisconsin,

Development of Behavior in the Loggerhead Turtle; Helen T.

Ness, Echo L. Price and Helen T. Parsons, University of Wis¬

consin, The Availability of Thiamine in Dried Yeasts ; H. F. Wil¬

son, University of Wisconsin, Electrostatic Effects Produced in

Dust Clouds Made with Finely Ground Minerals of Various Com¬

position; John C. Neess, University of Wisconsin (Introduced by

A. D. Hasler), A Contribution to the Ecology of the Bluntnosed

Minnow in an Artificial Pond; T. E. Allen and R. K. Chapman,

University of Wisconsin, Stimulation and Suppression of Some

Vegetable Plants by DDT Sprays and Dusts; K. R. Johansson,

University of Wisconsin, The Role of Diphtheroids in Bovine Tri¬

chomoniasis: Preliminary Studies; John Lilly and Mary Jane

Bradford, University of Wisconsin, Studies of Sugar Content and

Acidity of Corn as Related to Resistance to the European

Corn Borer; Bernice Cooper, State Teachers College, Superior,

The Abbe Prevost and the English Latitudinarians (By title) ;

C. L. Fluke and F. M. Hull, University of Wisconsin, Revision

of the Cartosyrphus Flies of North America (Syrphidae) (By

title) .

Chemistry Section

Saturday morning

Harry Posvic, Reduction Products of some Diphenyl Acids;

Calvin L. Stevens and S. M. McElvain, The Ortho Esters of

Phenylacetic Acids and Their Behavior; Warren J. Close and

A. L. Wilds; The Synthesis of Compounds Related to the Fe¬

male Sex Hormone Equilenin ; Kurt Rorig and S. M. McElvain,

Condensation of Benzaldehyde with l-Methyl-4-Piperidone ; B.

S. Sehweigert and C. A. Elvehjem, Nutritive Factors Needed

by the Monkey; W. R. Ruegamer and C. A. Elvehjem, Potas¬

sium Deficiency in the Dog; R. W. Rivett, J. J. Johnson and

Proceedings of the Academy

353

W. H. Peterson, Anti-biotics by Molds; Joseph Farber and

Paul Bender, Studies on Anthracene Transannular Peroxide. I.

Frequency Dependency for the Formation of Peroxide and Po-

larographic Analysis of the Peroxide.

Archeological, Folklore, and Museums Section

Saturday morning

Dorothy Moulding Brown, What Americana Meant to

Charles E. Brown; Sylvester Adrian, A Tavern Museum in

Muir Land; Albert 0. Barton, Early Contemporaries of Ella

Wheeler Wilcox; F. G. Cassidy, Pecatonica and Koshkonong

Place Name Origins; Herb Peters, Wisconsin's Only Covered

Bridge Museum; Dorothy Kundert, Monroe County Folktales;

Mitchell Red Cloud, Black Hawk Recollections; Mrs. Hope Nu-

zum, Black Hawk Trail Marking; Mrs. Willis Tyler, Aztalan

Museum Happenings; Palmer IJaugs, Aztalan Site Study; Cal

Peters, Prairie du Chien Museum; Walter Bubbert, Wisconsin's

Colorful Market Days; Theodore Mueller, A Milwaukee Legend

of “Yellow Dead" Island; Rollo Jamison, Beetown Tavern Mu¬

seum; Walter Bubbert, Census of Wisconsin Museum.

Junior Academy Section

Saturday afternoon

Robert Zusy, St. John Cathedral High School, Milwaukee,

Backyard Insect Collecting; Anita Kaufman, Chemistry Club,

Lincoln High School, Wisconsin Rapids, The Importance of Soil

Analysis ; Melborne Rabedeau, Seminar Club, Mary D. Bradford

High School, Kenosha, Cold Light; Kathryn Masterson, Science

Club P. J. Jacobs High School, Stevens Point, Applications of

Atomic Energy; Patricia Kasper, Dolores Demski, Mercedes

Ironside, Mercy Science Club, Mercy High School, Milwaukee,

Blood Will Tell; Robert Bard, Nature Club, Appleton High

School, Appleton, Astronomy Hobby; Lawrence Maurer, Sem¬

inar Club, Mary D. Bradford High School, Kenosha, Hydro¬

ponics; James Check, Science Club, P. J. Jacobs High School,

Stevens Point, Lift and Drag Coefficients of Airfoil Sections.

854 Wisconsin Academy of Sciences, Arts and Letters

Annual Academy Lecture

The annual Academy dinner was held Friday evening, April

12, at the Memorial Union. Seventy-eight members and guests

were present. Two addresses were made. President H. A.

Schuette presented his presidential speech on “Harm in the pot

no more.,, Bruce S. Wright, Lt. Commander R.C.N.V.R. gave

an illustrated talk on “The British Commandos.”

Academy Business Meeting

The annual business meeting was held in the Biology Build¬

ing on Friday afternoon, April 12, 1948.

The committee on nominations: H. C. Bradley (Chairman),

A. W. Schorger, Banner Bill Morgan, W. C. McKern, and P.

W. Boutwell presented the slate of officers for the next Academy

year. The election results are shown on the cover of this

Transactions.

The following persons were elected to life membership in

the Academy: Richard Fischer, Madison; Ruth Marshall, Wis¬

consin Dells, W. S. Marshall, Madison; C. E. Allen, Madison;

E. R. Maurer, Madison; and F. E. Turneaure, Madison.

Elected to honorary membership were Alexander Wetmore,

Frank Lloyd Wright, and Esther Forbes.

Proceedings of the Academy

355

Treasurer's Report

April 1, 1946

Receipts

Carried forward in Treasury, April 1, 1945 - $1,726.12

Receipts from the Junior Academy _ _ 101.00

Receipts from dues April 1, 1945 to April 1, 1946 — 1,052.79

Sale of publications _ 285.57

Interest on endowment _ 92.50

Total Receipts _ $3,257.98

Disbursements

Check cleared for payment April 2, 1945 _ $ 100.00

Purchase of U. S. Savings Bonds, Series G _ 300.00

Allowance to Secretary-Treasurer _ 200.00

Printing for Junior Academy _ 33.50

Purchase of Directory for Junior Academy _ .75

Stamps, envelopes, postal cards, return cards, express

charges, mimeograph material for newsletter _ 90.00

Printing for the Academy _ 34.10

Student help for wrapping, transactions _ 26.00

Safe Deposit Box _ 3.60

Printing cost for Volume 35, 1943 _ _ 1,317.94

Total disbursements _ _ $2,105.89

Balance, April 1, 1946 _ $1,152.09

Banner Bill Morgan

Secretary -Treasurer

The accounts of the Academy were found to be in order

and as reported above for the date April 1, 1946.

Auditing Committee

Raymond J. Roark ( signed )

Clement C. Williams ( signed )

356 Wisconsin Academy of Sciences , Arts and Letters

ENDOWMENTS AND ASSETS OF THE WISCONSIN

ACADEMY OF SCIENCES, ARTS AND LETTERS

1. U. S. Treasury Coupon Bond 1692B _ $1, 000.00

2. U. S. Treasury Coupon Bond 12894D _ 500.00

3. U. S. Savings Bond Registered Series G —

M1696059G _ 1,000.00

4. U. S. Savings Bond Registered Series G —

C1563347G _ 100.00

5. U. S. Savings Bond Registered Series G —

C1563348G _ 100.00

6. U. S. Savings Bond Series F — D494206F _ 500.00

7. U. S. Savings Bond Series F — M989457F _ 1,000.00

8. U. S. Savings Bond Series G — C3389339G _ 100.00

9. U. S. Savings Bond Series G — C3457898G _ 100.00

10. U. S. Savings Bond Series G — C3512841G _ 100.00

11. U. S. Savings Bond Series G — C3560656G _ 100.00

12. U. S. Savings Bond Series G — C3564110G _ 100.00

13. U. S. Savings Bond Series G — C415448IG _ 100.00

Total Amount of Endowment _ $4,800.00

14. U. S. Savings Bond Series G — C2386504G _ $ 100.00

15. U. S. Savings Bond Series G — C2386505G _ _ 100.00

16. U. S. Savings Bond Series G — C2386506G _ 100.00

17. U. S. Savings Bond Series G — C2386507G _ 100.00

Current Assets Invested in U. S. Bonds _ $ 400.00

18. Savings Account No. 3262, 1/1/46 _ _ $1,010.00

Total _ $6,210.00

Banner Bill Morgan

Secretary -Treasurer

The contents of the safety deposit box and the savings ac¬

count were found in order and as reported above for the date

April 1, 1946.

Auditing Committee

Raymond J. Roark

Clement C. Williams

THE CONSTITUTION OF THE WISCONSIN ACADEMY

OF SCIENCES, ARTS AND LETTERS

(April 11, 1947)

Article I — Name and Location

This association shall be known as the Wisconsin Academy of Sciences,

Arts and Letters, and shall be located at the city of Madison.

Article II — Object

The object of the Academy shall be the promotion of sciences, arts and

letters in the state of Wisconsin. Among the special objects shall be the

publication of the results of investigation and the formation of a library.

Article III — Membership

The Academy shall include four classes of members, viz.: life members,

honorary members, corresponding members and active members, to be

elected by ballot.

1. Life members shall be elected on account of special services ren¬

dered the Academy. Life membership may also be obtained by the payment

of one hundred dollars and election by the Academy. Life members shall

be allowed to vote and to hold office.

2. Honorary members shall be elected by the Academy and shall be

men who have rendered conspicuous services to science, arts or letters.

3. Corresponding members shall be elected from those who have been

active members of the Academy, but who have removed from the state.

By special vote of the Academy men of attainments in science or letters

may be elected corresponding members. They shall have no vote in the

meetings of the Academy.

4. Active members shall be elected by the Academy or by the council,

end shall enter upon membership on payment of the first annual dues.

Article IV — Officers

The officers of the Academy shall be a president, a vice-president for

each of the three departments, sciences, arts and letters, a secretary, a

librarian, a treasurer, and a custodian. These officers shall be chosen by

ballot, on recommendation of the committee on nomination of officers, by

the Academy at an annual meeting and shall hold office for one year.

357

358 Wisconsin Academy of Sciences, Arts and Letters

Their duties shall be those usually performed by officers thus named in

scientific societies. It shall be one of the duties of the president to pre¬

pare an address which shall be delivered before the Academy at the

annual meeting at which his term of office expires.

Article V — Council

The council of the Academy shall be entrusted with the management

of its affairs during the intervals between regular meetings, and shall con¬

sist of the president, the three vice-presidents, the secretary, the treasurer,

the librarian, and the past presidents who retain their residence in Wis¬

consin. Three members of the council shall constitute a quorum for the

transaction of business, provided the secretary and one of the presiding

officers be included in the number.

Article VI — Committees

The standing committees of the Academy shall be a committee on pub¬

lication, a library committee, and a membership committee. These com¬

mittees shall be elected at the annual meeting of the Academy in the same

manner as the other officers of the Academy, and shall hold office for the

same term.

1. The committee on publication shall consist of the president and

secretary and a third member elected by the Academy. They shall deter¬

mine the matter which shall be printed in the publications of the

Academy. They may at their discretion refer papers of a doubtful char¬

acter to specialists for their opinion as to scientific value and relevancy.

2. The library committee shall consist of five members, of which the

librarian shall be ex-officio chairman, and of which a majority shall not

be from the same city.

3. The membership committee shall consist of five members, one of

whom shall be the secretary of the Academy.

Article VII — Meetings

The annual meeting of the Academy shall be held at such time and

place as the council may designate. Summer field meetings shall be held

at such times and places as the Academy or the council may decide.

Special meetings may be called by the council.

Article VIII — Publications

The regular publication of the Academy shall be known as its Trans¬

actions, and shall include suitable papers, a record of its proceedings, and

any other matter pertaining to the Academy. This shall be printed by

the state as provided in the statutes of Wisconsin.

Constitution and By-Laws

Article IX — Amendments

359

Amendments to this constitution may be made at any annual meeting

by a vote of three-fourths of all members present; provided , that the

amendment has been proposed by five members, and that notice has been

sent to all the members at least one month before the meeting.

BY-LAWS OF THE WISCONSIN ACADEMY OF

SCIENCES, ARTS AND LETTERS

1. The annual dues shall be two dollars for each active member, to be

charged to his account on the first day of January of each year. Five

dollars, paid in advance, shall constitute full payment for three years'

annual dues.

2. The annual dues shall be remitted for the secretary-treasurer

and librarian during their term of office.

3. As soon as possible after January first of each year the secretary-

treasurer shall send to members statements of dues payable, and in case

of non-payment shall, within the succeeding four months, send a second

and, if necessary, a third notice.

4. The secretary-treasurer shall strike from the list of members the

names of those who are one year or more in arrears in the payment of

their dues, and shall notify such members of this action offering at the

same time to reinstate them upon receipt of the dues in arrears plus the

dues for the current year.

5. Each member of the Academy shall receive the current issue of the

Transactions provided that his dues are paid. Any member in arrears

at the time the Transactions are published shall receive his copy as

soon as his dues are paid.

6. The fee received from life members shall be set apart as a per¬

manent endowment fund to be invested exclusively in securities which are

legal as investments for Wisconsin trust companies or savings banks. The

income alone from such fund may be used for the general purposes of

the Academy.

7. The secretary- treasurer shall receive annually an allowance of two

hundred dollars for services.

8. The secretary-treasurer shall be charged with the special duty of

editing and overseeing the publication of the Transactions. In the per¬

formance of this duty he shall be advised by the committee on publication.

9. The Transactions shall contain in each volume: (a) a list of the

officers of the Academy, (b) the minutes of the annual meeting and

(c) such papers as are accepted under the provisions of Section 10 of

these By-Laws and no others.

10. Papers to be published in the Transactions must be approved as

to content and form by the committee on publication. They must repre¬

sent genuine original contributions to the knowledge of the subject dis¬

cussed. Preference shall be given to papers of special interest to the

\

360 Wisconsin Academy of Sciences , Arts and Letters

state of Wisconsin and to papers presented at a regular meeting of the

Academy. The privilege of publishing in the Transactions shall be re¬

served for the members of the Academy.

11. The Constitution and By-Laws and the names and addresses of

the members of the Academy shall be published every third year in the

Transactions. The Constitution and By-Laws shall also be available

in reprint form from the secretary-treasurer at any time.

12. Amendments to these By-Laws may be made at any annual meet¬

ing by vote of three-fourths of all the members present.

K