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TRANSACTIONS

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mss

OF THE

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WISCONSIN ACADEMY

OF

SCIENCES, ARTS, AND LETTERS

VOL. XVIII, PART I

Wi

H

MADISON, WISCONSIN

*915

;'.v'- -i *■ '-."i ■'■ /’: ", A A;--, :' V -^. ':' "/ ,/: <;•;'

The Transactions of the Wisconsin Academy of Sciences, Arts,

and Letters are issued in annual half -volumes, under the super¬

vision of the Secretary.

The price of this part is $1.00.

Arthur Beatty,

Secretary.

ff«!

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TRANSACTIONS

OF THE

WISCONSIN ACADEMY

t

OF

SCIENCES, ARTS, AND LETTERS

VOL. XVIII

INDEX

MADISON, WISCONSIN

INDEX

Page

Birge, Edward A., The Heat Budgets of American

and European Lakes. (With three Figures, and

five Tables) . 166

Birge, Edward A., The Work of the Wind in Warming

a Lake. (With Plates I-X) . 341

Charter of the Academy, Extracts from . 754

Davis, J. J., Notes on Parasitic Fungi in Wisconsin — I. 78

Davis, J. J., Notes on Parasitic Fungi in Wisconsin —

II . . . 93

Davis, J. J., Notes on Parasitic Fungi in Wisconsin —

III . 251

Grossenbacher, J. G., The Periodicity and Distribu¬

tion of Radial Growth in Trees and their Relation

to the Development of “Annual” Rings . 1

Harper, Edward T., Additional Species of Pholiota,

Stropharia, and Hypholoma in the Region of the

Great Lakes. (With Plates XI-XXIV) . 392

Juday, Chancey, Limnological Apparatus. (With

Plates XXXIV-XXXVIII) . 566

Juday, Chancey, Limnological Studies on Some Lakes

in Central America. (With four Figures and

three Tables) . 214

Kuhl, Ernest P., Chaucer’s Burgesses . 652

Mavor, James and Strasser, William, On a New

Myxosporidian, Hermeguya Wisconsinensis , n. sp.,

from the Urinary Bladder of the Yellow Perch,

Perea flavescens. (With three Figures) . 676

Members of the Academy, January 1, 1917 . 732

Munro, Dana C., Some Tendencies in History. Presi¬

dential Address, 1915 . 695

Page

Pierson, Merle, The Relation of the Corpus Christi

Procession to the Corpus Christi Play in England 110

Proceedings of the Academy, 1914, 1915, 1916 . 713

Smith, Gilbert M., A Monograph of the Algal Genus

Scenedesmus , based upon Pure Culture Studies.

(With Plates XXV-XXXIII) . 422

Smith, Gilbert M., A Preliminary List of Algae found

in Wisconsin Lakes . 531

Stewart, Alban, Some Observations Concerning the

Botanical Conditions on the Galapogos Islands . 272

Stewart, K.Bernice, and Watt, Homer A., Legends of

Paul Bunyan, Lumberjack . 639

Strasser, William, and Mavor, James, On a NewMy-

xosporidian . 676

Voss, Ernst, A True Bit of Instruction Showing why

we are under Obligations to Pay Taxes and Tithes

for the Preservation of Christian Peace and the

Avoidance of Trouble . 683

Watt, Homer A., and Stewart, K.Bernice, Legends of

Paul Bunyan, Lumberjack . . . 639

Young, Karl, William Gager’s Defence of the Aca¬

demic Stage . . . 593

TRANSACTIONS

OF THE

. "X;

V J U4_ 3 .1 j 1 6 '42

lYlus^

WISCONSIN ACADEMY

OF

SCIENCES, ARTS, AND LETTERS

VOL. XVIII, PART I

MADISON, WISCONSIN

1 91 5

t

i

I

j

TABLE OF CONTENTS OF VOLUME XVIII, PART I.

Page

The Periodicity and Distribution of Radial Growth in Trees

and their Relation to the Development of “Annual”

Rings. J. G. Grossenbacher . 1

Notes on Parasitic Fungi in Wisconsin — I. J. J. Davis. .. . 78

Notes on Parasitic Fungi in Wisconsin — II. J. J. Davis. . 93

The Relation of the Corpus Christi Procession to the Corpus

Christi Play in England. Merle Pierson . 110

The Heat Budgets of American and European Lakes. With

three Figures, and five Tables. Edward A. Biroe.. . 166

Limnological Studies on Some Lakes in Central America.

With four Figures and three Tables. Chancey Juday. 214

Notes on Parasitic Fungi in Wisconsin — III. J. J. Davis 251

Some Observations Concerning the Botanical Conditions on

the Galapagos Islands. Alban Stewart . . 272

THE PERIODICITY AND DISTRIBUTION OF RADIAL

GROWTH IN TREES AND THEIR RELATION TO

THE DEVELOPMENT OF “ANNUAL” RINGS.

J. G. Grossenbacher,

INTRODUCTION.

The study of the so-called ‘ 4 animal” rings in trees has re¬

ceived the attention of numerous investigators during past years

and still claims the interest of many. Research along that line,

however, is not as active as formerly apparently owing to the

general prevalence of the idea that the causes of ring formation

are beyond our ability to fathom at present ; although it is gen¬

erally conceded that an environment resulting in discontinuous

radial growth is somehow responsible for their occurrence.

In studying crown-rot of fruit trees1 I found that radial

growth and especially its distribution on trees during late sum¬

mer seemed to have a relation to the occurrence of the disease.

A number of more or less incidental remarks had been noted in

the literature concerning irregularities in the time of commence¬

ment and closing of cambial activity, but the irregularities oc¬

curring in fruit trees during late summer and fall were found

so marked that the literature was more carefully examined. The

number of significant papers on the subject proved so large and

the conclusions drawn so varied and contradictory that it seemed

desirable to discuss radial growth and the factors thought to

determine its distribution in a separate paper before writing up

1 Crown-rot, Arsenical poisoning and winter-injury. N. Y. State Agrl.

Expt. Sta. Tech. Bui. 12:367-411. 1909.

Crown-rot of fruit trees: field studies. N. Y. State Agrl. Expt. Sta.

Tecli. Bui. 23: 1-59. 1912.

1— -S. A.

2 Wisconsin Academy of Sciences, Arts, and Letters.

the results obtained from a histological study of the early stages

of crown-rot. $

The purpose of this paper, then, is to summarize in some de¬

tail most of the important hypotheses and investigations dealing

with the matter included in the title, to compare them with one

another and to bring out their relation to the writer’s observa¬

tions. Thus collecting the widely scattered ideas and summariz¬

ing the records of research along this line, it is hoped will stim¬

ulate a wider interest in the causes of periodic growth in trees

and encourage and lead to their reconsideration from a more

modern or quantitative standpoint. In the main the aim is to

restate the questions raised by the investigators, although some¬

times in a modified form. A restudy of the structural and

tension changes accompanying periodic growth may also lead to

an investigation of the enzymes active during radial growth and

to the effect which adverse changes of environment have

upon them while in an active condition. In any case studies of

this type will throw more light on the relation of a varying en¬

vironment to vegetative and reproductive processes in woody

plants and thereby increase the knowledge necessary for a com¬

prehensive investigation of their diseases. Most of the diseases

of trees which are of much economic importance and of most

scientific interest begin in the bark, and their origin seems to

have a definite relation to such radial growth and consequent

bark tensions, the normal adjustment of which is interfered with

by subsequent changes in the environment. Studies of that

kind will also help to clarify and perhaps correct some misappre¬

hension that may exist regarding the relation of mycology and

physiology to plant pathology.

Seasonal Periodicity of Growth.

It is generally held that seasonal periodicity or the alterna¬

tion of one or more growing and resting periods during the year

is a more or less unalterable inheritance of perennial plants of

temperate zones, but Klebs starting with his extensive investi¬

gations on the artificial control of periodicity in algae and fungi,2

has reached a very different conclusion regarding the periodic

3 Klebs, G. Willkiirliche Entwickelungsanderungen be Pflanzen. pp. 166.

Jena, 1903.

Grossenhacher — Radial Growth in Trees.

3

habit of such plants.* * 3 He maintains that the periodic or discon¬

tinuous habit of vegetative activity in plants is due to an alter¬

nation of favorable and unfavorable seasons of the year or to a

periodicity of the climate, and that it, therefore, may be made

continuous by modifying the environment. From his experi¬

mental work he concludes that dormancy is due to a reduction in

one or more of the factors essential for growth, such as tempera¬

ture, moisture and mineral nutrients, below7 the required amount ;

and that when such conditions occur the further manufacture

and accumulation of organic foods inhibits the action of the

enzymes necessary for growth. A timely increase in the limit¬

ing factor is said to either prevent or terminate a period of

dormancy in most cases. The reduction in the supply of mineral

foods was found to be a very important factor in inducing dor¬

mancy and, therefore, raising the temperature and increasing the

supply of water and mineral foods was often found to force

plants into growth. Berthold4 also concluded that a reduction

in the supply of nutrient salts is the chief factor inducing a ces¬

sation of terminal growth. This same conclusion was more re¬

cently drawn by Lakon5 who caused the buds of various decidu¬

ous trees and shrubs to open when cuttings were placed in

Enop’s solution. Klebs thinks that in many cases the individ¬

ual periodicity of the different branches and twigs of a tropical

plant are due to differences in transpiration and mineral nutrient

supply of such structures. It is thought probable that there

may be a periodicity in the supply of mineral nutrients in the

tropics which at times becomes a limiting factor inducing par¬

tial dormancy. On the other hand Smith6 maintained that

elongation growth of various Ceylon plants is controlled chiefly

by the temperature and water supply; sometimes one and then

the other or perhaps both acting together as the limiting factors.

In his interesting study of the second growth occurring on

* Klebs, G. liber die Rhythmik in der Entwieklung der Pflanzen.

Sitzungsber. Heidelber. Akad. Wiss. Math. Naturw. Klass. 23. 1911.

pp. 84.

4 Berthold, G. D. W. Untersuchungen zur Physiologie der pflanzli-

chen Organization. 2:131-257. 1904.

B Lakon, G. Die Beeinflussung der Winterruhe der Holzgewachse

durch die Nahrsalze. Ein neues Fruhtreibenverfahren. Zeit. Bot.

4:561-82. 1912.

6 Smith, A. M. On the application of the theory of the limiting factors

to measurements and observations of growth in Ceylon. Ann. Roy. Bot.

Gard. Peradeniya. 3:303-75. 1906.

4 Wisconsin Academy of Sciences, Arts, and Letters.

trees Spath7 comes to still another conclusion. According to

him the occurrence of the June elongation-growth which makes

its appearance fairly regularly on vigorous young trees of oak and

beech, is not determined by the environment but follows the

close of the spring elongation period after a fairly definite in¬

terval of time, and may even deveio pduring a drought or while

conditions are extremely unfavorable for growth. In Quercus

the resting period between the spring and June growth was

from 30 to 40 days and in Fagus from 15 to 20 days. It is said

to last 9 to 16 days in the former and 13 to 24 days in the lat¬

ter. The length of these second shoots is thought to depend

chiefly upon the amount of available water and is usually but

not always less than that of spring shoots. The species with the

June-elongation habit have a short but very active spring-

growth period as compared with those not having the June

growth. It was found impossible to prevent the June elonga¬

tion growth by reducing the food and water supply and by low¬

ering the temperature, nor could it be made to continue beyond

the ordinary period by supplying heat, moisture and food con¬

ditions favorable for growth.

Spath also found that the second growth is made up of three

types. In one kind the axillary buds of an elongating shoot devel¬

op into branches before they are fully formed. This happens in

Salix, Populus, Taxus, Buxus, Prunus, Pyrus, and is called syllep-

tic growth. In the second type known as June growth ( Johannes-

trieb) the buds are fully formed before they open after the ter¬

mination of the spring elongation growth. The third type is

called proleptic growth and is said to develop at any time during

summer from buds which normally would not have opened until

the following spring but which open early owing to wound or

some other strong environmental stimulus.

Neither sylleptic nor June elongation-growth was said to have

a zonation effect upon radial growth while the production of

proleptic shoots practically always resulted in more or less dis¬

tinct zonation of the radial growth. This was shown during

their development by the wood cells produced being wider than

those differentiated just before the new shoots appeared.

7 Spath, H. L. Der Johannistrieb. Ein Beitrag zur Kenntniss der

Pferiodizitat and Jahresringbildung sommergriiner Holzgewachse. Ber¬

lin, 1912. pp. 91.

Grossenbacher — Radial Growth in Trees.

5

The Beginning and Duration of Radial Growth.

Observations and statements regarding the commencement of

cambial activity or radial growth in spring are many but no

positive conclusion can as yet be drawn as to just where on any

particular species of tree it will begin one season after another.

In fact it seems to differ considerably for individuals of the same

species.

According to Strasburger8 in pines as well as in Picea, as

many as five layers of tracheids had been formed from the cam¬

bium in one-year shoots and considerable elongation growth had

occurred by the first of May, while at the bases and on the

trunks of eight-year old branches the cambium was still inactive.

In case of Robinia Pseudacacia and some other species, how¬

ever, radial growth was found to begin first on the trunk. But

in general cambial activity is said to begin in one-,year shoots

just back of the unfolding buds and to proceed downward to the

larger branches and trunks on which it usually begins uniform¬

ly and at about the same time from top to bottom. He found

that the cambium gives rise chiefly or almost exclusive to wood

cells9 in spring, and as the vegetative season advances, the pro¬

duction of phloem increases while that of wood cells decreases.

In trees of our zone wood formation is said to cease by mid-Au¬

gust while that of the phloem continues practically up to the

end of the vegetative season. Wood cells are therefore usually

matured before winter but phloem cells sometimes enter the

dormant season in an immature condition.

Pfeffer10 also says that “the secondary growth of xylem in

trees begins and ends sooner than that of the phloem.”

Hartig11 states that although no growth had occurred on April

15 on any of the sixteen-year-old trees under observation, by

May 5 the new radial growth in oak was about equal on all parts

of the trunk but that none had occurred underground; while

in maple, though the buds were farther advanced than in oak,

the growth as yet was confined chiefly to the one-year shoots.

8 Strasburger, E. Ueber den Ban und die Verrichtungen der Leitung-

sbahnen in den Pflanzen. Histologische Beitrage 3:494. 1891.

8 1. c. p. 282.

10 Pfeffer-Ewart. Tbe Physiology of Plants. 2nd revised Ed. 2:207.

1903.

11 Hartig, Th. Beitrage zur physiologischen Forst-Botanik. Aligem.

Fnrst-u. Jagd-Zeit. 1857: 281-96. 1857.

6 Wisconsin Academy of Sciences , Arts, and Letters.

On pine and larch the greatest growth had occurred at the base

of the trunks. By August 19 radial growth had ceased on

above-ground parts of broad-leaved trees, only a small amount

had occurred on the lateral roots and none on the fibrous roots.

In conifers radial growth was not entirely completed on aerial

parts and the roots were in about the same condition as those of

broad-leaved trees. In oak and maple radial growth on the

fibrous roots began about the 1st of August, in pine about the 1st

of September, in larch about mid-September. Hastings12 found

that radial growth started first back of opening terminal buds

in broad-leaved trees and proceeded basad. By the time the

five to six-year branches were producing new wood radial growth

had become general all over the trees. In case of pine radial

growth commenced on the two to three-year old portions of

branches and apparently before the buds opened. It was

thought that perhaps growth started on two-year branches

in pine because leaves are retained two years, for it was noted

that in the hemlock, where the leaves are retained six to seven

years, radial growth seemed to have started first on six-year-old

branches, while in the bald sypress radial growth started first

just back of the opening terminal buds as in broad-leaved trees.

On the other hand Knudson13 reports that radial growth begins

on young trees of the American larch in the fourth to six-year-

old branches. He holds that the cambium first gives rise to

phloem cells in spring and that wood cells are developed later

though his counts show only a few cells. The branches showing

the first radial growth were found in the middle region of the

tree. Here growth began at the apexes while in the trunk

xylem formation is said to start near the middle. Darkened

bark, owing to its heat absorbing qualities, is thought to induce

early growth.

According to Goff14 spring growth begins in many plants on

their roots. From his examinations in late March he reports

that the roots of Eibes vulgare had elongated as much as 7.5 cm.

(3 inches) before aerial growth had begun. Of the following

13 Hastings, G. T. When increase in thickness begins in our trees.

Plant World. 3:113-16. 1900. Sc. 12:585-86. 1900.

13 Knudson, L. Observations on the inception, season and duration

of cambium develonment in the American larch. Bui. Torr. Bot. Club.

40:271-93. 1913.

14 Goff, E. S. The resumption of root growth in spring. Wise. Agrl.

Expt. Sta. Ann. Rpt. 15:220-28. 1898.

Grossenbacher — Radial Growth in Trees.

7

species he says that root growth had also “ started more or less

in advance of the buds:” Picea excelsa, P. alba, P. pungens,

Pseudotsuga Douglasii, Abies concolor, Thuja occidentalis, Finns

sylvestris, Tsuga canadensis, Tamar ix amurensis , Acer sacchari-

num, Pyrus Mains, P. Communis, Prunns cerasns, P. virginiana,

Betula alba, Morns alba, Cornns stolonifer, Eleagnns hortensis,

Ribes nigrum and R. oxyacanthoides. "When these observations

are compared with those of von Mohl15 who found that, though

radial growth in conifers has practically ceased by winter and

that in deciduous trees it usually has not, it seems likely that Goff

overlooked the possibility that portions he held to be new spring

growth may have been very late growth of the preceding fall.

Hartig16 found that the roots of various forest and fruit trees

had ceased radial growth in January, as judged by the thickness

of the new ring and by the presence of starch in all of the ray

cells of the cambial region. Russow17 made similar observations

in regard to both forest and fruit trees. Hartig notes an excep¬

tion in the case of a species of willow where radial growth of the

roots had not been completed as shown by the thinness of the

ring as well as by the absence of starch in the ray cells of the

cambial region. Resa18 also made some observations which sup¬

port Goff in some cases at least. He found that the roots of

Picea and Fagus ceased growth in November and recommenced

in February and March, while in case of Aesculus Hippocastan-

um and Tilia root growth ceased in October and recommenced

in December or later. In Alnus glntinosa root growth began in

October and continued practically through the winter except

when the ground was frozen. Root growth began in late May in

Acer campestre and in June in Qnercus Robur. It is not usually

considered that such enormous variations occur in the root

growth of our trees and shrubs and for want of more detailed

information it seems necessary to admit that at least in some

16 von Mohl, H. Einige anatomische und physiologische Bemerkun-

gen uber das Holz der Baumwurzeln. Bot. Zeit. 20:225-30; 233-39;

268-78; 281-87; 289-95; 313-19; 321-27. 1862.

16 Hartig, Th. Ueber die Zeit des Zuwachses der Baume. Bot.

Zeit. 21:288-89. 1863.

17 Russow, E. uber den Inhalt der parenchymentischen Elemente der

Rinde vor und wahrend des Knospenaustriebes und Beginns der Cam-

biumthatigkeit in Stamm und Wurzel der einheimiscben Lignosen.

Stizungsber. Naturforscher-Ges. 6:386-88. 1884.

18 Resa, F. Ueber die Periode der Wurzelbildung. Inaug. Dissert.

Bonn. 1877. pp. 37.

8 Wisconsin Academy of Sciences , Arts , and Letters.

cases root growth may precede the growth of aerial parts of trees

in spring.

Schwarz19 found that radial growth may start in spring in

various parts of trees depending on the environment. In case of

a much shaded or overtopped tree it was found that radial

growth had begun at the base, while half way up the trunk the

cambium was still .dormant. In another instance 43% of the

ring had formed at the base of a tree by July 27, while 5.5 m. up

the trunk no growth had yet occurred. These irregularities are

held not to be attributable to differences in temperature occurring

at the different regions. Mechanical stimuli to be discussed

later are held to be the instigators and distributors of radial

growth.

THE RELATION OF FOOD DISTRIBUTION AND THE PRESENCE OF

ELONGATING STRUCTURES TO THE OCCURRENCE OF RADIAL

GROWTH.

It is of interest to know definitely what relation exists between

the occurrence of radial growth and elongation growth or

whether both are simply dependent upon the presence of certain

unknown amounts of elaborated and inorganic foods in connec¬

tion with the enzymes that may be involved in food transforma¬

tions and growth. The experiments of Jost20 indicate that a cas¬

ual relation exists between radial growth and some phases of elon¬

gation growth or the presence of unfolding buds, since on the re¬

moval of the buds from seedling beans radial growth practically

ceased although elongation might continue. Starch was present in

abundance and increased after the operation yet cambial activ¬

ity remained in abeyance. All the elongation buds were re¬

moved from several years growth of branches of Finns Laricio

on March 8 while the dwarf branches and their leaves were al¬

lowed to stay. The dwarf branches wrhich were nearly terminal

then developed elongation buds. By the end of May but few

tracheids had developed in the decapitated branches while in

normal branches a new layer of about twelve tracheids was pres¬

ent, and they had become lignified. A month later the mutilated

19 Schwarz, F. Physiologische Untersuchungen fiber Dickenwachstum

and Holzqualitat von Pinus silvestris. Berlin. 1899. pp. 371.

50 Jost, L. Ueber Dickenwachstum and Jahresringbildung. Bot.

Zeit. 49:485-95; 501-10; 525-31; 541-47; 557-63; 573-79; 589-96;

605-11; 625-30. 1891.

Grossenbacher — Radial Growth in Trees.

9

branches had a layer of tracheids not to exceed five or six while

a branch from which all dwarf-branches or assimilating leaves

had been removed on March 8 but on which the terminal buds

had been left, had developed a layer of eighteen to twenty

tracheids.

In another experiment dost removed buds from branches in

early May. When examined in fall it was found that at a cer¬

tain point or line in the year’s growth the radial diameter of the

tracheids was suddenly reduced and then increased again, thus

indicating the time when the buds were removed. The doubl¬

ing effect on the wood ring resulting from the removal of the

leaves at a certain time, has since been investigated by Kiihne as

noted below.

In a later paper dost21 reports some further experiments along

this line. Defoliated pine branches were found to undergo nor¬

mal radial growth provided the terminal buds are not removed,

though • they may be kept in the dark ; while when the last

grown leaves and the terminal buds wTere removed very little or

no radial growth occurred. Practically the same results were

obtained following a similar experiment with Rhododendron.

Holes wrere bored into the trunks of various trees in late Sep¬

tember and covered to prevent evaporation. By mid-October

callus formation had occurred in all but Tilia, even, though gen¬

eral growth had ceased. That is, it appears that although

cambial activity is usually started by leaf or shoot elongation

wounding may also induce it, and that not the availability of

food but a distal connection with some growing leaf-structures or

buds is necessary for the occurrence of radial growth. This

same phenomenon is also indicated by the results of an experi¬

ment with Periploca. Although this plant has bicollateral

bundles, removing a girdle of bark prevented radial growth on

the basad side of the girdle. Nordlinger22 had noted that in

case of most trees from which the branches are removed in win¬

ter practically no radial growth occurred during the following

vegetative season although in some instances slight growth re¬

sulted-

21 Jost, L. IJeber Beziehungen zwischen der Blattentwicklung und

der Gefassbildung in der Pflanze. Bot. Zeit. 51:89-138. 1893.

22 Nordinger, H. Der Holzring als Grundlage des Baumkorpers.

Stuttgart. 1871. pp. 47.

23 Vochting, H. Zur experimentellen Anatomie. Nachrichten Kgl.

Ges. Wiss. Gottingen. 1902:278-83. 1902.

10 Wisconsin Academy of Sciences, Arts, and Letters.

Vochting23 also found that decapitating herbaceous plants re¬

sulted in the cessation of radial growth of the stele though in¬

crease in diameter may result from the growth of the pith and

cortical parenchyma. After such decapitated plants were

budded cambial activity was resumed.

Reiche24 also notes regarding trees of Chili that radial growth

begins after the buds burst and that it does not occur unless bud

development precedes it.

The more detailed experiments by Lutz 25 also give support

to J ost ’s conclusions regarding the relation of growing leaves or

buds to radial growth, and they show besides that other things

being equal the distribution of food may also be a determining

factor in the occurrence of radial growth. All the buds and

leaves of six to ten-year old trees of Fagus silvatica and some of

Finns silvestris five to seventeen years of age were removed at

intervals from spring through the summer and the amounts of

reserve starch and growth were determined. The buds were re¬

moved on March 20 from a Fagus which was about a meter high.

Branches were examined for the distribution of starch and for

radial growth on June 15, July 1, 15 and 30, August 10 and 20,

on the 10th of September, October and November, as well as De¬

cember 5 and 23. The adventitious buds were removed but con¬

tinued to reappear, some large ones being removed on October

10. Only minor fluctuations in the starch content of the pith

rays, wood and bark of the branches were noted through the sum¬

mer with an almost entire disappearance of starch in December.

The branches remained healthy-looking but no radial growth re¬

sulted. Similar trees were defoliated on May 20, June 15,

July 1, 15 and 30, and August 28 respectively, and also freed of

buds at intervals during the remainder of the growing season.

Branches of these trees were also examined on the above dates.

In the tree defoliated on May 20 no starch was found in the

branches aside from traces which occurred in the pith and

broad rays during midsummer, and even that had disappeared

by August 20. Only a small amount of radial growth took place

which had all occurred by July. On October 30 the stem or

trunk was found to contain considerable starch at the ground or

24 Reiche, K. Zur Kentniss der Lebensthatigkeit einiger chilenischen

Holzgewachse. Jahrb. Wiss. Bot. 30:81-115. 1897.

85 Lutz, K. G. Beitrage zur Physiologie der Holzgewachse. Beit-

rage Wiss. Bot. 1:1-8. 1897.

Grossenbacher — Radial Growth in Trees.

11

crown in the pith, rays, and bark yet no radial growth had oc¬

curred at that point, while, 20 cm. above ground where no starch

was present, about 4% of the normal amount of radial growth

had occurred. The thickness of the new growth in the trunk

increased upward until at 75 cm. above ground a maximum of

30% of the normal amount had occurred although no starch was

present there. A little starch was present in the main root near

the crown but none occurred in the laterals and no radial growth

had occurred in them.

Corresponding results were also obtained with the other trees.

The starch content and radial growth were found to have in¬

creased in each case, until, in the tree defoliated on August 28,

the amounts of both starch and growth were normal. It should

be noted, however, in cases where defoliation induced much re¬

duction of food and growth of the trunks, that a radial growth

maximum usually occurred about 75 to 80 cm. above ground,

such as given above in detail. The year’s growth of full-leaved,

young trees was found to be in excess of that occurring in pre¬

ceding years and their starch content was very high throughout

the summer.

Five young trees of Pinus silvestris were used in similar ex¬

periments; one being defoliated on each of the following dates:

March 20, May 20, June 15, July 1, and August 30. The buds

which had been left on the tree defoliated March 20 had burst by

May 20, although the needles had not reached full size. On

July 1 and 30 some more buds had burst and begun to develop

needles. On June 15 small amounts of starch were present in

the branches. On August 20 no starch was present and only

from 4 to 20% of normal radial growth was found. On Octo¬

ber 10, when the tree was taken out traces of starch were still

present in the base or crown of the trunk but none occurred in

the roots. The roots had died and their bark had become loose

and infested with nematodes. Brown spots occurred on the

bark of the stem and the twigs were being eaten by insects. The

new growth was very irregularly distributed over the stem.

Around the circumference just above the ground growth varied

from none to 8% of the normal thickness and from this point

upward the variations were equally as marked.

In the tree defoliated on May 20 no starch was found during

the summer, yet from 10 to 60% of the normal increase in thick

12 Wisconsin Academy of Sciences , Arts , and Letters.

ness had occurred. In the remaining three trees traces of starch

were present which soon disappeared. The radial growth

ranged from 25% to normal. The tree defoliated June 15 was

dead by October and the one defoliated in August by the follow¬

ing May.

The stems of the first four and of some untreated young pines

were cut in 15 to 30 cm. pieces in October and by December the

bark on the treated- tree pieces was found to have loosened espe¬

cially where considerable radial growth had occurred. The bark

had split and was shrunken both in length and circumference;

while that on the pieces from untreated trees adhered firmly to

the wood. In December pieces were also cut from the branches

of the last treated tree, the bark of which had lost its turgidity

after the operation but regained it again. A discolored circle

was found in the cambial region. Groups of undifferentiated

wood cells had been ruptured or broken down and were discol¬

ored.

In his researches on the reserve food of trees du Sablon26

found that the carbohydrate content underwent farily definite

seasonal changes which apparently occurred irrespective of the

weather. On March 17 the roots of pear trees contained much

more sugar and very much more starch than the stems and the

total carbohydrate content of roots was also higher. In stems

of chestnut trees the carbohydrate content reached a maximum

in October and a minimum in May, while in roots the maximum

came in September and the minimum in May. In case of quince

the maximum in both root and stem was found in January with

a minimum in stems in May and in roots in June. In peach the

minimum in both root and stem came in May and the maximum

in the stem in July and in roots in November. In willow both

stem and roots were found to have a minimum of carbohydrates

in April and a maximum in October, but both the maximum and

minimum were more extreme in the roots. In the case of rasp¬

berry bushes the roots had a minimum in April and a maximum

in October, while in the biennial stems a high carbohydrate con¬

tent was maintained during the first summer with a maximum in

October, followed by a slight depression and subsequently a les¬

ser maximum in the second April. Afterwards a fairly constant

20 du Sablon, Leclerc. Recherches physiologiques sur les matieres de

reserves des arbres. Rev. Gen. Bot. 16:339-68; 386-401. 1904.

Grossenbacher — Radial Growth in Trees.

13

decrease in the carbohydrates occurred until the end of the

stem’s life.

The observations by Fabricius27 on the distribution of food in

large spruce trees throughout the entire year seems also to throw

some light on the possible relation this may have to the inception

of radial growth in spring. The first tree was cut in February.

It was 25 m. high, had 68 growth rings at the base and its low¬

est branches were 14 m. above the ground. The bark of the

stem 30 cm. above ground had considerable starch in the medul¬

lary rays, and less in the parenchyma. The older phelloderm

and ray cells contained less starch than the younger ones. Prac¬

tically the same distribution of starch obtained in the entire bark

of the trunk up to the first branches. From the branches up¬

ward the starch gradually increased to a maximum at 21 m. and

diminished again near the distal tip where but little was present.

The twenty-five outer rings in the lower part of tne trunk had

live, starch-bearing wood-rays and gum-canal cells and only the

outer half of the youngest wood-ring contained no starch. Fif¬

teen meters above ground where the stem had 36 rings only 19

contained live cells and at 18 m. about a tenth of the ray cells

were alive and starch bearing in the innermost of the 21 rings.

In the one-year shoot only about half of the pith contained starch.

The distribution of the fats was similar to that of the starch but

it was much less in amount except in the youngest twigs. The

decrease of reserve food near the distal portions was thought to

be due to a loss through respiration during winter.

The starch content of small roots was slight but usually in¬

creased with their diameter up to 1 to 2 mm. An excentric root

having 55 rings on one side and 37 on the other contained starch

in the outer 20 rings of the thicker side and in the outer 15 of the

thinner. Only the roots over 2.5 cm. in diameter contained fats.

In some eases excentric roots were found to have a difference of

as much as 50 growth rings between the broad and narrow sides,

yet the cambium on the thinner side was normal, although it was

evident that it often remained inactive during several years.

The relative amounts of starch stored on the different sides of an

excentric root was proportional to the amount of growth on any

side.

27 Fabricius, L. Untersuchungen liber den Starke-und Fettgehalt der

Fichte auf der oberbayerischen Hochebene. Naturw. Zeit. Land-u.

Forstw. 3:137-76. 1906.

14 Wisconsin Academy of Sciences , Arts , and Letters.

A tree with 82 rings at its base and 22 m. high was cut in

March. The bark was fairly rich in starch from the ground up.

The 32 outer rings of wood contained starch. At the first

branches 12 m. above ground, where the stem had thirty rings,

only the fifteen outer rings were alive and starch bearing. At a

height of 18 m. eleven or twelve of the fourteen rings present

contained starch. Considerable starch occurred in the wood at

the tree’s base and decreased rapidly upward to a minimum

about 3 m. above ground, above which it gradually increased

again to a maximum just below the branches. From this point

upward a decrease occurred which reached a second minimum

18 m. above ground, and was followed by a second increase up¬

ward to a maximum at the point where the stem had but six

wood rings. No fats could be found in the bark and very little

in the wood. Apparently fats had been changed to starch.

More starch was present in the small branches of this tree than

of the one cut in February. Both the wood and bark of the

roots contained considerable starch except the youngest phloem

cells which were devoid of it. In excentric roots the starch dis¬

tribution was similar to that found in the former tree.

Another tree which was much like the one cut in March as to

size, age, etc., was cut in late April. Its bark was rich in starch

with the exception of the phloem about 8 m. above ground where

none occurred. The reduction in the number of live, starch¬

bearing wood rings from below upward was about the same as

in the other cases. The wood rays near the cambium were de¬

void of starch. A slight amount of fat was present in the bark,

while that of the wood increased from a small amount at the

base of the tree upward to a maximum in the smallest twigs

where it exceeded the starch. In this case a starch maximum

occurred also at the base of the trunk, while in the branch bear¬

ing part of the stem the starch was evidently being dissolved

from the cambium inward and in increasing extent upward.

Fats were abundant throughout the trunks and also present in

the wood of the larger roots but absent from the bark of roots.

But very little starch was present in the wood-rays at the base

of the trunk and the season’s growth of wood was devoid of

starch, while the previous year’s growth was almost free of it in

mid-June. From this region upward starch-free peripheral

wood increased up to the first branches, where it included the

Grossenbacher — Radial Growth in Trees.

15

outer four rings. In the branch-bearing portion of the stem the

outer rings again showed some starch. All the wood was rich

in fats which usually exceeded the starch present. In the

phloem only the youngest cells had appreciable amounts of fat.

That is, in mid- July more fat and less starch is present in spruce

than in June.

Very little starch occurred in the one-year roots but it in¬

creased in amount toward the thicker roots so that in four-year

roots as much starch was present as there had been in the trees

cut before. The bark also contained much starch but very little

fat. No fat was present in the wood of the smallest roots but it

occurred in the larger ones and increased upward. The new

elongation growth of the roots and the bark on the thin ones, as

well as the young wood and phloem, were devoid of starch al¬

though considerable was present in the large roots. Fat oc¬

curred in the root wood and in occasional places in the bark.

By the last of August an additional reduction had occurred in

the fat content of the bark and the starch in the bark had also

decreased from the ground upward while nearly the entire wood

cylinder had become practically starch-free. The bark of the

larger roots contained considerable starch but it was irregular¬

ly distributed. In the youngest phloem it was absent. The

wood-rays in the larger roots and stumps had fairly large

amounts of fat present. In general it may be said that the

starch decreased in the aerial parts and increased underground

since last examined in July. The transition occurring at the

crown or stump where starch was less and fat more abundant

than earlier in the summer.

On September 25 the bark of the stem contained considerable

starch but it was present in decreasing amount from the first

branches upward to practically none in the season’s growth of

shoots. Nearly the entire wood cylinder was devoid of starch

excepting a small amount at its base or crown and in the inner

living rings. Both bark and wood were rich in fats especially

in the rays. The maximum fat content occurred about 3 m.

above ground where starch was practically absent. All except

the thin roots were comparatively rich in starch. In the wood

starch increased toward the stump. The larger roots also con¬

tained considerable fat while the small ones had none.

On October 28 the bark of the stem near the ground contained

16 Wisconsin Academy of Sciences , Arts , and Letters.

very large quantities of starch, which gradually diminished up¬

ward to the branches where it increased again but none was pres¬

ent in the season’s shoots. In the wood of the stump the starch

was also abundant especially in the rays. It decreased upward

to the branches and in the season’s shoots only a little was pres¬

ent near the pith. The fat content of the bark increased from

the ground upward but beyond the four-year-old branches there

was but little fat present. In general less fat than starch was

present in the wood of the stem but it gradually increased from

the ground up to the branches.

A marked starch increase in the wood since September was evi¬

dent while the fat content had not been correspondingly reduced,

in fact it was considerable in the branch-bearing part of the

trunk. The distribution and relative amounts of reserve food

was very similar to that found on the preceding February. It

is therefore thought evident that starch does not diminish early

in the dormant season and that it is retained as starch through¬

out winter.

The bark of the roots had an increasing amount of starch

toward the stump until a maximum was reached in roots 2 to 3

cm. in diameter after which it diminished. The wood contained

considerable starch in as many as thirty of the outer rings near

the stump and then the number of starch bearing rings decreased

peripherally as it did in the stem from the ground upward. In

an excentric root with a radius of 44 mm. on one side and of 7

mm. on the other twenty rings contained starch on the thicker

side and ten on the other. The thicker side had 70 rings and

the opposite side 20 showing that during 50 growing seasons no

radial growth had occurred on the thinner side. The roots con¬

tained considerable fat which diminished toward the stump.

In this ease as well as in the tree cut in February the young¬

est phloem and the included portions of the phloem rays besides

the outer cortex contained very little starch while that portion of

the bark between them contained much starch. Fabricius thinks

that the characteristic browning of the inner phloem so com¬

monly noted in late winter and spring, which has been attributed

to the action of atmospheric electricity by Tebeuf ,28 probably has

a relation to this distribution of reserve food in the bark.

18 Tubeuf, K. von. Beobachtung iiber elektrische Erscheinungen im

Walde.

Naturw. Zeit. Land-u. Forstw. 3:493-507. 1905.

Grosseribacher — Radial Growth in Trees.

17

From these j observations on reserve food distribution in large

trees it seems evident that most of the starch is converted to fat

during spring and early summer, and reconverted to starch

again beginning in late September, so that the smaller portion

of reserve food passes the 'winter as fat. Fischer’s29 observa¬

tions do not agree with those of Fabricius but, since the former

based practically all his conclusions on specimens from stems

ten years old or less his conclusions are not surprising.

According to Fabricius there is a general increase of starch

also in spring but it is of short duration. By April 22 it had

largely disappeared from both 'sides of the cambial region and

more especially toward the top of the tree, i. e., apparently in

proportion to cambial activity. At the same time the process

of converting the reserve starch in the older rings to fat (which

continues all summer) is1 also going on. The redeposition of re¬

serve food is begun in the bark in the form of starch. In the

wood this process does not 'begin till about the last of September

and not until October is the fat in the wood converted into

starch. The fat in the bark' is used up during summer and, from

the peripheral shoots downward, followed by a redeposition of

starch as the second growth is finished in late summer. Elonga¬

tion growth of roots is said to occur chiefly in June and July ' and

again to a slight extent in October. During those periods they

contained considerable fat which afterwards disappeared.

This series of examinations has shown 'that the fat content of

roots is practically proportional to the amount of elongation

growth in progress and that when this growth ceases very little

or no fat is present, i. e., a causal relation seems to exist between

fat content and elongation growth. It is thought that perhaps

the growing tip secrets an enzyme which is carried up the root

by the “transpiration current,” and which converts starch to

fats. After the cessation of growth the fats are again changed

to starch.

A more recent contribution to this discussion is by Preston

and Phillips,30 but it also is based chiefly on determinations made

on young trees. The study covered the period from October to

29 Fischer, A. Beitrage zur Physiologie der Holzgewachse. Jahrb.

Wiss. Bot 22:73-160. 1891.

80 Preston, J. F., and Phillips, F. J. Seasonal variation in the food

reserve of trees. Forest Quarterly 9:232-43. 1911.

2— S. A.

18 Wisconsin Academy of Sciences, Arts , and Letters.

J une and included both hard and soft wood trees. It was found

that all starch disappeared in winter from Populus deltoides,

Salix alba and Juniper us virginiana , while Quercus rubra , Ulmus

americana, Acer saccharum and Juglans nigra retained consid¬

erable starch in the wood through the winter. Tilia americana

underwent a starch reduction but retained some in the phloem,

medullary rays, and xylem, while Carya glabra lost its starch

in small stems but retained about a fourth of it in larger stems.

None of these trees except Carya showed a reduction of starch

in the roots during winter. Large amounts of sugar were found

present only in spring as the buds were unfolding. The trees

tested had a maximum fat content in late fall and a minimum in

spring. These tests seem to show that broad-leaved hard wood

trees cannot be called starch trees nor those with soft wood fat

trees, as had been done by Fischer.

Niklewski31 concluded from his study that the starch conver¬

sion in soft wood trees like Tilia, Betula, etc. is practically com¬

plete on the approach of winter, while in hardwood trees like'

Prunus and Syringa it is only partial. It was found that fats

are more abundant in winter and also that a rise in temperature

increased the amount.

According to Wotczal32 starch transformation begins in spring

in the distal parts of shoots and roots and proceeds towards the

older portions of the tree, although it starts later in roots than

in the shoots. But normally these two waves of starch trans¬

formation starting in the roots and shoots do not encounter one

another, and in this way a starch residue remains in the older

wood and in the region of the root-crown. The deposition of

starch then occurs in the reverse manner throughout the tree,

i. e. it begins in the oldest parts and around the root-crown and

proceeds wave-like toward the distal ends of the shoots and roots.

The work by Fabricius reviewed above shows that remarkable

and apparently wave-like progressive changes occur in the state

and distribution of reserve foods in trees and that maxima and

minima of the different types occur in certain parts at rather

definite periods of the seasonal history. The above cited experi-

81 Niklewski, B. Untersuchungen liber die Umwandlung einiger

stickstoffreier Reservestoffe wahrend der Winterperiode der Baume.

Beihefte Bot. Centralbl. 19 Abt. 1:68-117. 1906.

32 Wotczal, E. Die Starkeablagerung in den Holzgewachsen. Bot.

Centralbl. 41:99-100. 1890.

Grossenbacher — Radial Growth in Trees.

19

merits by Lutz show in addition that food distribution in stems

is related to the source and amount of elaborated food descend¬

ing from the leaves although such factors do not seem to pre¬

vent the eventual regional distribution so strongly brought out

by the observations of Fabricius, except in cases where the sup¬

ply is very limited and apparently all used up or deposited on

its way down the tree. At any rate, in such instances too little

reaches the lower part of the trunk and roots to permit the oc¬

currence of radial growth in those regions. Some recent defolia¬

tion experiments by Kuhns33 show that the radial growth occurr¬

ing after defoliation usually does not extend to the base of the

stem and, therefore, results in an incomplete double ring. In

this case as in those cited by Hartig, Rubner, etc., the conclusion

seems warranted that radial growth w^s omitted on the lower

part of the trunk and roots chiefly because the downward stream

of elaborated food is used up before reaching that part of the

trunk. When the growth of excentric roots and an irregular

distribution of radial growth at any given circumference of a

tree-trunk, as noted by Lutz, are considered in relation to the

occurrence of reserve food, the problem becomes more complex.

Such cases make it necessary either to assume that elaborated

food is thus irregularly distributed in a tree or else that other

factors are involved in the distribution of radial growth.

Fabricius found that food is stored in a larger number of rings

on the thicker side of an eccentric root, but that does not neces¬

sarily mean that the oldest starch-bearing rings on that side are

any older than the oldest starch-bearing ones on the thinner side

since rings are often entirely omitted on the narrower side. It

is at least possible that radial growth begins in spring in that

portion of a tree in which the greatest amount of food is stored

and in view of the fairly well established fact that growth con¬

tinues longest in fall in such regions of maximum food content

this possibility is somewhat emphasized. Perhaps it might be

of interest first to consider the causes of excentric growth as far

as they have been determined before taking up the factors which

have been advanced by several authors as the cause not only of

the distribution of reserve foods but of the general form of tree

trunks.

83 Kiilms, R. Die Verdoppelung des Jahresringes durch kiinstliche

Entlaubung. Bibiio. Bot. 70:1-53. 1910.

20 Wisconsin Academy of Sciences , Arts , and Letters.

THE CAUSES AND THE OCCURRENCE OF EXCENTRIC RADIAL GROWTH.

In a study of the distribution of excentric radial growth on

trees it is well to note that excentricity may conceivably come

about in one or more of four ways and that in a sense such an

uneven growth of a stem at any height corresponds to the wave¬

like uneven distribution at different heights of a tree. The four

ordinary ways excentric stems may be built up are (1) by the

entire omission of radial growth in a part of the circumference,

(2) by the unequal rate of growth on different sides of stems,

(3) by the entire omission of summer growth on one side and,

(4) by the omission of spring growth on a part of the cir¬

cumference and its occurrence at other places. In looking over

papers on excentric stems, etc., it is sometimes difficult to deter¬

mine to which of the four classes the case under consideration

belongs but usually that is apparent.

Gravity and other factors of the environment as w7ell as the

anatomic or physiologic characteristics of a species seem to be

the causes of excentric radial growth but as yet the matter is

not fully understood. That a difference may be found in trees

of different groups in regard to excentric growth, when subjected

to the same environment, is shown by some observations by

Nbrdlinger.34 He cites an instance in which saplings of conifers,

beech, and oak had been bent over by the heavy snows of 1868

and afterwards grew in slanting positions. Three years later

sections taken at any point of the stems showed that pine, spruce,

and larch had developed three excentric rings with the larger

radius below while on the oaks and beeches the three last rings

were thicker above. In one spruce only one very narrow ring

had been laid down on the upper side while the other rings had

been wholly omitted on that side. In both oak and beech radial

growth had been extremely slight on the under side during the

three years. This show^s that different trees subjected to the

same environment may respond differently. That is, the specific

characteristics of a plant to a certain extent determine the man¬

ner of response to the environment.

Muller’s35 observations seem to indicate that if excentric

84 l. c.

80 Muller, N. J. C. Beitrage zur Entwicklungseschichte der Baum-

krone. Bot. Untersuchungen 1:512-24. 1877. Heidelberg.

Grossenbacher — Radial Growth in Trees.

21

growth is due to the environment the branches on the upper and

lower parts of the same tree must be dominated by different

factors. On measuring the cross sections of 100 large horizontal

branches of beech trees he found that of those arising on the

stems between eight and fifteen meters above ground 36% were

epinastic, 60% hyponastic and 4% of equal radius above and

below ; of those arising between fifteen and twenty meters above

ground 36% were epi — and 39% hyponastic and 24% had equal

radii above and below ; of those taken twenty to twenty-four me¬

ters above ground 64% were epi — and 28% hyponastic, with only

7% having equal radii above and below.

This opened up a phase of the problem, which is often left out

of consideration. It shows that the branches of some trees are

chiefly hyponastic on the lower part of trunks while they may

be predominately epinastic in the upper regions. From his tab¬

ulated data the unmentioned and highly interesting fact may also

be gleaned that, of the 100 branches measured, 47 had the great¬

est diameter in the horizontal plane and only 28 had the greatest

diameter in the direction of gravity, while the other 25 weke

isodiametric. Although no special attention was directed to

these facts by Muller he apparently was fully aware of them for

he concluded that gravity is not a factor in the distribution of

excentric radial growth, but that its distribution depends upon

illumination and the relative proximity to the channels of most

direct or greatest water and food conductance. Wiesner36 who

has given this problem much attention, says that all inclined

stems of conifers are hyponastic or what he calls hypotrophic,

and that those of broad-leaved trees with little or no anisophylly

becouie first epinastic or epitrophic and eventually often become

greatly hyponastic, while species with marked anisophylly are

first hypotrophic and subsequently become epitrophic, and finally

hypotrophic again. He maintained that excentric or heterotro-

phic radial growth of a branch is due to its position both in rela¬

tion to gravity and to the axis from which it arises. On the

other hand Gabnay37 concludes that the difference in the specific

gravity of the elaborated food or of the cell content and the de¬

gree of regenerative power possessed by the different classes of

38 Wiesner, J. Ueber das ungleichseitige Dickenwachsthnm des Holz-

korpers in Polge der Lage. Ber. Deut. Bot. Ges. 10:605-10. 1892.

87 Gabnay, F. Die Excentrizitat der Baume. Just’s Bot. Jahresber.

20:100. 1894.

22 Wisconsin Academy of Sciences, Arts, and Letters.

trees are the factors determining whether excentric growth shall

be epi- or hypotrophic. The specific gravity of the elaborated

food of conifers was found appreciably greater than that of

broad-leaved trees. The regenerative power of a tree is said to

be inversely proportional to the specific gravity of its elaborated

food and it is held that the greater the regenerative power of a

tree the more epitrophic it is, while the lower its regenerative

power the more hypotrophic.

From his observations on the influence of the environment on

radial growth Kny38 concludes that the exeentricity of horizon¬

tal branches is not only a reaction to gravity but that it is also

influenced by the relative illumination, transverse bark tension,

etc., as well as by some unknown factors. In some plants the

greatest thickness of one wood ring is on the lower side of a

branch while subsequent rings may be thicker above. The

branches of most of the broad-leaved woody plants were found

to have the upper half of the wood cylinder of greater thickness

than the lower, but quite a number of exceptions were also noted,

e. g. Tilia, Cydonia, Fraxinus, Gleditschia, Corylus and Alnus.

The branches of conifers on the other hand are thickened in ex¬

cess chiefly on the lower side. In general it was found that one

type of exeentricity is characteristic of certain natural groups

of plants, but isolated exceptions were often noted indicating

that gravity plays a minor part in the distribution of radial

growth. The upper side of branches is subject to greater varia¬

tions of light, temperature and moisture than the lower and it

was thought that perhaps bark tension might be less on the up¬

per side owing to the greater distension of the bark on that side

by the variations of the temperature; yet since the results may

be just opposite in neighboring trees of different groups having

the same environment no conclusions were thought admissible.

It was observed that, owing to the fact that all leaves and buds

attached to the under side of a lateral branch develop and grow

most strongly, the axis is usually thicker on the lower side dur¬

ing the first year, while in subsequent years the branches on the

upper side of a horizontal branch grow more rapidly than those

on the lower and thus result in changing hyponastic to epinastic

branches. A case is cited where the stems of Ficus stipulata

38 Kny, L. Ueber das Dickenwachsthum des Holzkoerpers in seiner

Abhaengigkeit von aeussern Einfiuesen. pp. 136. Berlin 1882.

Grossenbacher — Radial Growth in Trees.

23

clambering up vertical walls were found to have both the wood

and phloem portions of the bundles thicker and of larger cells on

the wall than on the free side of ascending branches which is as¬

sumed to have become inherited dorsiventrality.

Kny’s study of the roots of both hyponastic and epinastic

species showed that no regularity occurs in the excentricity of

radial growth and it was thought that local pressure relations

may determine the excentricity in roots. The lateral roots were

cut from small seedlings of Tilia, Picea and Gleditschia and,

after they had begun to develop new roots, they were placed in

darkened Knop ’s solution and allowed to grow. No excentricity

resulted except in some cases where the upper radius was greater

at the origin of the root from the axis. An examination of hori¬

zontal roots which had been exposed for years, showed that their

excentricity is the same as that of the branches of the same tree.

In a more recent paper he39 came to practically the same con¬

clusions and maintained that the same factors which induce ex-

centric growth in aerial structures are in the main responsible

for their occurrence in roots. The atmospheric environment was

thought somehow to be the causal agent.

A new and rather striking application of the bark-pressure

hypothesis of Bachs and de Yries was made by Detlefsen40 in ex¬

plaining excentric radial growth. He pointed out the obvious

fact that on the concave side of a curved stem radial growth must

necessarily decrease while on the convex side it increases bark

pressure chiefly because of the effect such growth has upon lon¬

gitudinal tension of the bark. Owing to the presence of the

hard-bast fibers in the bark the reduction of the pressure on the

cambium becomes effective some distance on both sides of the

curve. The bark was usually found to be considerably thicker

on the side of a stem having the greater radius and it was fre¬

quently wrinkled or at least more rugged. He held, therefore,

that the excessive thickening in the upper angles of large lateral

roots and in the lower angle of branches is due to the reduced

bark pressure at those places following radial growth, and that

the ridges extending from such roots up the trunks are secondary

28Kny, L. tiber das Dickenwachstum des Holzkorpers der Wurzeln

in seiner Beziehung zur Lothlinie. Ber. Dent. Bot. Ges. 26:19-50. 1907.

40 Detlefsen, E. Versuche einer mechanischen Erklarung des ex-

centrisehen Diekenwachsthums verholzter Aschen und Wurzeln.

Arbeit. Bot. Inst. Wurzburg. 2:670-88. 1882.

24 Wisconsin Academy of Sciences , Arts, and Letters.

effects of the same thing. In case of branches, it was assumed

that their weight increases the longitudinal bark tension above

and reduces it underneath. Trees having one-sided tops were

said to also be affected by the increase of bark tension on the side

with fewer branches and a decrease on the top-heavy side, thus

resulting in excentric growth of the stem with the greater radius

on the side having more branches. A case was described in

which a large horizontal branch had a sharp lateral bend on the

concave side, which had resulted in a marked increase in radial

growth with only a slight increase on the lower side. On such

an assumption as this of Detlefsen it is conceivable that, after

the excentricity in the upper angles of lateral roots has once be¬

come marked and a tree has attained some age, it may become

more and more pronounced until a buttress-like structure results.

However, he failed to mention epinastic branches.

Kny41 has also noted that bending roots of herbaceous plants

and allowing them to grow in the bent position results in exces¬

sive growth of both xylem and cortex on the concave side.

According to Mer42 the two chief causes for excentric radial

growth are those affecting the manufacture of organic food and

those influencing cambial activity. The factors affecting the for¬

mer are the slope of the land, proximity to other trees, fertility

of the soil, exposure, etc., while those influencing cambial activ¬

ity are thought, to be mechanical strains due to wind, gravity,

traumatism, etc. Sloping ground is said to induce an increased

growth on the hill and a reduced growth on tne valley side.

Trunks were more commonly found excentric in thick than in

thin forest stands and the excentricity was confined chiefly to

the lower parts. When affected by the proximity of another

tree the radius toward the influencing tree was shorter. Curva¬

ture was held to be the most frequent cause of excentric growth.

Wounds were found to induce an excessive radial growth on the

opposite side of the stem ; and excentricity was found to be con¬

ducive to the occurrence of frost clefts.

41 Kny, L. Ueber den Einfluss von Zu g und Druck auf die Richtung

der Scheidewande in sich theilenden Pflanzenzellen. Jahrb. Wiss. Bot.

37:55-98. 1902.

42 Mer, E. Recherches sur les causes d’ excentricite de la moelle dans

le sapins. Rev. Eaux et Forets. Ser. 2:461-71; 523-30; 562-72. 1888.

- 3:19-27; 67-71; 119-30; 151-63; 197-217. 1889.

Grossenbacher — Radial Growth in Trees.

25

Cieslar43 performed an experiment which suggests the above

cited observations by Nbrdlinger in that he bent over the tops of

four eight-year-old spruce trees and tied them in a horizontal po¬

sition in early summer, one was bent toward each of the four

cardinal points. All ascending branches were also fastened

horizontally. The trees were cut during the second winter fol¬

lowing the beginning of the experiment and the radial growth

was found to have become greater on the upright Dasal portion

of the stems on the side of the bent-over tops. The excentricity

increased from near the ground up to a maximum beyond the

middle point of the turn where the stem was horizontal. Start¬

ing in the outer ring some distance above the inception of ex-

centric growth and extending even into the outer part of the

third ring, the wood on the side having the longer radius had a

reddish color, which also became darker upward in proportion to

the increase in the radius. That is, the rings produced the year

before the trees were bent were also affected by the bending. It

is also shown that the spring-growth of the affected rings is not

discolored in the lower part of the stained region.

Such “red-wood” as described above is very commonly present

in the under half at the base of pine and spruce branches. The

physical properties of “red wood” have been studied in some de¬

tail and its histological characteristics have also received some

attention. Although it seems not to occur in stem structure de¬

void of excentric growth, excentricity is not always accompanied

by ‘ ‘ red wood. 9 ’ The fact brought out in the above cited paper

by Cieslar that the summer wood may be affected while the

spring wood of the same ring is normal is especially noteworthy

because it shows that the factors producing “red- wood” are not

effective throughout the year.

Hartig44 made an investigation of the occurrence and distri¬

bution of “red-wood” in spruce and found that it is always pres¬

ent on trees which have excentric trunks and are located in iso¬

lated places or in thin and interrupted forest stands. Since

“red-wood” occurs in portions of trees which appear to be sub¬

ject to the greatest strains, Hartig thinks it arises in response to

the mechanical requirements of stems. He found that inclined

43 Cieslar, A. Das Rothholz der Fichte. Centbl. Gesam. Forstwesen.

22:149-65. 1896.

44 Hartig, R. Das Rothholz der Fichte. Forst. Naturw. Zeit. 5:96-109;

157-69. 1896.

26 Wisconsin Academy of Sciences , Arts, and Letters.

tree-trunks had a greater radius on the side toward which they

slant and also have 4 'red-wood’ 7 present on the side with the

longest radius. In one instance a tree on the west edge of a for¬

est and therefore having most of its branches on the west side

was found to have a longer radius as well as abundant 4 4 red¬

wood ’ ’ on the east side. In another case trees along the western

edge of a forest had the typical excessive growth and 4 4 red¬

wood ” on the east side of the trunks up to the age of about 80

to 90 years, after which the new rings showed a lesser excentric-

ity and a smaller amount of 4 4 red-wood.” The change seemed

to have resulted from the presence of a new planting on the west

side which had attained some size by that time. Hartig con¬

cluded that the mechanical or swaying effects of wind not only

causes excentric radial growth but also induces the formation of

4 4 red- wood” on the side of trunks subjected to longitudinal com¬

pression. An instance is also cited in which the leeward side of

a tree-trunk is excessively thickened from the base up but which

was devoid of 4 4 red-wood ’ ] near the ground although it was abun¬

dant farther up. A case is described where the distal part of a

young spruce stem had been bent into a complete turn and had

grown in that position during 27 years. Sections cut at various

points of the curve showed the occurrence of the greatest radial

growth and of 4 4 red- wood” on the sides where gravity and lon¬

gitudinal compression resulting from the top-weight and wind

action would require it. The excentricity of large spruce

branches and the accompanying 4 4 red- wood” was found to ex¬

tend only about four meters out from trunks.

According to Hartig 4 4 red-wood” has comparatively large

intercellular spaces and the cells seem not to be very firmly at¬

tached since they frequently fell apart in sections. The tra-

cheids are said to have especially thick walls the innermost thick¬

ening layers of which are arranged spirally.

In a more recent summary of his investigations of wood Har

tig45 claims to have proved the relative influence of gravity and

longitudinal compression in inducing the formation of 4 4 red¬

wood.” Spruce trees planted in large tubs were suspended in

an inverted position in a greenhouse and the distal part of the

stems were bent upward and allowed to grow during one sea-

45 Hartig, R. Holzuntersuchungen. Altes und Neues. Berlin. 1901.

pp. 99.

Grossenbacher — Radial Growth in Trees.

27

son. The excessive growth at the curve and the accompanying

“red-wood” was found to have developed on the under or con¬

vex side of the curve. This was assumed to indicate that grav¬

ity has more influence in the production of “red- wood” than

longitudinal compression.

Rubner46 has given us some interesting observations on ex-

centrix as well as of more irregularly distributed radial growth

of trees. He called attention to the fluted or furrowed trunks

and buttressed trunk-bases so characteristic of certain species.

He attributed the ridges to excessive and the valleys to subnor¬

mal radial growth. In Carpinus the deep, wide grooves in the

stem were found to occur at places where several compound

medullary rays are grouped together, while lesser depressions

or channels occurred along each individual compound ray, but

these lesser grooves were practically compensated for by the

greater phloem production so that the outer surface of the bark

did not show them. In portions of trunks represented by the

ridges the rays were small and it was assumed by Rubner that

the distribution of the large and small rays influences the rel¬

ative amounts of radial growth of the ridges and valleys in the

wood cylinder. While Nordlinger47 assumed that the valleys

are due to an excessive bark pressure along the large rays owing

to the development of stone cells or abnormally long phloem-ray

cells in the bark at such places. He notes the absence of marked

valleys and grooves in oaks devoid of broad rays, and that on

very large, old trees the outer rings often have the valleys be¬

tween the large rays while the ridges occur along the rays. The

armpit-like depressions below some branches, according to Rub-

ner, occur under branches whose leaves elaborate only enough

food for their own use thereby leaving the region just below7 the

branch bases insufficiently supplied, owing to the deflection the

branch-bases cause in the downward current of food in the trunk.

These depressions are said to be chiefly confined to epinastic

species. In the valleys Rubner found the wood to consist main¬

ly of thick- walled fibers and the radial arrangement of the cells

wras perfect, apparently because the valley-wood is devoid of ves-

sels. The large “false rays” present in the valleys of Carpinus

46 Rubner, K. Das Hungern des Cambiums und das Aussetzen der

Jahrringe. Naturw. Zeit. Forst-u. Landw. 8:212-62 1910.

47 Nordlinger, H. Wirkung des Rindendruckes auf die Form der

Holzringe. Centralbl. Gesam. Forstwesen. 6:407-13. 1880.

28 Wisconsin Academy of Sciences, Arts, and Letters.

were found to develop in the second and subsequent annual rings

by the elimination of most of the wood cells between adjoining

rays. Eames48 has noted a similar compounding of the simple

rays of white oaks. Rubner found that the ray cells in the val¬

ley wood are shorter than those in the ridge-wood. The wood

in valleys often showed no indication of rings because the cells

were frequently all of the summer-wood type with a reduced

radial diameter. In the deep valleys many rings were found to

converge into a homogeneous layer of small cells many of which

had brownish contents. In some cases as many as twenty-two

year’s growth had occurred on the ridges while no growth re¬

sulted in the valleys. In some such instances the cambium in

the valleys had become thick-walled and apparently lost its

power of growth and in others it had died and turned brown.

In the smaller valleys of trunks phloem production wras found

excessive while on the ridges it was only slight. Rubner also de¬

scribed instances in which no radial growth resulted on the lower

portion of tree-trunks during a number of years. He found

that long branches with sparse foliage have very irregularly dis¬

tributed radial growth, often being wholly omitted in some por¬

tions and present in others, although at times with imperfectly

differentiated cells. Similar irregularities were also noted by

Ursprung49 in branches of teak wood from the tropics ; cross sec¬

tions showed that in some growing seasons the cambium had

been active in only a part of the circumference.

The work reviewed above shows that several types of excen-

tric radial growth occur both in horizontal and upright struc¬

tures and that some of them are apparently due to differences in

bark pressure and to an excentric distribution of the transpira-

iton current and metabolized food, while in others the cause of

the excentricity is not shown. For instance these authors have

not determined why radial growth should be distributed in scat¬

tered patches on branches or tree-trunks which have an inade¬

quate supply of food or why fluted trunks and buttressed stumps

should occur, although Detlefson made some interesting sugges¬

tions regarding the latter. Rubner has shown that radial

growth is very slight in the valleys or grooves occurring in the

48 Eames, A. J. On the origin of the broad ray in Quercus. Bot. Gaz.

49:161-66. 1910.

48 Ursprung, A. Zur Periodizitat des Dickenwachstums in den

Tropen. Bot. Zeit. 62: Abt. 1:189-210. 1904.

Grossenbacher — Radial Growth in Trees.

29

tranks of Carpinus, etc., and that the wood of these valleys con¬

tains the large aggregate rays while that in the ridges has simple

ones. That the presence of the aggregate rays has induced the

valleys by their early cessation of growth as Sorauer50 held does

not necessarily follow, though it may be true, as it is more re¬

cently implied by Bailey51 and others. In. a number of recent

papers written by Jeffrey’s students52 it is maintained that the

different types of rays and their method of development are of

great phylogenetic significance in showing the paths of evolu¬

tionary development. Yet in the above cited paper by Bailey

it is also noted that changed nutrition may markedly modify the

rays and their distribution.

Some of Kny’s53 results obtained in his experiments seem to

indicate that the pressure under which rays differentiate in the

cambial zone has much to do in determining their sizei He

found on applying a pinch-cock to twigs of Salix and Aesculus

Hippo cast anum in spring that not only was radial growth almost

entirely inhibited on the compressed sides but that the ray cells

were broader in tangential direction and that in some cases a

doubling of the typically simple rays had occurred in both trees.

In the above cited paper on the causes of excentric growth Mer

also calls attention to the increase of radial growth on trunks

opposite a wound. This observation of Mer’s is of interest here

chiefly because the occurrence of traumatic rays54 in wood pro-

eo Sorauer, P. Handbuch der Pflanzenkrankheiten. Zweite Auflage.

1:537. 1886.

61 Bailey, I. W. The relation of the leaf-trace to the formation of

compound rays in the lower Dicotyledons. Ann. Bot. 25:225-41. 1911.

52 Bailey, I. W. Reversionary character of traumatic oak woods.

Bot. Gaz. 50:374-80. 1910.

Eames, A. J. On the origin of the herbaceous type in the Angio-

sperms. Ann. Bot. 25:215-24. 1911.

Thompson, W. P. On the origin of the multiseriate ray of the Dico¬

tyledons. Ann. Bot. 25:1005-14. 1911.

Holden, R. Reduction and reversion in the North American Sali-

cales. Ann. Bot. 26:165-73. 1912.

Bailey, I. W. The evolutionary history of the .foliar ray in the wood

of the Dicotyledons, and its phylogenetic significance. Ann. Bot.

26:647-61. 1912.

53 Kny, L. Ueber den Einfluss von Zug und Druck auf die Reichtung

der Scheidewande in sichtheilenden Pflanzenzellen. Jahrb. Wiss. Bot.

37:55-98. 1902.

64 Jeffrey, E. C. Traumatic ray-tracheids in Cunninghamia sinensis.

Ann. Bot. 22:593-602. 1908.

Bailey, I. W. Reversionary characters of traumatic oak woods. Bot.

Gaz. 50:374-80. 1910.

30 Wisconsin Academy of Sciences , Arts , and Letters.

duced on the side of a stem opposite a wound is assumed to have

phylogenetic significance.

According to Groom55 the evolution of the rays in Quercus is

not as simple as presented by Eames, Bailey, Thompson and

others for he found cases where the primary rays seemed to

branch like those of beech described by Jost56 as well as others

where the aggregations occurred in the manner described in the

above cited papers. Groom is inclined to the view that ray de¬

velopment and architecture is based on physiological rather than

on phylogenetic factors and that it is impossible at present to

decide whether the narrow or the broad-rayed type is the more

primitive.

It is also worth noting that, although Nordlinger57 found the

valleys originating along the groups of broad rays and that oak3

without the broad rays are devoid of valleys, in case of very

large old trees the ridges were often found to occur along the

broad rays, while valleys were present between them, i. e. just

the reverse of the conditions obtaining in younger specimens.

Perhaps it might prove worth while to find out whether the

occurrence of valleys and ridges in such trees is due to differ¬

ences between the rate of growth in the wood and in the rays

rather than being due to an early cessation of ray growth as

Sorauer had assumed. In case the formation and radial elonga¬

tion of ray cells were very slow as compared to the radial in¬

crease in the wood cylinder in general, it is conceivable that the

solid broad rays may have a dominating influence and retard

radial growth on both sides of them because of the firm attach¬

ment between the raj^s and the surrounding tissues. If the claim

made by Klebs57 that the presence of large quantities of elab¬

orated food retards radial growth should prove correct and since

these large rays are the storage reservoirs for elaborated foods

it would also be understandable how they might be comparative¬

ly slow growing in youth and comparatively more rapid in old

age, when radial growth has become slow.

The conspicuous ridges on the lower part of trunks correspond

56 Groom, P. The evolution of the annual ring and medullary ray in

Quercus. Ann. Bot. 25:983-1003. 1911.

66 Jost, L. Ueher einige Eigenthiimlichkeiten des Cambiums der

Baume. Bot. Zeit. 59:1-24. 1901.

47 Nordlinger, H. Wirkung des Rindendruckes auf die Form der

Holzringe. Centbl. Gesam. Forstwesen. 6:407-13. 1880.

67 1. c.

Grossenbacher — Radial Growth in Trees.

31

with the occurrence of the upper lateral roots. In trees like the

elms, ironwoods, and oaks the excessive thickening in the upper

angle primary roots make with trunks are often exaggerated into

buttress-like enlargements which are continued as ridge-like pro¬

longations extending some distance up the trunks. According

to Detlefsen58 the excessive radial growth in the upper angle of

lateral roots and in the lower angle which large branches make

with the trunks is chiefly due to a continued decrease of the bark

pressure at these places which results from radial growth. This

hypothetical explanation, however, requires an experimental

basis. The fact that the bark at these places is often cleft or

ruptured rather shows that radial bark pressure, at least, occurs

there. The pressure exerted against the bark by the growing

wood is not only sufficient to bring about tension at the root and

branch ridges but tension of sufficient magnitude to rupture the

bark in many instances. The experiments by Vochting59 in

which the distal tips were cut from Helianthus and other plants

with the result that the stems became somewhat fleshy and in

some cases rib-like thickenings developed over the leaf traces and

ran some distance down the stem, can scarcely be said to apply

owing to the fact that in Vnch ting’s experiments the excessive

thickening was chiefly due to increase in the pith and cortical

parenchyma instead of radial growth of the stele.

It has been suggested or inferred by some of the above as well

as by other writers that greater cambial activity occurs in the

upper angle of roots at their origin from the stump than takes

place in the lower angle, because the downward current of meta¬

bolized food is checked and accumulates more or less in the up¬

per angle. The lower angle of the root is said to be more indi¬

rectly and, therefore, more sparsely supplied with food and for

that reason one sided radial growth results. An additional

factor, which contributes to this excentricity, is doubtless the

pressure of the tree’s weight on the cambium of the underside

and another may be the reduced longitudinal bark tension sug¬

gested by Detlefsen. Even in case of a tree with a deeply pene¬

trating tap root a very marked radial increase on the lower side

of large primary laterals would tend to elevate the entire tree,

and a tree without a tap root must be carried chiefly by the large

68 1. c.

69 1. c.

32 Wisconsin Academy of Sciences, Arts, and Letters.

primary laterals and therefore exerts great pressure on the

cambium as Detlefsen60 maintained.

According to another group of investigators to be cited in the

discussion on the distribution of radial growth, excentric growth

is not due to an independent distribution of metabolized food and

the other factors commonly assumed to be effective. Both food

and growth are held to be distributed by the mechanical effects

of the environment in conjunction with the weight effects of the

structure in question or by the rate and path of the transpiration

current.

THE GENERAL FORM OF TREE-TRUNKS AND THE DISTRIBUTION OF

RADIAL GROWTH.

The distribution of radial growth on trees determines the form

of the stem and therefore its value as timber. Owing to the

economic importance of the shape of tree-trunks to the lumber¬

ing industry foresters studied the distribution of radial growth

and its relation to the environment very extensively and have

collected many valuable data. Since the stem of a tree grown

in a fairly dense and uniform forest stand is relatively longer

and less tapering toward its upper end, free of branches and

therefore of more lumbering value than one grown in the open,

the differences in the environment of the two types have re¬

ceived much attention.

Nordlinger61 noted that the yearly increase in thickness on the

branchless and branched parts of stems grown in a forest dif¬

fered from each other. The annual distribution of radial growth

on the branch -bearing portion in a forest stand was found to be

similar to that on the entire trunk of a free-standing tree, which

bears branches nearly to its base. The thickness of the wood

rings in the branch-bearing part of stems was found to decrease

from the base upward. On the branchless portion of trunks in

dense forest stands the thickness of the recent rings was noticed

to have decreased from the branches downward although in some

cases the thickness of the new yearly growth remained practically

constant at the base of trunks. He thought that the presence

of elaborated food was not the only requisite for the occurrence

00 1. c.

61 1. c.

Grossenbacher — Radial Growth in Trees . 33

of radial growth in any particular region of a tree-trunk for the

reason that the radial growth maxima in dense stands move up¬

ward more rapidly than would be demanded by the reduction in

metabolized food.

Sanio62 noted that in case of a dwarfed fourteen-year-old sap¬

ling of Fraxinus excelsior growing in a swamp the spring wood

was for the most part very thin and usually had but a single

row of vessels while in some parts of the stem the rings were de¬

void of vessels. He thought it likely that spring growth had

been wholly omitted at such places and that the ring there con¬

tained only summer-growth wood.

R. Hartig63 has probably published more on the general dis¬

tribution of radial growth than any other investigator. From

a study of overtopped pines and spruces between 20 to 30 years

old, he found that the rings became thinner from the branched

top downward and that in some cases as many as seven rings had

been entirely omitted on the lower part of stems. When rings

had been omitted during a series of years the lower edges of

the new rings or wood-sheaths were found to have receded farther

from the base each year. In another paper he64 called attention

to the fact that in overtopped trees a reduction occurs in the

yearly amount of wood produced from the branches downward.

In general a stem is said to have three more or less distinct

growth regions in each of which a typical distribution occurs.65

In the main axis of the branched top the cross sectional area of

the growth rings is said to increase from above downward. The

rings on the branchless shaft also increased in thickness from

the branches downward in trees having a well developed top,

but as stated above, the reverse was found true of a dominated

tree with a small top.

A more detailed study of the distribution of radial growth was

carried out by Hammerle66 in connection with his observations

66 Hammerle, J. Zur Organization von Acer Pseudoplatanus. Biblio.

Bot. 50:1-101. 1900.

82 Sanio, K. Verleichende Untersuchungen iiber die Znsammenset-

zung des Holzkorpers. Bot. Zeit. 21:391-99. 1863.

83 Hartig, R. Das Aussetzen der Jahresringe bei unterdriickten Stam-

men. Zeit. Forst.-u. Jagdwesen. 1:471-76. 1869.

84 Hartig, R. Zur Lehre vom Dickenwachsthum der Waldbaume.

Bot. Zeit. 28:505-13; 521-29. 1870.

85 Hartig, R. Ueber den Entwicklungsgang der Fichte im Geschlos-

senen Be-stande nach hohe, Form und Inhalt. Forst. Naturwiss. Zeit.

1:169-85. 1892.

3— S. A.

34 Wisconsin Academy of Sciences , Arts , and Letters.

on the elongation growth of young maple trees. He found that

the greatest thickness of each ring normally occurred in the

hypocotyledonary or crown region of young trees. The second

ring of the branches was thicker toward the end than in the mid¬

dle but subsequent rings decreased regularly toward the distal

end. The third ring of a rather dwarfed, overtopped specimen

had its greatest thickness in the three-year-old branches and di¬

minished toward the base until at the height to which the tree

had grown by the end of its first year, the ring was almost invis¬

ible ; at the hypocotyl or crown region and at least as far as 19

cm. downward on the roots no growth at all had occurred during

the third year. The bark in all cases was thickest at the hy¬

pocotyl or crown region.

From the papers cited in this section as well as from others

noted elsewhere it is very evident that the distribution of radial

growth is at least quite strongly influenced if not entirely deter¬

mined by the environment and it will be interesting to examine

some of the papers in which the factors that have been advanced

as being the regulators of this distribution are discussed.

The publication of Schwendener ’s67 epoch-making paper on

the mechanical principles underlying the structure of Mono¬

cotyledons gave a view of plant anatomy from a new angle and

still exerts a marked influence on both physiology and anatomy.

Many measurements and calculations obtained from typical

Monocotyledons are presented in this paper in support of the

hypothesis that plant structures take on forms and have the sup¬

porting tissues distributed in them in such a fashion as to meet

the mechanical requirement necessary to make such structures

most efficient in carrying their own weight as well as in resisting

injurious bending by the wind, etc. In replying to some severe

criticism of this paper he68 admitted that many inaccuracies oc¬

cur in the calculations but maintained that on the whole it is

correct. The general principle developed in the first paper is

here also reinforced in its application to Dicotyledons but in a

less thoroughgoing way. It was noted that radial growth in a

tree-trunk seems to be distributed in a manner so as to meet the

87 Schwendener, S. Das mechanische Princip im anatomischen Bau der

Monocotylen mit vergleichenden Ausblicken anf die ubrigen Pflanzen-

klassen. pp. 179. 1874.

88 Schwendener, S. Zur Lehre von der Festigkeit der Gewachse.

Sitznngsher. K. Preuss. Akad. Wiss. Berlin. 1884:1045-70. 1884.

Grossenbacher — Radial Growth in Trees.

35

mechanical needs in supporting the top in its environment. The

general form of trunks was found to conform more or less com¬

pletely with shafts constructed to be of equal endurance through¬

out and capable of supporting a given load (top) and wind-pres¬

sure. It is said that owing to this fact a tree trunk grown in the

open and therefore bearing branches nearly to the ground is

thicker at the base than one grown in a forest and crowded by

other trees.

Some years later Metzger69 published some results and obser¬

vations from which the striking conclusion is drawn that light,

warmth, moisture and food enable a tree to grow but that the

wind determines how it shall grow. He points out the self-evi¬

dent but none the less interesting fact that a tree-trunk must not

only carry its own weight and that of the branched top but also

resist the wind action as it shifts the center of gravity while

swaying to and fro. The tree stems are said to be the pillars of

the forest and in order that the forest exist they must be both

rigid and at the same time elastic enough to withstand strong

winds. This is illustrated by him by imagining a wooden shaft

firmly fixed in a horizontal position at one end and weighted at

the other, thus resulting in the greatest strain at the place of at¬

tachment. If such a shaft is to be equally liable to break at any

point of its entire length its cross sectional area must decrease

from the point of support to the application of the weight or

force in accordance with the physical laws involved, and the most

economical use of the material of the shaft would require such

a construction. By making numerous measurements and calcu¬

lations it was found that the proportional thickness and form of

tree-trunks below the branch-bearing tops was practically that

required of the shaft described above, except that most of them

are enlarged at the base or root-crown beyond the hypothetical

requirements. It is noted that tap-rooted trees in deep soil are

devoid of the excessive basal enlargement, and it is therefore

thought that the enlargement is only a result of developing an

adequate root anchorage for the tree. That portion of the stem

in the branching top was also found to conform to such a shaft.

In case of horizontal branches it is held that their own weight

overbalances wind action as a formative factor, while in upright

69 Metzger, A. Der Wind als massgebender Faktor fur das Wach-

sthum der Baume. Miindener Forst. Hefts. 3:35-86. 1893.

36 Wisconsin Academy of Sciences , Arts , and Letters.

branches like in trunks wind effects predominate over the weight

of the structures themselves as formative stimuli. Branches in

positions intermediate between these two extremes are said to be

correspondingly influenced by the two factors. Since conifers

of various sizes were found to conform very closely to the hy¬

pothetical requirements, Metzger thought it logical to assume

that wind and the weight of the supported structures themselves

are the factors instrumental in shaping tree-trunks or distribut¬

ing radial growth on them. When the lower branches of a free¬

standing tree were removed, it was found that the annual

growths on the lower portion of the trunk were reduced in cross-

sectional area in very nearly the proportion required by the

hypothetical considerations of the upward movement of the

point of greatest stress. When a free-standing tree is encom¬

passed by young trees radial growth of its trunk decreases from

above downward as required by this hypothesis. When forest

trees are left free-standing by the removal of surrounding trees

radial growth is found to increase on their trunks from above

downward and to decrease below normal on the upper part of

the stems. In conformity also with the above hypothetical re¬

quirements the tall or over-topping trees in a forest of mixed

sizes undergo most radial growth on the lower parts of the trunks

while the overtopped trees grow more on the upper part of

trunks.

Although these conclusions were based on data, which were

obtained from spruce, Metzger70 thinks them applicable to the

distribution of radial growth of trees in general. According to

him the wind, acting as a stimulus through its mechanical effects

upon trees, also regulates in a general way, the distribution of

the elaborated food as well as that of radial and elongation

growth in accordance with the relation of the form of the top,

etc. to vdnd-exposure. It is said that during the first and sec¬

ond year after the thinning of a forest most of the available food

is used up in increasing radial growth on the lower part of the

tranks so as to increase the wind resisting power of the suddenly

exposed trees, but afterwards elongation growth proceeds rap¬

idly. In some cases of this kind it is held that the top may be

70 Metzger, A. Studien iiber den Aufbau der Waldbaume und

Bestande nach statischen Gesetzen. Miindener Forstl. Hefte. 5:61-74.

1894. Miindener Forstl. Hefte. 6:94-119. 1894.

Grossenbacher — Radial Growth in Trees.

37

deprived of marked radial as well as elongation growth for sev¬

eral years, and the long-continued scarcity of food in the upper

part of the top is said often to result in the dying back of the

upper branches and thus gives rise to stag-horn effects. The

length of time required for adjustment to the new environment

is said to depend upon the extent of a tree’s leaf surface. The

sprouts, which often arise on long bare trunks, are thought to

be induced by the swaying action of the wind thus tending to

develop a lower head.

An enormous amount of data and calculations on the relation

of the environment to radial growth and its distribution was also

callected by Schwarz71 and published as a monograph which in

addition contains many very important observations on the life

and seasonal historj^ of Pinus silvestris. It is noted that yearly

radial growth as measured by the area of its cross section in¬

creases in trees until the age of about 20 to 30 years is reached,

but under very favorable environmental conditions its growth

may increase to the age of 100 years. His general conclusions

regarding the wind in its relation to the distribution of radial

growth are practically the same as those put forth by Schwen-

dener and Metzger. Some instances are cited where the tops of

trees had been broken off when about 30 years old and which had

since grown about 60 years with lateral branches diverted to

function as the main axis. In the region of curvature of the

branch which assumed the functions of the main axis excentric-

ity became very marked, with the greater radius on the under

side. It is thought that the excessive pressure or weight on the

under side was the stimulus to increased radial growth on that

side. In one case, in which the curvature induced had been such

as to exert the greater pressure on the upper side in one place,

it was found that this upper side had the greater radius. Many

measurements on vertical stems also showed a greater radius on

the leeward side in regard to the prevailing wind. By tying a

young pine tree in a bent position excessive growth resulted on

the compressed side, i. e. it seems that a fixed, bent position ex¬

erts the same influences on radial growth as the discontinuous

pressure due to wind swaying. Other measurements on slightly

inclined trees also showed a greater radius on the side toward

which the trees inclined. It is held that relative amounts of

1. c.

71

38 Wisconsin Academy of Sciences, Arts, and Letters.

elaborated foods present in different regions of trunks is not

primarily responsible for the distribution of radial growth, for

on such an assumption the greatest growth would always occur

on the stem just below the branches, while as a matter of fact it

usually occurs within two meters of the ground. In fact it is

claimed that both the distribution of metabolized food and radial

growth are regulated by the wind-pressure-and-weight stimuli.

The wind effects are thought to induce the transfer of most of

the food elaborated in the leaves of a recently isolated tree to the

lower part of the trunk where increased radial growth is caused

by the increase of the mechanical wind-stimulation. Attention

is called to the fact that in case of excentric annual rings the ex-

centrieity is chiefly due to an excessive production of the so-

called summer wood, thus upholding the view that swaying and

weight stimuli are especially effective during the latter part of

the period of radial growth. The data seemed also to show that

after trees with excentric rings are perhaps about 73 years old

or have begun to decline in their rate of growth the new rings

decrease markedly in excentricity and in conformity with that

it is noted that late season growth is less in trees which have

reached the age of decline in growth rate.

Schweinfurth72 reported that about the Red Sea tree trunks

all have a greater radius on the south side owing to the occur¬

rence there of a continued and strong north wind during the

summer. The presence of reduced branches on the north side is

thought to have caused the reduced growth on that side.

A more detailed application to Schwendener ’s mechanical

principles of plant structure to excentric radial growth in

branches was made by Ursprung.73 He maintained that the dis¬

tribution of radial growth of both stem and branches is deter¬

mined by the compression-strain stimulus resulting from the

weight of the structure and the action of the wind. Non-verti¬

cal stems and branches were usually found to have an elliptical

cross section with the longer diameter in the direction of gravity.

This is said to increase the carrying capacity of the wood be¬

cause the force required to bend such a branch in a vertical plane

is proportional to the third power of the vertical diameter and to

72 Schweinfurth. Sitzungsber. Ges. Naturfor. Freunde. Berlin 1887.

p. 4.

73 Ursprung, A. Beitrag zur Erklarung des excentrischen Dicken-

wachstum. Ber. Deut. Bot. Ges. 19:313-26. 1901.

Grossenhacher — Radial Growth in Trees.

39

only the first power of its horizontal diameter. He also con¬

cluded that vertical stems may become excentric owing to one¬

sided action of wind but that the effect on some trees might be

different on account of variations in the shape and the conse¬

quent distribution of the weight of the top. The crooks in a

tree trunk are assumed also to be gradually eliminated by the

distribution of the radial growth in response to strain stimuli.

The same laws are thought to apply to the radial growth in roots

but because of the variation in the environing soil they are not

always so regularly effective.

Vochting74 cut the tips from some potted one-year-old savoy

plants and placed them with their pots in a horizontal position.

He attached weights to some near their decapitated tips and al¬

lowed them to vegetate during some months. The vertical diam¬

eter of the stems was markedly increased in the regions of great¬

est strain while the stems of the check plants retained their

cylindrical forms.

The far-reaching applicability of this wind-gravity hpyothe-

sis originating with Schwendener and elaborated by Metzger and

others, according to which tree-trunks and other stem structures

have a form required of a shaft of equal endurance throughout,

has recently been questioned by Jaccard.75 He holds that the

hypothesis is untenable because measurements and calculations

made by him on a number of spruce trees resulted in a noncon¬

formity of the hypothetical and actual forms of their trunks.

It was found that the portions of the trunks beginning with 5

m. above ground and extending to about 9 m. above ground were

practically of the form and dimensions required of such a shaft

but above and below that region the trunks were thicker than

required by the laws of mechanics. In one instance described

in detail, however, the trunk of a spruce practically conformed

to the required hypothetical shaft.

Although much more frequent strong winds are said to occur

in western Switzerland the trees there were not found to differ

appreciably from those of eastern Switzerland where strong

winds are few. Jaccard maintained that during the growing sea¬

son the wind is too spasmodic to be a factor in the distribution

74 1. c.

76 Jaccard, P. Eine neue Auffassung liber die Ursachen des Dieken-

wachstums. Naturw. Zeit. Forst-u. Landwirts. 11:241-79. 1913.

40 Wisconsin Academy of Sciences, Arts, and Letters.

of radial growth, and besides, he holds that the distribution of

growth on a tree-trunk having concentric rings could not

conceivably be dependent upon wind action. From the meas¬

urements and calculations it is concluded, however, that tree-

trunks are shafts of equal water conductance throughout. From

insufficient data and non-convincing arguments it is concluded

tion mentioned above, though larger than necessary for the wind-

that the diameter of tree trunks above and below the 5 to 9 m. por¬

tion mentioned above, though larger than necessary for the wind-

gravity hypothesis are of just the size required of a shaft of

equal water conductance throughout. The morph'ogenic power

of the water current is thought to be proportional to the rate of

metabolism and transpiration. The rate of cambial division is

held to depend upon and be controlled by turgidity, and the in¬

fluence of the environment is thought to affect radial growth

chiefly through the transpiration stream. In the calculation up¬

on which this hypothesis is founded it was assumed that the

water conduction is confined to the outermost ring or wood

sheath.

This hypothesis has some defects in common with the one it is

supposed to supplant in that the distribution of radial growth is

assumed to be controlled chiefly by one factor, other factors be¬

ing effective only in so far as the basic one is influenced. Jac-

card has many difficult problems to solve before his hypothesis

to account for the actual distribution of radial growth in trees

can be considered a theory. The relation of the first radial

growth and its distribution in trees to the transpiration stream

in cases where such growth precedes actual unfolding of the

leaves will need to be explained in the promised detailed study

he is to publish in a future paper. Nor is it permissible to as¬

sume as a fact that the water current is confined to the outer¬

most ring of wood, especially when it is recalled that in certain

portions of trunks radial growth may be wholly omitted during

a number of successive years, and that many cases of girdling are

also on record in which trees operated on vegetated and fruited

normally during several years.

Wieler76 concluded that practically all water is conducted in

76 Wieler, A. Ueber den Antheil des secundaren Holzes der dicotyle-

donen Gewachse an der Saftleitung nnd iiber die Bedeutung der Anas-

tomosen fiir die Wasser-versorgung der transpirirenden Flachen. Jahrb.

Wiss. Bot. 19: 82-137. 1888.

Grossenbacher — Radial Growth in Trees.

41

the last ring but in a more recent study Jahn77 made it appear

that the entire alburnum may be more or less active in water

conduction although perhaps as much as half or more of the

water is thought to be carried up the last ring.

Some evidence of the fact that wind is both a formative and

a limiting factor in plant growth is afforded by several scat¬

tered papers on the influence of wind on vegetation, a few of

which might be briefly noted in this connection.

While making an experimental study of the effects of wind

on vegetation Bernbeck78 obtained some interesting results.

He found that both shoots and leaves of plants subjected to

wind of 14 m. or less per second were injured in proportion to

the amount of swaying and bending induced and that even del¬

icate leaves of shade plants are not injured by the wind if they

are firmly held to prevent swaying or bending during the ex¬

posure. It was found that the production of organic food was

reduced in leaves exposed to wind as . compared to that accum¬

ulating in protected leaves.

Gilchrist79 reported that potted plants of Helia/nthus annuus

subjected to artificial wind swaying and rocking did not grow

as tall as the checks while the diameter of their stems exceeded

that of the check plants. Some more recent observations by

Cavara80 show a similar effect of wind exposure on the struc¬

ture of Iresine, Coleus, Aster, Zinnia, and Sempervivum.

Esbjerg81 found that protecting various herbaceous plants

from strong winds by means of screens resulted in an increased

yield. An increase of 16 to 31% above that of the checks was

secured in the yield of grain from rye; the yield of ruta-baga

roots was increased from 7 to 17% and of mangels from 3 to

18%, while clovers and grasses showed a gain of from 4 to 23%

as a result of wind protection.

77 Jahn, E. Holz nnd Mark an den Grenzen der Jahrestriebe. Bot.

Centbl. 59:257-67; 321-29; 356-62. 1894.

78 Bernbeck, O. Der Wind als pflanzen-pathologischer Faktor. In¬

augural Dissert. Bonn. 1907. pp. 116.

79 Gilchrist, M. Effect of swaying by the wind on the formation of

mechanical tissue. Report Mich. Acad. Sc. 10:45. 1908.

80 Cavara, F. Some investigations on the action of wind on plant

growth. Expt. Sta. Record. 25:224-25. 1912.

81 Esbjerg, N. Experiments with windbreaks. Expt. Sta. Record

23:435. 1910.

42 Wisconsin Academy of Sciences , Arts , and Letters.

Similar facts are also reported by Waldron82 from North

Dakota. While from Porto Rico83 we learn that the northeast

wind prevailing there causes citrus trees to grow slowly and

one-sided in unprotected places; the bark looks dead and the

new shoots are variously twisted. A case is cited where two

similarly planted citrus groves are located across the road from

each other but one is protected by a windbreak while the other

is fully exposed. The trees had all been set three years and

were bearing in the protected grove while in the exposed one

they looked as though they “had just been set.” Wind-exposed

trees were also found heavily infested by scale-insects while the

protected ones were practically free from the pest.

In a very recent paper84 it is stated that the wind induces

dwarfing and the rosette habit, although the structural modifi¬

cations are attributed to excessive transpiration.

A like conclusion was recently also drawn by Choux.85 He

found that the stems of Neptunia prostrata and of Ipomea rep-

tans grown during the tropical dry season were not only smaller

but that their vascular systems were much more strongly devel¬

oped than in those produced during the wet season. Starch was

abundant in the dry season plants and practically absent from

those grown in the wet season.

The hypothesis advanced by Schwendener and subsequently

elaborated by Metzger and Schwarz and the more recent one by

Jaccard are so simple and imbued with such insidious directness

that they are fascinating although not wholly convincing. Af¬

ter making a brief survey of the observations and experiments

by Jost, Lutz, Fabricius, Rubner, etc., it seems as though the

occurrence and distribution of radial growth could not be de¬

pendent on a single factor. It appears for instance that the

distribution of elaborated food must in part at least depend upon

its place of manufacture and on the channels of its transport,

especially when the amount available is somewhat below the

sa Waldron, C. B. Windbreaks and hedges. N. Dk. Agrl. Expt. Sta.

Bui. 88. 1910. pp. 11.

ss Tower, W. Y. Insects injurious to citrus fruits and methods for

combating them. Porto Rico Agrl. Expt. Sta. Bui. 10:16-20; 35. 1911.

84 Kroll, G. H. Wind und Pflanzenwelt. Beihefte Bot. Centralbl.

30 Abt. 1:122-40. 1913.

85 Choux, P. De Pinfluence de l’humidite et de la secheresse sur la

structure anatomique de deux plantes tropicales. Rev. Gen. Bot. 25:

153-72. 1913.

Grossenbacher — Radial Growth in Trees.

43

actual needs. On the other hand from the work of both Jost

and Lutz it is also evident that the presence of food, transpira¬

tion current and suitable environment alone do not result in

radial growth when no developing buds or shoots are present;

i. e., cambial activity seems somehow to be dependent upon

elongation growth or some enzyme activated or produced by it.

The determinations by Fabricius, however, have made it ap¬

parent that the distribution of reserve food in tree-trunks seems

to be in accordance with some unknown law, which brings about

maxima and minima of food storage in more or less definitely

alternating regions. The marked differences in the amounts of

reserve food in the regions of maxima and minima could not be

attributed to differences in the storage capacity of the regions

for such differences would have been noted, nor to the distribu¬

tion of the branches because the wave-like succession of maxima

and minima also occurred and it was usually most marked on

the branchless portion of trunks. There is some indirect evi¬

dence to be had from the cited papers which tends to show that

the places of the inception and longest duration of radial growth

in a general way are the places of maximum food storage, and

therefore gives support to Mer’s86 contention to the effect that

radial growth begins first where most food is stored and is most

active and persists longest in such regions. The Schwendener-

Metzger-Schwarz hypothesis suggests another way out of the

difficulty by its assumption that wind action is responsible for

the distribution of both metabolized food and radial growth.

But we cannot admit the far-reaching claim of these investiga¬

tors that wind and gravity are the only formative factors con¬

cerned in the distribution of radial growth especially since light

and transpiration have been shown to be powerful formative

agents.

OBSERVATIONS ON THE DISTRIBUTION OF LATE RADIAL GROWTH ON

FRUIT TREES.

While studying crown-rot of fruit trees during a series of

years, I found that the initial bark injuries which afterwards

result in the disease usually occurred in places at the base of

86 Mer, E. Sur les causes de variation de la densite’ des bois. Bui.

Soc. Bot. France. 39: 95-105. 1892.

44 Wisconsin Academy of Sciences , Arts , and Letters .

tree trunks where radial growth continues late in fall. The

observations made to determine the distribution of late radial

growth showed that it is very irregularly distributed, yet that

when it occurs it is confined to certain parts of trees. Crane-

field87 has called attention to the general variation of radial

growth in branches. After two seasons observations he con¬

cluded ‘ ‘ that a wide difference existed between trees of the same

variety, age and external appearance, and that the difference

was often greater between different branches of one tree than

between different trees.” In 1899 he found that the bark

peeled readily on all branches of apple, pear, plum and cherry

as late as August 15, and after that date the bark still peeled

easily for some time on the larger branches. In 1900 the bark

of branches over 1 cm. in diameter slipped easily enough to make

whistles as late as September 15, while two weeks later it would

not peel from any of the branches.

Although observations like these of Cranefield show that

marked variations may occur in the distribution of the last ra¬

dial growth, it is apparent that its actual variation can only be

determined by much more detailed examinations at numerous

points not only of any one tree but of any one branch. Some

of the above cited observations on the general distribution of

radial growth and more especially those on excentric growth also

suggest the inference that late growth is often very irregularly

distributed and that it is perhaps frequently confined to regions

of trunks and branches where excentric growth occurs. In a

general way that represents the distribution of the late growth

occurring in fruit trees.

Radial growth in apple and other fruit trees was most com¬

monly found to continue latest in fall around the base of the

trunk and its upper roots as well as about the bases of branches

and around crotches ; but in some cases other regions also under¬

went late growth. The distribution of late growth about the

base of the trunk is apparently subject to many variations de¬

pending upon the place of origin of the large upper roots as

well as on the size of the top. Usually the last growth occurs

on the ridges of the roots approximately in the center of the

rounded angle a root makes with the trunk, although in some

87 Cranefield, P. Duration of the growth period in fruit trees. Wise.

Agrl. Expt. Sta. Ann. Rpt. 17:300-8. 1900.

Grosserib acker — Radial Growth in Trees.

45

cases it was found to occur equally late in the upward extension

of such a root-ridge on the trunk. Again, in some instances in

which trees had only two large lateral roots making a rather

narrow angle with each other, very late growth was found to oc¬

cur in the valley-like angle between them. From an earlier pa¬

per88 on crown-rot and the papers cited there it is interesting

to notice that the distribution of that disease on fruit trees con¬

forms fairly closely to the distribution of late radial-growth oc¬

curring at the root-crown region. It was found that in cases

where only a part of the bark was affected it was confined to the

upper angles of lateral roots, or to the very deep angles between

two large laterals.

Pruning fruit trees very heavily often results in a decided re¬

duction in the thickness of the next annual ring toward the base

of the trunk. This was found by pruning some fruit trees in

one of the seedling apple orchards of the New York State Agri¬

cultural Experiment Station in early spring of 1912. The ra¬

dial growth on the lower part of such heavily pruned trees also

continued several weeks later than it did on nearby checks.* *

The result seems to agree with those obtained by Jost, Lutz,

and Kuhns89 in that a reduction of the foliage beyond a certain

amount resulted in greatly reducing growth toward the base of

the stem.

As stated above observations made regarding the occurrence of

crown-rot on fruit trees seemed also to show a possible relation

of that disease to the distribution of late growth. Some New

York apple orchards may be used to illustrate this relation. In

one instance90 two varieties almost equally susceptible to crown-

rot were grown side by side and received the same treatment ex¬

cept that the Baldwin variety was pruned up high while the

other or Ben Davis variety was allowed to grow largely unpruned

and therefore low headed. The Ben Davis trees had been set

for fillers and were not deemed worth the care bestowed on the

88 Crown-rot, arsenical poisoning and winter-injury. N. Y. State

Agrl. Expt. Sta. Tech. Bui. 12:389-94. 1909.

* The writer wishes to thank G. H .Howe of that station for having

the pruning done, and R. Wellington, now of the Minnesota Experiment

Station, for making some of the collections of specimens from these

trees into killing fluids.

89 1. c.

90 Crown-rot of fruit trees: field studies. N. Y. State Agrl. Expt. Sta.

Tech. Bui. 23:18-20, 48, and plate 7. 1912.

46 Wisconsin Academy of Sciences, Arts, and Letters.

other variety since they were to be removed after the Baldwins

had attained some size. Nearly all of the Baldwin trees had the

bark injured about a decimeter above ground during the winter

of 1910-11, and over 80% had practically entire girdles of loos¬

ened or injured bark so that they had become worthless, while

none of the low headed Ben Davis trees were affected. In an¬

other case91 bark injury resulted high up the trunks of bearing

trees after a severe pruning.

It was also found that radial growth is often very late in thick

callus rolls about old cankers and sometimes on the under side,

or on the concave side of crooks in horizontal branches. The

bases of water sprouts or adventitious ascending shoots that arise

on the larger branches of excessively pruned young apple trees

also undergo very late radial growth and apparently for that

reason are winter-injured in those regions ; as in some cases dis¬

cussed on pages 40 to 42 of the above cited paper on crown-rot.

Very similar observations regarding the distribution of winter-

injury in the bark of trees had been made by Nordlinger.92 He

also assumed that such places are injured because of their late

growth.

The reasons for the occurrence of late radial growth at certain

places on trees are doubtless the same as those underlying the

general distribution of excentric growth, and have not been fully

determined as yet. It seems, however, that the re-distribution of

bark pressure incident to radial growth, the distribution of

elaborated food, the location of the channels for water conduc¬

tion, and the gravity-wind pressure effects advocated as factors

which regulate the distribution of radial growth, may afford at

least a partial explanation of the localization of late growth after

they have been submitted to a more careful quantitative study.

WHAT CAUSES RADIAL GROWTH TO APPEAR AS “ANNUAL” RINGS.

The general distribution of radial growth in trees has also an

indirect relation to the development of “annual” rings in that

the proportion of spring and summer wood of a ring at any level

of a stem is doubtless dependent upon the comparative distribu-

01 1. c. p. 24-27.

92 Nordlinger, H. Die September-Froste 1877 nnd der Astwurzel-

schaden (Astwurzelkrebs) an Baumen. Centbl. Gesam. Forstw. 4:489-

90. 1878.

Grossenbacher — Radial Growth in Trees. 47

tion and duration of growth, in the early and late season, over

the different parts of a tree. That is, if in any particular re¬

gion of a trunk radial growth starts very early in spring and

continues rapidly to the end of the spring-growth period a con¬

siderable layer of spring wood will occur in that region; while

if spring growth starts late, proceeds slowly and stops rather

early the thickness of spring wood would be slight. If the dis¬

tribution of summer growth is such as to add but little to a re¬

gion where spring growth had been heavy and much where

spring growth had been slight, the rings resulting in the two

regions would have a very different appearance. To continue

the illustration further, if for some reason radial growth failed

to occur in certain parts of a tree-trunk until after the produc¬

tion of summer wood had begun such parts would show only

small-lumened, thick-walled cells in the ring ; while had the sum¬

mer growth been eliminated in regions where spring growth oc¬

curred the resulting ring would consist of spring wood only.

From the papers cited above on the distribution of radial growth

it is evident that all the cases illustrated here do actually occur

even in the extreme forms used in the last illustration. It is ap¬

parent, therefore, that in some environments and especially on

certain parts of trees the distribution of radial growth may have

a marked influence not only on the type of the resulting ring

but even on the nature of the wood in such portions of stems.

This evident relation between the seasonal distribution of radial

growth on a tree to the type of wood ring to be produced has

reseived practically no attention, although in von Mohl ’s93 paper

on the anatomy of roots it is noted that rings with only the spring

type of wood seem to result owing to the entire omission of the

summer growth; while Sanio94 suggested a similar idea regard¬

ing the absence of spring growth in parts of some rings of a

dwarfed Fraxinus grown in a swamp. Lutz95 also noted the ab¬

sence of summer wood in a pine, from which the buds had been

removed in March, the little growth that occurred was spring

wood. When the wood of roots or stems grown m certain en¬

vironments consist largely of so-called spring wood, elaborate

explanations are usually manufactured to show that the high

"1. c.

84 1. c.

*1. c.

48 Wisconsin Academy of Sciences , Arts , and Letters.

water requirements of such habitats induce the formation of

large vessels throughout the wood for the conduction of the

water needed. This may be typically illustrated by a paper of

von Lazniewski93* on alpine plans in which attention is called to

the fact that the rings in mountain willows are much thinner and

have a greater proportion of vessels per ring than those in trees

of the same species grown in the valleys. Yet it was noted that

the outer parts of the wood rings were usually only partially

lignified, indicating that radial growth had been prematurely

checked. The excessive number of vessels per ring of the alpine

trees was interpreted as being due to the greater demands for

water on the mountains, while the probable fact that the sum¬

mer-wood portion of the rings had perhaps been wholly elimi¬

nated by the environment was not even mentioned. Practically

the same observations although on a larger scale were made by

Rosenthal96 in a later paper and the conclusion was drawn that

the larger number of vessels per unit area of cross section in

willows grown on the mountains is an adaptation to a higher

transpiration rate.

A number of hypotheses have been elaborated in an endeavor

to explain “annual” rings, and more or less data has been col¬

lected by their supporters to substantiate them but with indiffer¬

ent success as judged by Krabbe97, who some years after publish¬

ing his last researches on the subject, maintained that ring

formation cannot be satisfactorily explained with our present

knowledge of the factors determining the size differentiating

cells attain in different parts of the growing season, and of

the ones regulating the thickness of cell walls in different parts

of the rings.

It was recently pointed out by Klebs98 that periodicity in

plant growth occurs in all regions of the world having a periodic

climate, and that the dormant periods coincide with the cold pe¬

riods of temperate climates and with the dry periods of the

tropics. He noted too, that some trees have partial and irregu-

9G* Lazniewski, von, W. Beitrage zur Biologie der Alpenpflanzen.

Flora, 82:224-67. 1896.

98 Rosenthal, M. Ueber die Aushildung der Jahresringe an der Grenze

des Baumwuchses in den Alpen. Inang. Dissertation. Berlin, pp. 24.

1904.

97 Krabbe, G. Einige Anmerkungen zu den neusten Erklarungsver-

suchen der Jahringbildung. Ber. Deut. Bot. Ges. 5:222-32. 1887.

98 1. c.

Grossenbacher — Radial Growth in Trees.

49

lar periodicity even in regions of the tropics having what ap¬

pears to be a practically non-periodic climate.

In central Uruguay" where the temperature never goes much

below freezing and where late summer is a dry season, some trees

have distinct yearly wood-rings, while in others more than one

ring is produced in a year. Robinia Pseudkcacia and Melia

azedarach have fairly evident annual zones, but they also have

imperfect secondary zones due to a concentric arrangement of

large vessels. In Acacia the yearly zonation is less distinct but

the last wood is usually made up of cells with a reduced radial

diameter.

The measurements by Hall100 show that the trunks of trees in

Uruguay usually increase in circumference during nearly ten

months of the year, and that in some cases they even increased

during the months of May and June (winter). He found, how¬

ever, that the circumference of most trees decreased more or less

during winter, the deciduous trees more noticeably than the ever¬

greens. Ursprung101 found that a number of the evergreen trees

and shrubs of a tropical locality without any appreciable peri¬

odicity of climate showed a zonation in cross sections of the stems

without the presence of any evident histological difference in the

wood of the different parts of zones. Some of these species are

said to become deciduous in localities having a periodicity in the

water supply wTith the result that the zonation of their wood be¬

comes more marked. Holtermann102 also studied the relation of

climate to radial growth in the tropics and came to the conclu¬

sion that the formation of growth rings in the wood is intimately

connected with the occurrence of periods of markedly different

transpiration rates, and that the larger vessels are developed to

meet the demands of increased transpiration. He holds that

tropical trees growing in a saturated atmosphere most of the

time have no indication of zonation in the wood even though they

99 Christison, D. On the difficulty of ascertaining the age of certain

species of trees in Uruguay, from the number of rings. Trans. Bot. Soc.

Edinburgh. 18:447-55. 1891.

190 Hall, C. E. Notes on the measurements, made monthly at San

Jorge, Uruguay, from January 12, 1885, to January 12, 1890. Trans.

Bot. Soc. Edinburgh. 18:456-68. 1891.

101 1. c.

102 Holtermann, C. Der Einfluss des Klimas auf den Bau der Pflan-

zengewache. Anatomisch Physiologische Untersuchungen in den

Tropen. pp. 249. 1907. Leipzig.

4— S. A.

50 Wisconsin Academy of Sciences , Arts , and Letters.

are deciduous like some species of Leguminosae, Guttifereae and

Ficus. On the other hand it is noted that a seven-year-old tree

of Theobroma Cacao had developed 22 radial-growth rings, and

since it cast its leaves three times a year it is evident that the

number of rings corresponded with the vegetative seasons of the

tree. The real cause of zonation is thought to be an inherent

characteristic of a plant though the environment induces its

manifestation.

According to Dingier103 leaf-fall is more dependent on the age

of the leaves than on the environment, for by cutting back decid¬

uous trees in Ceylon some time before the normal period of leaf-

fail the new crop of leaves which immediately came out was re¬

tained throughout the dormant season which is dry and very hot.

Unfortunately the effect upon radial growth was not noted but

from evidence given above it seems very likely that the periodic¬

ity of radial growth always follows foliar periodicity in decid¬

uous trees whether natural or induced.

In another paper he104 reported that the folier periodicity of

European fruit and forest trees grown in the highlands of Cey¬

lon is very irregular even in different branches of individual

trees. In late October the trees of Quercus pedunculata could

be divided into five classes in regard to the condition of their

foliage, ranging all the way from cases in which chiefly old

spotted leaves were present (though some scattered buds were

swelling) to instances wThere no old leaves were present and the

new shoots occurred in all stages of elongation, although most of

them were full grown. Quercus Cerris had a more uniform

periodicity. In late October all trees bore two generations of

leaves : the old ones hard and spotted, alth ought still green, and

the young ones not yet full grown. In late November the old

leaves had practically all fallen and the new elongation growth

had been completed. European pears, peaches, cherries, plums

and apples were found to have practically the same periodicity,

producing two crops of leaves and flowers, though but one crop

of fruit per year. The trees are often almost leafless some time

los Dingier, H. Versuche iiber die Periodizitat einiger Holzgewachse

in den Tropen. Sitzungsber. Math.-Physical. Kl. Kgl. Bayer. Akad.

Wiss. Miinchen. 1911:127-43. 1911.

104 Dingier, H. uber Periodizitat sommergriiner Baume Mittele-

uropas im Gebirgesklima Ceylons. Sitzungsber. Math.-Physical. Kl.

Kgl. Bayer. Akad. Wiss. Miinchen. 1911:217-47. 1911.

Grossenbacher — Radial Growth in Trees.

51

in February or March. All stages of bud and leaf are said

usually to occur in these trees.

An experiment similar to that performed by Dingier had pre¬

viously been made by Wright105 in Ceylon. He lopped trees of

Mangifera indica and Terminalia Catappa in May and new

leaves developed from July to September, with the result that

no new leaves were produced on these trees in February and

March when others of those species developed new crops of

leaves. Some of the plants develop new leaves once or twice and

others several times annually, and immature leaves may be found

during every month of the year. Only a comparatively small

percentage of the Ceylon trees are said to be deciduous. Some

rapidly growing species were found to become defoliated at the

end of the first year and others at the end of the second ; while

the more slowly growing ones may vegetate as evergreens until

the close of the fifth or sixth year before losing their leaves.

Usually, after a tree has once lost its leaves it loses them annually

but some species are deciduous only in youth and become ever¬

green later. Some of the so-called evergreen trees are said to

also lose all the leaves in occasional years before the new crop ap¬

pears. In some species periods of sparse foliation occur two or

three times per year and in others the foliage is more copious

on alternate years. It is held that the absence of any very

marked periodicity in the environment permits some plants to

follow their inherent periodicity of growth, while the annual

variation in the transpiration rate and atmospheric moisture are

thought to be the cause of the deciduous habit of others.

These observations on foliar periodicity by Dingier, Wright

and others seem to show that Dingier may be correct in his con¬

tention that leaf-fall is more dependent upon the normal dura¬

tion of life of the leaves than upon the environment. However,

if that should prove to be a fact, it would necessarily follow that

certain plants are deciduous not because of the leaf-fall but on

account of the failure of a new crop of leaves to develop before

the old ones drop. Such a view centers attention upon the causes

inhibiting growth rather than upon the causes of leaf -fall in the

study of periodicity, a method of attack adopted by Klebs in the

paper cited above.

106 Wright, H. Foliar periodicity of endemic and indigenous trees in

Ceylon. Ann. Roy. Bot. Gard. Peradeniya 2:415-516. 1905.

52 Wisconsin Academy of Sciences, Arts, and Letters .

It seems then that although trees having annual or more prop¬

erly radial-growth rings are distributed all over the arborescent

world, one or more factors of their envionment must be effective

periodically in order that marked zonation occur. The more or

less regular recurrence of cold or dry seasons are the factors

usually noted in connection with periodically recurrent vegeta¬

tive seasons, but doubtless any other recurrent environmental

factor influencing growth may also affect zonation, e. g., periodic

variation in the supply of inorganic foods as was suggested by

Klebs.106 It should be noted, however, that wood zonations re¬

sulting from recurrent dry periods of the tropics even in decid¬

uous trees are not as marked as those occurring in temperate

zones where the dormant period is chiefly due to seasonal varia¬

tions in the temperature and where consequently a greater sea¬

sonal change occurs in the bark pressure.

The causes of the formation of radial-growth rings have been

studied mainly in the north temperate zone and, therefore, ex¬

planations are largely based on the environmental factors that

seem to be operative in that region. Seasonal changes in bark

pressure, in the supply of metabolized food to the cambium, and

in the rate of transpiration have been either separately or in

partial combination advanced as explanations for the occurrence

of the large-celled spring-wood alternating with small-celled

summer-wood.

The bark-pressure hypothesis : — Sachs107 seems to have been

the first to suggest that the difference between spring and sum¬

mer wood may be due to a difference in the bark tension or pres¬

sure obtaining in spring and summer. The idea was then tested

experimentally by de Vries108 with the result that Sachs ’ hy¬

pothesis seemed to have been sustained. The experiments by de

Vries consisted in making some longitudinal slits in the outer

bark of various trees in spring and of applying ligatures to the

stems of others. On the following winter it was found that only

about one-half as many cells had been produced under the liga¬

tures as occurred on other parts of the past season ’s ring ; while

in the regions where the outer bark had been slit the number of

106 1. e.

107 Sachs, von, F. G. J. Lehrbuch der Botanik. 1. Aufl. 1868, p. 409.

108 Vries, de, H. Ueber den Einfluss des Rindendruckes auf den ana-

tomischen Ban des Holzes. Vorlaufige Mitlheilung. Flora. 33:97-102.

1875.

Grossenbacher — Radial Growth in Trees.

53

cells had become two to three times that produced in the normal

portions of the ring. Similar experiments also showed that the

amount of radial and tangential growth of cells differentiating

from the cambium is inversely proportional to the pressure ex¬

erted on them. It also seemed that pressure acts as a selecting

agent in determining the proportion of vessels to wood fibers;

i. e. the greater the pressure the fewer the vessels and the more

numerous the wood fibers to be produced. De Vries concluded

therefore that bark pressure influences the rate of cambial di¬

vision as well as the relative size cells may attain during differ¬

entiation. Since bark-growth follows the enlargement of the

wood cylinder it was thought evident that bark pressure is

greater toward the end of the radial-growth period than at its

beginning. For these reasons de Vries held that a seasonal

change in bark pressure is the chief cause of seasonal growth ap¬

pearing as “ annual” rings.

In some later experiments, while studying wound wood, he109

found on lifting loose strips of bark with a knife on the concave

side of young tree-trunks held in a bent position, and then tying

it in place again in such a way as to prevent evaporation, that

numerous large vessels developed in the new wood produced un¬

der the strips. He reiterated his former conclusion that bark

pressure is an important factor in determining the size of wood

cells and that it is largely responsible for the difference between

spring and summer wood.

That bark tension does occur on enlarging stem structures had

been shown by Kraus109 as well as by Nordlinger110 but neither

of them secured quantitative results of value.

The influence of pressure on cambial activity and cell differ¬

entiation have since been investigated from various viewpoints

and have led to different conclusions. Hohnei* * 111 found sharp¬

angled transverse displacements in the bast fibers of many

Dicots at points where neighboring cells make an abrupt uneven

109 Vries, de, H. Ueber Wundholz. Flora. 34:2-8; 17-25; 38-45;

49-55; 81-88; 97-108; 113-21; 129-39. 1876.

109 Kraus, G. Die Gewebespannung des Stammes und ihre Folgen.

Bot. Zeit. 25:105-19; 121-33; 137-42. 1867.

110 Nordlinger, H. Spannt die Baumrinde im Sommer nicht? Kritische

Blat. Forst-u. Jagdwiss. 52: (1) : 253-55. 1870.

111 Hohnei, von, F. Ueber den Einfluss des Rindendruckes auf die

Beschaffenheit der Bastfasern der Dicotylen. Jahrb. Wiss. Bot.

15:311-26. 1884.

54 Wisconsin Academy of Sciences , Arts , and Letters.

joint. Such transverse displacements or sharp double-bends

were found in about two-thirds of the fifty to sixty species ex¬

amined. They were especially prevalent in Urticaceae, Apocy-

naeeae, Asclepidaceae, Linaceae, etc., while in other families the

double-bends occurred only in certain genera. None were found

in the Rosaceae including the pomaceous group, nor in the Tilia-

ceae and Cupuliferae.

It was held that the sharp bends are due to bark pressure, as

indicated by the fact that in the plants in which these bends

commonly occur the bast-fibers are but slightly or not at all

lignified. Hohnel held that if the double bends were not due to

growth or bark pressure they would not always appear at points

in the fibers where joints or breaks occur in the cells of the sur¬

rounding tissues. The failure of the bends to become evident

until after the tissues are fully differentiated was taken to indi¬

cate that bark-pressure becomes greater during the latter part of

the differentiation period. It also seemed that in case of Urtica,

Cannabis and Linum the bark pressure was often greater in the

lower part of the stem than above, for the angular bends were

frequently present on the fibers of the lower part while none oc¬

curred in the upper. The transverse displacements were found

to be made up of two successive sharp bends which were notice¬

able in all layers of the wall. In many cases some of the layers

were actually ruptured.

Krabbe112 made extensive studies of bark pressure and tried

to obtain some quantitative measurements. He increased bark

pressure by encircling tree-trunks with a chain much like that

now used on bicycles, except that it was wider. One end of the

chain was fixed to an iron peg driven into the tree and the other

ran over a pulley and had a weight pan attached. A piece of tin

a little wider than the chain was placed about the trunk under

the chain to distribute the pressure more evenly and to reduce

friction. Weights were put into the pans in accordance with the

determinations of bark pressure obtained before, and it was

found that the bark pressure had to be doubled and even quad¬

rupled before any influence on the size of the cells or the thick¬

ness of the yearly growth became evident.

112 Krabbe, G. tiber die Beziehung der Rindenspannung zur Bildung

der Jahrringe and zur Ablenkung der Markstrablen. Sitzungsber.

Akad. Wiss. Berlin 1882: 1093-1143. 1882.

Grossenbacher — Radial Growth in Trees .

56

The “normal” bark pressure was determined by stretching

rings of bark over a smooth cylinder by means of weights until

the bark had attained the length it had while still attached to

the tree. In his later work113 the rings of bark were straightened

out and weighted at one end to determine the force required to

stretch the bark to its former length, for it was found that the

results obtained in this way were the same as those gotten with

the more elaborate apparatus. The bark pressure of conifers was

found to be usually under one-half an atmosphere and that

of broad-leaved trees about twice as great. In case of conifers

the pressure seemed to increase in fall on an average about

0.8 gm. per square millimeter of cross section, while the average

of similar measurements on a number of broad-leaved trees indi¬

cated a decrease of pressure in fall equal to 12.5 gm. per square

millimeter of cross section. He maintained that the breaking

strain of bark is never reached by growth pressure. Bark pres¬

sure was found greatest in regions of most rapid radial growth,

for instance on the side of excentric stems with the longer radius.

By using pressures from five to eight atmospheres the sum¬

mer-wood type of radial growth was induced in spring on trees

having comparatively little difference in the size of spring and

summer-wood cells, while on trees having very marked differ¬

ences between spring and summer wood it was practically impos¬

sible to induce the formation of the summer-size of cells in

spring by increasing the bark pressure. In reducing the bark

pressure by means of longitudinal slits in the outer bark in sum¬

mer, typical spring wood vessels developed in trees which nor¬

mally have only a slight difference between size of spring and

summer wood cells; but in trees like Quercus and Fraxinus in

which a marked difference occurs between spring and summer

wood, the spring wood vessels could not be thus induced.

Krabbe therefore concluded that bark pressure remains practi¬

cally the same throughout the growing season and that changes

in bark pressure could not be the cause of ring formation be¬

cause it requires such a great increase to influence the size of the

wood cells.

118 fiber das Wachsthum des Verdickungsringes und der j ungen Holz-

zellen in seiner Abhangigkeit von Druekwirkungen. Abhandl. Kgl.

Akad. Wisa. Berlin. 1884. Anhang. 1:1-80. 1885.

56 Wisconsin Academy of Sciences , Arts , and Letters.

Gehmacher114 also performed some experiments in the increase

and decrease of bark pressure on three to six-year-old trees and

shrubs. The outer cortex was slit in February and nearby on

the same stem a ligature of tightly wound wire was applied and

the stem allowed to grow until the end of the season.

The number of cork cells varied inversely as the pressure and

their radial diameter was decreased by 11% under increased bark

pressure, while under reduced pressure an increase of 13% above

normal resulted. A similar effect was noted on the cortical

parenchyma cells except that both the radial and tangential di¬

ameters were decreased under increased pressure and the inter¬

cellular spaces were obliterated, while under reduced pressure

the cells became globular and the intercellular spaces were in¬

creased in size above the normal. The difference between the

thickness of the cortical parenchyma under increased and that

under decreased pressure was enormous. In the wood the num¬

ber of fibers increased and that of vessels decreased under added

pressure, while the number of bast fibers was greatly reduced by

increased pressure. Gemacher ’s conclusion was that it does not

require the enormous differences of bark tension to influence the

size of wood cells as had been maintained by Krabbe.

Hoffman115 also investigated the influence of pressure on cell

division and differentiation in the cambium of trees and con¬

cluded that the forces which contribute to the development of

cylindrical stems rather than some other form are (1) bark ten¬

sion and the consequent bark pressure, (2) radial-growth pres¬

sure, and (3) the passive resistance of the wood. Cambial di¬

vision and growth are said to continue only as long as growth

pressure exceeds bark pressure and it is thought that if bark

pressure is equal on all sides the axis .must either be or soon will

become cylindrical on occurrence of continued radial growth.

This is shown by the fact that angular young shoots become

cylindrical on growing older. Even when the tension of the

bark is the same all around a branch bark pressure may be dif¬

ferent at different points, being considerable at prominences and

114 Gehmacher, A. Untersuchungen uber den Einfluss des Rinden-

druckes auf das Wachstum nnd den Bau der Rinden. Stizungsber. K.

Akad. Wiss. Wien. 88 Abt. 1:878-96. 1884.

116 Hoffman, R. Untersuchungen uber die Wirkung mechanischer

Krafte auf die Teilung, Anordnung und Ausbildung der Zellen beim

Aufbau des Stammes der Laub- und Nadelholzer. Inaug. Dissertation.

Berlin. 1885. pp. 24.

Grossenbacher — Radial Growth in Trees.

57

perhaps zero or even negative in depressions. Among the nu¬

merous angular young twigs examined the greater pressure at

the angles did not prevent the development of normal spring

wood, but larger numbers of both spring and summer wood cells

were produced in the depressions than on the ridges until the

twig became cylindrical.

It was found that when a tree-trunk or branch presses against

some non-yielding object or the bases of the component branches

of a forked stem press against each other, radial growth is re¬

duced on the side of contact when the pressure has reached a

certain intensity and that the rays spread outward and eventu¬

ally became parallel to the obstructing surface. The continu¬

ance of radial growth tends to separate or pull apart the com¬

ponents of a forked stem or widen the upper angle a branch

makes with its axis. Branches thus firmly pressed against each

other eventually fuse and the rays then come to radiate from

the common center and further radial growth tends to result-

in a cylindrical, united structure. It was found that the callus

developing at the cut end of a twig in water also conformed to

the general law of the mechanics of radial growth in that its

cross sections become semicircular with a rough outline ; but the

surface becomes smooth as tension is developed by further

growth. When a rectangular piece of bark was cut from a tree

the first division of the cambium in the formation of a callus is

said to be by a radial wall or one at right angles to the wall

formed under normal conditions. Further growth and division

was also found to occur in accordance with the resistance to

growth and resulted in a structure having its center at the place

where the first cambial divisions took place. The rays in the

bark on both sides of the piece cut out become diverted not only

by the contraction of the bark at the time the piece was cut but

also by the lack of surface growth in the bark surrounding the

wound. The omission of surface growth is said to be due to the

lack of accustomed tangential pull formerly exercised by the ex¬

cised piece. Growth is resumed only after the callus bark has

reached a tension comparable to that of the piece removed. This

resulted in increased radial growth in the entire region in¬

fluenced by the wounding, as shown by a count of the number of

cells produced here as compared to that produced in other places.

When the cambium was first freed from its normal bark pres-

58 Wisconsin Academy of Sciences , Arts , and Letters.

sure its cells took on isodiametric forms which were retained un¬

til the hark pressure became appreciable again and then reverted

back to the elongate form normal to the species. It is held that

the upper and lower edges of a wound do not produce callus as

copiously as the lateral ones because of the lesser reduction of

bark pressure, and the death of the cut cells which extend some

distance above and below the wound.

From his experiments in which ligatures were applied to

stems Sorauer116 concluded that slow radial growth combined

with high bark pressure results in twisted grain and that a re¬

duction of bark pressure below normal not only induces more

cells to form from the cambium, but cells having a greater di¬

ameter and a reduced length.

Newcombe117 found that when external conditions prevent

growth, the unfinished tissues remain unaltered and thin walled ;

that mechanical resistance or pressure prolongs the differentiat-

tion period, the cells remaining smaller and thinner walled.

The occurrence of numerous cocoons of bag- worms on various

species of trees and the fact that the narrow silken bands by

which they are attached to the twigs are often too strong for ra¬

dial growth pressure to break, afforded von Schrenk118 an occa¬

sion for a study of the effects of excessive pressure on radial

growth. In most cases the silken bands encircling the twigs are

burst early in the summer of the year following the time of the

attachment of the bags. In some instances in which the liga¬

tures were too strong to be ruptured by the thickening twigs the

transfer of elaborated food was eventually checked and an en¬

largement developed on the distal side of the constricting band.

In other cases the ligature was sufficiently distended by growth

to permit of some food transfer and resulted in the formation of

welts on both sides of bands. In some instances the pairs of

welts fused above the ligatures and reestablished normal connec¬

tion and pressure. In arbor vitae the wood fibers of the first

116 Sorauer, P. Handbuch der Pflanzenkrankheiten. Dritle Auflage.

1:764-66. 1909.

117 Newcombe, F. C. The influence of mechanical resistance on the

development and life-period of cells. Bot. Gaz. 19:149-57; 191-99;

229-36. 1894.

118 Schrenk, von, H. Constriction of twigs by the bag-worm and in¬

cident evidence of growth pressure. Ann. Rpt. Mo. Bot. Gard. 17 : 153-81.

1906.

Grossenbacher — Radial Growth in Trees.

59

year’s growth were often found arranged at right angles to the

axis, under unbroken bands.

In the latter part of the second summer following the attach¬

ment of the bags the portion of the twigs distad to the constric¬

tion had much starch in the bark rays and pith, while that on

the basad side was practically devoid of it.

In hard-wood trees both bark and wood were found to have

continued growing under unbroken bands though welts developed

on both sides. The first wood cells formed under the ligatures

were normal but those developing afterwards had a shorter ra¬

dial diameter and thicker walls than those under normal pres¬

sure. The number of vessels appeared to decreaes in proportion

to the pressure. The wood fibers developing under high pres¬

sure were found to have their long axis at right angles to the

twig or parallel with the compressing band, and the rays were

bent or buckled laterally unded pressure. It is held that the in¬

creased pressure induces the formation of smaller wood cells not

because cambial division occurs before the cells have attained the

normal size but because the pressure hinders their enlargement

during subsequent differentiation.

A large number of tests made to determine the breaking strain

of the bands from both conifers and broad-leaved trees showed it

to be about 40 atmospheres; and, therefore, indicates that

Krabbe’s experimental results showing a growth pressure of 15

atmospheres are too low, since von Schrenk’s observations show

that the majority of the bag-worm ligatures are ruptured by

the enlarging twigs.

An osmotic-pressure hypothesis. — In a paper on the devel¬

opment of pits in the wood cells of the Abietineae Russow119

suggested another explanation of ‘ ‘ annual ’ ’ rings. He claimed

that the bark pressure hypothesis of Sachs which de Tries en¬

deavored to support by experiment, cannot account for the oc¬

currence of growth rings in the wood because the last phloem

cells of a season do not have a reduced radial diameter and on

account of the fact that two rings may be induced by defoliat¬

ing trees. The bark-pressure hypothesis is also held to be dis¬

credited by the occurrence of growth rings in the tropics where

119 Russow, E. tiber die Entwicklung des Hoftiipfels, der Membran

der Holzzellen und des Jahresringes bei den Abietineen, in erster Linie

von Pinus silvestris L., Sitzungsber. Naturfor. Ges. Dorpat 6: 147-57.

1884.

60 Wisconsin Academy of Sciences, Arts, and Letters.

the bark is not distended by low temperature during a dormant

season. In another paper he120 added that in accordance with

the bark-pressure hypothesis the wood cells in roots ought to be

small while as a matter of fact they are large. On the other

hand he held that the changes in the radial diameter of cells

from spring to fall can easily be explained by assuming the

presence in them of highly osmotic substances, which induce a

high hydrostatic pressure and as a result give rise to large cells

in spring, while toward the end of the radial-growth period the

hydrostatic pressure in differentiating cells is reduced owing

to a reduction of the osmotic pressure in them. By using solu¬

tions of glycerine as plasmolysing agents Wieler121 found that

osmotic pressure in herbaceous plants was less than that in the

living wood and ray cells of trees where it ranged from 13 to 21

atmospheres. No difference was found, however, between the

osmotic pressure in differentiating wood vessels and of that in

the cambium cells. He thought that the walls of differentiating

spring-wood cells are more distensible than those of summer

wood owing to their lower cellulose content.

Seasonal variation in the available elaborated food as the

cause of “annual” rings: — After years of intimate study of

forest trees Hartig122 concluded that since radial growth begins

in spring under suboptimal environmental conditions and while

the new leaves are very small or the buds are just bursting, the

nutritive conditions of the cambium must also be suboptimal

and for that reason the spring wood has thin cell walls. As the

season advances the leaves attain full size which in connection

with the accompanying seasonal changes are conducive to the man.

ufacture of the larger quantities of organic foods which, accord¬

ing to Hartig, are responsible for the production of the thicker

walled summer-wood cells. It is held that the chief difference be¬

tween spring and summer wood consists essentially in the thick¬

ness of the cell walls and that the improvement in the nutrition of

the cambium from early spring until the later summer is re-

120 Russow, E. liber den Inhalt der parenchymatischen Elemente der

Rinde vor und wahrend des Knospenaustriebes nnd Beginns der Cam-

biumthatigkeit in Stamm und Wurzel der einheimiseben Lignosen.

Sitzungsber. Naturfor. Ges. Dorpat. 6: 388-89. 1884.

121 Wieler, A. Beitrage zur Kentniss der Jahresringbildung und des

Dickenwachstums. Jahrb. Wiss. Bot. 18: 70-132. 1887.

122 Hartig, R. Ein Ringlungsversuch. Allgem. Forst-u. Jagd-Zeit.

65: 365-73; 401-410. 1889.

Grossenbacher — Radial Growth in Trees.

61

sponsible for the occurrence of “annual rings. ” Hartig stated

however, that the differences in the nutritive conditions cannot

account for the change in radial diameter of wood cells nor for

the presence of the larger proportion of vessels in spring wood,

and maintained that the transpiration current determines their

size. He suggested that the reason so little difference exists in

the radial diameter of spring and summer wood cells of Populus,

Salix, Acer, etc., is to be found in the fact that these trees con¬

tinue producing new leaves throughout most of the radial

growth period’ and because they have no duramen. Since the

water current in trees with duramen is necessarily confined to

the outer layers of wood its effects on cells differentiating from

the cambium are thought to be more marked and therefore re¬

sult in greater differences in the diameter of spring and summer

wood cells, e. g. in oaks, etc. According to Hartig, then, “ an¬

nual” rings are primarily due to the poor nutritive conditions

of the cambium in spring being followed by a period of more

abundant supply of metabolized food in summer, and secondari¬

ly to a decrease in the intensity of the transpiration current

toward the end of the radial-growth period.

Wieler123 came to a diametrically opposed conclusion regard¬

ing the differences in the nutritive conditions about the cambium

in spring and summer. He thought that since the character¬

istics of 4 4 annual ’ ’ rings lie in the type of wood produced in the

early and late growing season and not in the succession of rings,

the relation of different nutritive conditions to the formation

of spring and summer xylem could be more easily determined

experimentally with herbaceous than with woody plants. This

was deemed permissible owing to the fact that in an examination

of 54 species of herbs belonging to 21 families the characteris¬

tic reduction in the size of the xylem cells toward the end of the

growing season as is typical of the “annual” rings of woody

plants, was found in over half of them.

Seedlings of Ricinus communis were set into the soil of one-

fourth to one-half liter pots in spring, well watered and given

optimum light and temperature conditions, but they grew slow¬

ly and remained dwarfs. In early summer four of them were

transplanted to the soil in a field and three of them into good

soil in four liter pots. Those remaining in small pots were

123 1. c.

62 Wisconsin Academy of Sciences , Arts , and Letters.

only about 27 cm. high in January and their stems about 21 mm.

in circumference, while those in four liter pots were about 90

cm. high and 50 mm. in circumference. Those transplanted to

the field became large plants with woody stems. Five dwarfed

plants, which were subsequently transplanted to a forcing bed,

had since made a rank growth and were retransplanted to four

liter pots. They wilted but eventually recovered their turgid-

ity, although the older leaves died.

Cross sections showed the xylem cells of the field plants

to be larger and the vessels more numerous than in those re¬

tained in the small pots. In the plants transplanted to the field

the xylem cells around the pith were small and were surrounded

by larger ones toward the periphery. In case of those trans¬

planted to four liter pots the same inversion of the normal po¬

sition of large and small celled xylem occurred, but in addition

the outermost rows again had a much reduced radial diameter.

In the field plants which had been retransplanted to pots the

outermost cells also had a reduced radial diameter and thick

walls while within them was a zone of large, thin-walled cells

which had apparently been formed just before the last trans¬

planting and as a result their walls remained unthickened.

Similar results were also obtained with Helianthus annuus.

Wieler concluded from these experiments that the abundant

supply of metabolized food to the cambium is the most important

factor in the production of spring wood and that the shortage

of such a food supply induces the formation of summer wood,

and that therefore “annual” rings of trees are due to an abun¬

dant supply of organic food to the cambium in spring and a

reduced supply in summer.

Lutz124 was of the opinion that when the food supply to a

rapidly dividing cambium is comparatively low while water is

abundant the cells become large and thin-walled as is charac¬

teristic of spring wood, while if the food supply is good and the

water is low the cells become small and thick-walled as in sum¬

mer wood.

In a later paper Wieler125 reiterated his former conclusions

though he admits his inability to prove that the small radial

124 1. c.

125 Wieler, A. Ueber die Abhangigkeit der Jahresringbildnng von den

Ernahrungsverhaltnissen. Allgem. Forst-u. Jagd-Zeit. 67: 82-89. 1891.

Grossenbacher — Radial Growth in Trees.

63

diameter of summer-wood cells results from a reduced supply

of food to the cambial region; nevertheless, it is held to be a

more likely contention than that maintained by Hartig to the

effect that summer-wood results from an increase in the supply

of metabolized food.

In this paper "Wieler cited similar experiments by Sachs126 in

support of his conclusions, although Sachs noted that the fre¬

quent addition of abundant nutrient solution failed to induce

more growth in small pots. Sachs held the dwarfing in small

pots to be due to a crowding of the root system into mats in such

a way as to greatly impair their absorptive functions.

The relation of rest and food supply to the production of

wood rings: — Mer127 held that the winter rest of the cambium

and its consequent great activity in spring in connection with

the abundance of plastic materials at that time are the causes

of the production of large-celled spring wood. The cell walls

of spring wood are thought to remain relatively thin because

the food transfer through such a thick differentiating zone of

cells is comparatively slow, and the thick walls of summer wood

cells are assumed to be due to slow rate of cambial division or

to the thinness of the differentiating zone and consequent ready

access of organic food to its cells. The sudden and consider¬

able decrease in the radial diameter of the peripheral few rows

of wood cells in a year’s growth is held to be due to an arrest

of their development as a result of enfeebled cambial activity

rather than to an increase of bark pressure as maintained by

Sachs, de Yries and others.

A summary and comparison of the hypotheses • — The work

of Kraus, de Yries, Nordlinger, Detlefsen, von Hohnel, Ge-

macher, Hoffman, Kny, Newcombe, von Schrenk, and Sorauer,

have made it apparent that pressure on the cambium affects the

rate of cell division as well as the size differentiating wood cells

may attain, but owing to the fact that no method has as yet

been developed by means of which quantitative measurements of

bark pressure can be made it is impossible to determine just

what relation bark pressure has to the production of “ annual”

rings.

126 Sachs, von, F. G. J. Vorlesungen fiber Pflanzenphysiologie. Leip¬

zig. 1882. p. 623.

127 1. c.

64 Wisconsin Academy of Sciences , Arts, and Letters.

The different degrees of hydrostatic pressure assumed by Rus-

sow as the cause of the difference between spring and summer

wood has apparenty also been implied by Hartig, Mer and

others in speaking of growth force, etc., but even more than in

the former case do the few qualitative tests need to be replaced

by quantitative measurements before the validity of the idea

could be tested.

Hartig has collected a mass of observational and even some

indirect quantitative data that seem to support his hypothesis

that the relative abundance of elaborated food determines the

thickness of cell walls and that the relative intensity of the

transpiration stream determines the length of the radial diam¬

eter of wood cells, but the experiments of Jost, Lutz and others

show that although food and water may be present in great

abundance very little or no radial growth occurs when termi¬

nal growth is prevented.

Wieler’s hypothesis that the abundance of metabolized food

in the cambial region in spring induces the formation of spring

wood and its reduction, summer wood is also lacking in that it

does not account for the cessation of radial growth on the re¬

moval of the elongation structures. Besides, the experiments

with which he assumes to have made his contention probably in¬

volved too many unknown variables to afford even a satisfactory

test of the hypothesis.

The results obtained by Morgulis128 in his experiments in al¬

ternately feeding and starving salamanders tend also to make

one skeptical regarding the value of the hypotheses of both

Hartig and Wieler as explanations of ring formation because

Morgulis found “That the rate of growth is independent of the

amount of nutrition ’ ’ and that ‘ ‘ The impulse to grow plays the

leading part’’ and “determines the degree of utilization of the

nutriment.” Finally, he found too that “From all that has

preceded, the conclusion can be drawn that periodic starvation

is more detrimental to the organism than acute starvation fol¬

lowed by a liberal supply of food. In the former case the in¬

dividual remains below the level of the normally fed animals;

in the latter case, on the contrary, provided the inanition has

128 Morgulis, S. The influence of protracted and intermittent fasting

upon growth. Amer. Nat. 47: 477-87. 1913.

Grossenbacher — Radial Growth in Trees.

65

not been carried too far, the restorative process may go even be¬

yond the limit attainable under normal conditions. ’ ’

Since Hartig laid especial stress on the difference in the thick¬

ness of cell walls rather than the size of cells as the essential

difference between spring and summer wood his secondary fac¬

tor, the relative intensity of the transpiration current, would

come in for first consideration because it is claimed to regulate

the size of cells. It seems possible that the full report prom¬

ised by Jaccard129 on the tree-trunk as a shaft of equal water

conductance may throw more light on Hartig ’s idea.

The possible relation of enzymes to the formation of “annual”

rings: — In cases of this kind in which the hypotheses are so

numerous and the advocates of each can marshal at least a por¬

tion of the observed facts in support of their views the truth

usually lies somewhere between them, and each conflicting ex¬

planation will eventually contribute certain fragments to a

theory that will account for the known facts. The time for such

a theory has not yet come. However, since none of the pro¬

posed hypotheses gives promise of becoming such an explana¬

tory theory it may be pardonable to submit yet another with

the hope that the viewpoint thus suggested might lead to a new

attack on the problem.

From our present knowledge it seems that to be of any value

as a basis for work or a stimulus for the further study of radial

growth rings such an hypothesis must, by using all known and

some probable but undetermined facts explain how it is that

wood cells have a smaller radial diameter in summer than in

spring and why vessels are often wholly lacking in the later

summer wood.

It has been shown that an “annual” ring consists essentially

of a sheath or ring of wood produced during one more or less

continuous radial-growth period and that it is made up of two

types of wood which may merge gradually into each other or

join at a rather abrupt line. That portion of the ring devel¬

oped in “spring” or during the early part of a new elongation-

growth period has larger cells than that produced in “summer”

or after the closing of the first elongation, following the princi¬

pal dormant season. In the case of trees in temperate zones

and many of those in the tropics which produce new leaves near-

129 1. c.

5— s. A.

66 Wisconsin Academy of Sciences , Arts, and Letters.

ly throughout the vegetative season the growth rings are not

very marked though they are usually apparent. Generally the

most reliable criterion for distinguishing the rings Is the reduc¬

tion in the radial diameter of at least the last row or two of

wood cells; yet in the tropics histological distinctions are said

to be practically absent in some trees, and their rings may only

be distinguished by slight demarking lines.

The work reviewed in this paper has shown that the environ¬

mental factors which control elongation growth also influence

radial growth and that ordinarily the prevention of elongation

by the removal of vegetative points hinders growth in thick¬

ness even when the environmental conditions are optimal and

the food and water supply abundant. Klebs130 assumed, in fact,

that large quantities of organic foods accumulating in plants

inactivates the enzymes concerned in elongation and therefore

brings about a cessation of growth in length. According to

him a timely increase in the water and inorganic nutrients may

reactivate or prevent inactivation of the growth enzymes and

thereby shorten or eliminate the dormant period.

With such a precedent one may also assume the presence .of

enzymes which incite and maintain radial growth since there

are a number of phenomena to be noticed in connection with

growth in thickness that support such an assumption, as may

be gathered from the following papers.

In an investigation on the reserve food in seeds Reiss131 found

that cellulose is laid down on the inner side of cell walls of

many seeds and that it is largely redissolved on germination.

Schulze132 made a similar study of lupine seeds and found con¬

vincing evidence that the inner layers of the cotyledonary cell

walls are used up during germination. It seemed that the dis¬

solving part of the walls is a hemiceliulose which gives rise to

galactose and arabinose on hydrolysis. Griiss133 also noted the

occurrence of the hemicelluloses, galaetan and araban, in plant

i33 Gr(jSS) j. Ueber Losung und Bildung der aus Hemiceliulose besteh-

enden Zellwande und ihre Beiziehung zur Gummosis. Biblio. Bot. 39.

1896. pp. 14.

130 1. c.

131 Reiss, R. Ueber die Natur der Reservecellulose und liber ihre Auf-

losungsweise bei der Keimung der Samen. Ber. Deut. Bot. Ges. 7 : 322-

29. 1889.

132 Schulze, E. Ueber die Zellwandbestandtheile der Cotyledonen von

Lupinus lutens und Lupinus angustifolius und liber ihr Verhalten wah-

rend des Keimungsvorgangs. Ber. Deut. Bot. Ges. 14: 66-71. 1896.

Grossenbacher — Radial Growth in Trees.

67

cells, and that they may be dissolved or converted into gum by

enzymes. Potter134 called attention to the presence of an inner

cellulose layer in the xylem cells of many normal trees, and to

its especial abundance in the wood fibers of Quercus, Fagus,

Aesculus, Salix, Ulmus, Alnus, and Betula. He found that

after keeping wood in water during some days cellulose linings

became apparent in many cells in which none had been noted

before the water treatment.

Du Sablon135 concluded that when starch disappears in late

fall much of it is converted into reserve cellulose which is de¬

posited on the inner side of wood-cell walls. In some cases this

lining was found to be comparativey thick and occasionally it

even had folds extending into the lumen of cells. It is said to

be readily soluble in dilute hydrochloric acid.

Schellenberg136 made a more thorough study of the deposi¬

tion and partial solution of hemicellulose in the wood and bark

of trees. He found a hemicellulose lining on the walls of fibers

in both spring and summer wood of Aesculus Hippo cast anum,

Betula and other trees but it was not dissolved in spring. Since

similar hemicellulose linings in the cells of the phloem and corti¬

cal parenchyma were found corroded in spring he concluded

that the lining did not dissolve in the fibers because protoplasm

was absent there. In the wood fibers of Yitis and Robina

Pseudacacia he noted the occurrence of especially thick hemi-

celulose layers in well matured wood and of much thinner ones

in those of immature wood. The protoplasm remains alive in

the wood fibers of Yitis and he accordingly found the inner lay¬

ers corroded and dissolved in spring. He also found the same

solution of the inner unlignified layers in the bast fibers and

cortica Iparenchyma and collenchyma of Fraxinus excelsior.

Usually from a third to half of the unlignified layer in the cor¬

tical parenchyma is dissolved when the buds open. He was of

the opinion that the deposition of hemicellulose in the bark

parenchyma continues after the leaves fall.

From these papers it is evident that a hemicellulose dissolv¬

ing enzyme is active during the early part of a vegetative sea-

134 Potter, M. C. On the occurrence of cellulose in the xylem of woody

stems. Ann. Bot. 18: 121-40. 1904.

186 1. c.

188 Schellenberg, H. C. Ueber Hemicellulosen als Reservestoffe bei un-

sern Waldbaumen. Ber. Deut. Bot. Ges. 23: 36-45. 1905.

68 Wisconsin Academy of Sciences , Arts, and Letters.

&

son and that such an enzyme is not present or is inactive in the

latter part of the growing period as indicated by the fact that

hemicellulose is deposited in both the wood and bark at that

time. Sanio137 found that in Finns silvestris lignification did

not occur in spring wood until after the deposition of the secon¬

dary thickening had been completed, that it began at the angles

of the cells and then involved the radial walls and later the tan¬

gential walls. In the summer wood, however, the primary walls

were found to have lignified before the deposition of the secon¬

dary thickening began, and it occurred in cells which were only

a few removed from the cambium. The final composition of

the cell walls of spring and summer wood seem also to differ, for

according to Wider, 138 the walls of spring wood contain a lower

percentage of cellulose than those of summer wood.

If the deposition and lignification of cellulose are in any way

dependent upon enzymotic action, there must be at least two

enzymes concerned because the two processes appear to be inde¬

pendent of each other as indicated by Sanio ’s observations. It

is evident that either of the processes would necessarily impede

or check further enlargement of cells differentiating from the

cambium. It, therefore, appears permissible to assume that the

enzymes involved in the solution of hemicellulose and the tardi¬

ness of the lignification process in spring are important factors

in permitting the development of larger wood cells in spring

than those produced in summer, when the cellulose dissolving

enzymes are inactive and lignification occurs so quickly after a

cell is formed that in some cases it takes place even before sec¬

ondary thickening has begun. The experiments by Jost and by

Lutz also give support to the idea that radial growth is largely

controlled by enzymotic activities which are somehow dependent

upon the process of terminal elongation. Perhaps the enzymes

concerned are liberated or activated in enlarging and bursting

buds in different parts of trees and are carried downward in the

metabolized food, or possibly enzymes produced in the enlarging

buds simply initiate certain activities which are transmitted

without the further aid of the enzymes as was assumed by

137 Sanio, K. Anatomie der gemeinen Kiefer (Pinus silvestris L.).

Jahrb. Wiss. Bot. 9: 66-68. 1873.

188 1. c.

Grossenbacher — Radial Growth in Trees.

69

Fick139 regarding the action of the enzymes which coagulate

blood and milk.

The fact that stems and branches of trees are more pliable and

easily bent while in the midst of active spring growth than they

are at any other time, indicates that perhaps some enzymotic

softening of the mature wood occurs during the period of most

active growth. The upward bending of a branch on a decapi¬

tated conifer also argues for the presence of some softening

agent during the time of most vigorous growth because of the

fact that such branches often bend in response to gravity at

places where lignification had previously occurred. In other

words, it seems that one of the most important factors in the

production of large wood cells in spring and smaller ones in

summer may be the presence of enzymes which retard lignifica¬

tion and prevent rapid thickening of the walls and thereby per¬

mit growth or hydrostatic pressure to develop large cells in

spring; while the absence or inactive condition of those enzymes

induces rapid thickening and early lignification of the walls in

summer and thus checks the enlargement of summer-wood cells.

It may be that the idea of growth force expressed by Detlef-

sen, Mer and others as well as “the impulse to grow” em¬

phasized by Morgulis imply the same sort of notion as that ad¬

vanced in the above scheme regarding the possible relation of

enzymes to ring formation, but in any case the hypothesis is

only a guess based on rather suggestive indirect evidence. Mer’s

conclusion that the winter rest of the cambium induces its

greater activity in spring seems to have something in common

with the outcome of some feeding experiments by Morgulis, to

the effect that in subjecting salamanders to alternate periods of

fasting and liberal feeding a greater growth resulted than by

more frequent and abundant feedings. A theory to account for

wood rings must also make use of the evidence brought out re¬

garding the effect of variations in bark tension both longitudinal

and transverse, as well as of the influence of the transpiration

stream as suggested by Hartig and more recently elaborated by

Jaccard in his discussion of the distribution of radial growth.

It should be remembered, however, that transpiration is per¬

haps greater during the time summer-wood is formed than it is

while spring wood develops; to say that larger cells are pro¬

duced in spring to meet the higher water requirements of the

approaching summer explains nothing.

139 Pick, A. Ueber die Wirkungsart der Gerinnungsfermente. Archiv.

Gesam. Physiol. Mens. Thiere. 45: 293-96. 1889.

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Morgnlis, S. The influence of protracted and intermittent fast¬

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75

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76 Wisconsin Academy of Sciences, Arts, and Letters.

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78 Wisconsin Academy of Sciences, Arts , and Letters.

NOTES ON PARASITIC FUNGI IN WISCONSIN-!.

J. J. Davis.

These notes are intended to be supplementary to ‘ ‘ A provisional

List of Parasitic Fungi in Wisconsin” published in Trans¬

actions of the Wisconsin Academy of Sciences, Arts and

Letters. Vol. XVII, pt. 2, pp. 846-984.

Plasmopara Jiumuli Miyabe & Takahashi. This was collected

on wild Humulus Lupulus growing along the river bank at

Racine in 1909-10 since which time the station has not been

visited. The following notes of this fungus were made at

Racine: Spots small, angular at first, limited by the vein-

lets, brown-red or purplish above, below of a darker green

than the leaf, giving the “water soaked” appearance. The

spots are surrounded by an indeterminate yellowish discoloration

especially early in the season, less marked as the leaves become

firmer, and finally assume the lethal brown with the death

of the tissues included. Conidiphores hypophyllous, grey,

17 5-325x5-61/2/a with usually two lateral branches each of which

is about equal in development to the terminal portion and 1-3

times branched, ultimate branchlets tapering, subacute ; conidia

fuligineous tinted, elliptical, somewhat acute at each end, fur¬

nished with an apical papilla of dehiscence, 20-33 x 12-17/x, usu¬

ally about 26 x 15 /x ; oospores scattered in the leaves ; oogonia ir¬

regularly thickened, brown, subglobose, 36-40/x long, oospores

filling the oogonia 30-33 y long.

Aster ina plant ag inis Ellis. This is referred to Mycosphaer-

ella by Theissen (Ann. Mycol. 10:2:196. (Apr. 1912).

Asterina rxibicola Ell. & Evht. This is described by Theissen

in the same communication (p. 195) but no new combination is

proposed.

Davis — Parasitic Fungi in Wisconsin — I.

79

Gnomcmiella fimbriata (Pers.) Sacc. This was inserted in the

provisional list because of an immature specimen in the herbar¬

ium of the University of Wisconsin which is perhaps of this

species. It was collected at Osceola by E. Sheldon in 1892.

Phyllosticta destruens Desm. In writing the provisional list I

followed Ellis & Everhart (North American Phyllostictas, 40) in

referring to this species specimens on Prunus virginiana and al¬

so on Amelanchier. 'The former has been described under the

name Phoma virginiana Ell. & Hals. (Journ. My col. 4:8. (1888) )

the latter as Phyllosticta innumerabilis Pk. (Bull. Torr. Bot. Club.

36:336 (1909)). Specimens on Amelanchier were distributed

under the name Phyllosticta destructens Desm. in Fungi Colum-

biani continued 1447. I have seen no European specimens of

Ph. destruens Desm. which is said to occur on Celtis as well as

Prunus and to have epiphyllous pycnidia, but I infer that Mr.

Ellis had good reason for using that name. Before the list was

printed I removed Amelanchier as a host of Phyllosticta

destruens Desm. with the intention of inserting Ph. innumerabilis

Pk. an intention which I failed to carry out, so that Amelanchier

as bearing the Phyllosticta appears only in the host index. Mor¬

phologically I see no distinction between the fungi on the two

hosts.

In the provisional list of parasitic fungi in Wisconsin Pa-

touillard is given as the author of the binominal Protomyces

andinus as is done in the Sylloge Fungorum. An examination

of Patouillard ’s paper however shows that it was published as

Protomyces andinus Lagh. sp. nov. Lagerheim, not Spegazzini,

collected the type material in Ecuador, not Chili.

Phyllosticta mulgedii Davis, a name that was proposed in the

4th supplementary list (No. 709), was omitted from the provi¬

sional list. The fungus has not been collected again and is prob¬

ably one of the Phomae that have been described as occurring on

the leaves of Compositae.

Phyllosticta desmodii Ell. & Evht. This was described ( J ourn.

My col. 5: 146: 1889) from a single small collection in Walworth

Co. Much better material has been collected at Madison on Des-

modium canescens. The pycnidia are epiphyllous, brown, sub-

spherical, 125-160/* in diameter ; sporules oblong, often somewhat

80 Wisconsin Academy of Sciences, Arts, and Letters.

narrower in the middle, ends rounded, conspicuously 2-4 guttu-

late, 6-12x3y. The appearance of the sporules suggests that

later they may become septate.

Phyllosticta cruenta (Fr.) Kickx. The reference of this species

to Macrophoma as proposed by Ferraris (Ann. My col. 10:3:288

(Jn. 1912)) would make a generic distinction between the two

forms that were given as varieties in the provisional list because

the globose sporules of Ph. pallidior Pk. are only about 10^ in

length, although they equal in content the longer and narrower

sporules of the type of PJi. discincta.

Ascochyta pisi Lib. has been shown by R. E. Stone to be a con-

idial form of Mycosphaerella pinodes (Berk. & Blox.) Niessl

(Ann. Mycol. 10: 564 et seq. (Dec. 1912). Also R. E. Vaughan,

Phytopathology 3:11 [1913].

Actinonema rosae, (Lib.) Fr. Diedicke calls this Marssonina

rosae (Lib.) Trail (Ann. Mycol. 10: 146 (Apr. 1912) ). F. A.

Wolf has developed the ascosporous stage which he refers to the

Microthyriaceae and makes the type of a new genus and calls it

Diplocarpon rosae F. A. Wolf. (Bat. Gaz. 54, 231 (Sept. 1912) ).

Septoria nubilosa Ell. & Evht. (Proc. Acad. Nat. Sci. Phil.

1891, p. 76) which was founded on Wisconsin material on Eele-

nium autumnale has not been included in the Wisconsin lists be¬

cause it is merely a form of Septoria helenii Ell. & Evht. in

which the spots are not well developed. The type was collected

on the north side of plants bearing typical S. helenii and was

sent to the authors merely to show the variation.

Septoria ribis Desm. Some of the specimens that I have re¬

ferred to this species are perhaps S. grossidariae var. longispora

Ferraris (Ann. Mycol. 10:291). Typical S. grossulariae (Lib.)

West. I have collected but once.

Septoria saccharina Ell. & Evht. Specimens from Price Co.

bear sporules about 30x2%/*. The Acer — inhabiting fungi,

having triseptate sporules borne in acervuli and varying

in length from 20-70/*, and in width from 1%-5/a seem to me to

constitute a group the relation of the members of which can

be determined by inoculation methods only. The form with short

and thick sporules has been called Ascochyta aceris Lib. and later

Davis — Parasitic Fungi in Wisconsin — I.

81

Phleospora aceris (Lib.) Sacc. and with this have been included

narrower spored forms but usually the latter have been referred

to Septoria. Diedicke has recently referred the European forms

with sporules 3 /a or less in thickness to Cylindrosporium, recog¬

nizing three species. (Ann. My col. 10: 486 (Oct. 1912) ). Sep¬

toria saccharina E. & E., however, while producing similar spor¬

ules develops pycnidia in definite arid spots but Cylindrospor¬

ium saccharinum Ell. & Ev. agrees with the form on Acer rub-

rum which was recorded in the “provisional list” under the

name Phleospora aceris (Lib.) Sacc.

Septoria musiva Pk. The specimens of Septoria that have been

collected on Populus in Wisconsin may be divided into groups as

follows :

1. Spots subcircular to subangular, black becoming white and

arid except the peripheral portion, 3-5 mm. in diameter ; pycni¬

dia epiphyllous, superficial-collapsing, hemispherical in section,

about 100/a in diameter; sporules cylindrical, curved, obtuse,

3(2-4) septate, 25-60 x 2-3 /a. On Populus bdlsamifera.

2. Similar to the above except that the central portion of the

spot becomes alutaceous or cinereous instead of white and the

pycnidia are more scattered and lie deeper. Also on P. balsami-

fera.

3. Spots subcircular to angular, limited by the veinlets, at

first brown, becoming grey by the loosening of the cuticle, 1-3

mm. in diameter, becoming confluent into larger areas, %-l cm.

in diameter; pycnidia innate, discharging on either surface but

usually below, having a thin but distinct wall and a large open¬

ing, 70-100/a in diameter; sporules cylindrical, curved, obtuse,

3 (2-4) -septate, 25-65 x 2-3/*. On Populus deltoides.

4. Spots roundish to irregular and angular, dark brown above

becoming grey with age, light brown below, 2-5 mm. in diameter,

often confluent; pycnidia hypophyllous, punctiform; sporules

cylindrical, curved, obtuse, 3-5 septate, 45-65 x 3/a. On P.

balsamifera.

5. Spots angular, blackish brown above, paler below, becoming

lighter and mottled with age, %-l cm. in diameter; pycnidia

scattered, innate, thin walled ; sporules filiform, narrowed to one

end, 3-6 septate, 40-70x11/2~2%/a. On Populus tremuloides.

In the provisional list all of these forms were included in Sep¬

toria musiva Pk., although group 5 may prove to be distinct, but

6— S. A.

82 Wisconsin Academy of Sc'ences, Arts , and Letters.

one can hardly decide that on morphological grounds without an

abundance of material. I find this in Wisconsin only to the

north and with few pycnidia. Much the best specimens that 1

have seen were collected at North Yakima, Wash. (Wis. Acad.

Sci. Arts & Lett. 15 :778).

In Farlow’s Host Index, Populus balsamifera is given as a

host of Septoria salicina Pk. I presume that this refers to what

I have called group 1. This form looks quite distinct but it

merges into group 2 and that again into group 3 in a way that

makes it difficult to draw a line of separation, while the charac¬

ters of the sporules are identical. Septoria salicina Pk. differs in

its uniseptate sporules. Of Septoria populi Desm., which has

unisept ate sporules, I have seen no American specimens. Fungi

Columbiani 2873, issued under this name seems to be the same

as 2872 which is labeled Septoria musiva Pk. On neither have

I found a Septoria. I may mention that F. Col. 1587 on Populus

tremuloides collected by J. B. Ellis at Newfield, N. J. and issued

as S. musiva Pk. bears Marssonina rhabdospora (E. & E.) Magn.

at least in the two copies which I have seen. F. Col. 3486 on

Populus balsamifera collected at St. Johnsbury, Vt. by W. P.

Carr and issued as Septoria populi Desm. is of the form with

black bordered alutaceous spots that I have placed in group 2.

Pacific Slope Fungi 1723 on Populus Fremonti collected in Cali¬

fornia by Copeland and issued by Baker as Septoria populi Desm.

is somewhat intermediate between groups 2 and 3. Fungi Col¬

umbiani 1257 on Populus angustifolia collected at Golden, Colo¬

rado, by Bethel, and issued as Septoria populi Desm. has subcir¬

cular spots of a yellowish white or sordid white color with an ir¬

regular grey-brown border 2-4 mm. in diameter ; the pycnidia are

hypophyllous, broad and collapsing ; the sporules continuous 23-

35 x 31/2-4/*. While this does not correspond with S. populi

Desm. it is different from any Septoria on Populus that I have

seen and judging from the single specimen may prove to be dis¬

tinct. Cylindrosporntm oculatum Ell. & Evht. on Populus del-

toides (Put-in-Bay, Ohio) has hemispheric-superficial pycnidia

and obtuse sporules 30-50 x 3/* becoming 3 or more septate. This

forms circular grey to sordid spots about % cm. in diameter with

a narrow dark border. I would include it in Septoria musiva Pk.

as representing forms 1 and 2 on Populus deltoides. Specimens

Davis — Parasitic Fungi in Wisconsin — I.

83

on this host collected at Ithaca, N. Y., by Higgins (com. Barthol¬

omew) bear both this and form 3 on the same leaves.

After this was written a collection was made from the exami¬

nation of which the following characters were noted : spots angu-

lar-suborbicular, at first brown with a narrow darker margin,

then grey and finally mottled with small angular cream colored

areas, sometimes confluent, 2-3 mm. in diameter ; pycnidia mostly

hypophyllous and inconspicuous, 65-75/*, in diameter; sporules,

hyaline, filiform, acute, 3-6 septate, 38-60 x 21/2-S1/2y. On Popu¬

lous grandidentata. Devils Lake, Wisconsin, Aug. 6, 1913.

In this connection a still more recent (Aug. 21, 1914) collec¬

tion on Populus deltoides is of interest. Phyllosticta populina

Sacc. is said to occur in association with Septoria populi Desm. in

Europe. Having made a collection of the former at Prescott the

associated Septoria was examined with some interest. The spots

are orbicular, cinereous with a narrow dark margin and re¬

semble those of form 2 except in the grey color which in the

older spots changes to white. On some of the leaves are small

angular, confluent spots like those of typical Septoria musiva

Pk. Of the first mount from this material it was noted “spor-

ules mostly 18-22 x 2-3/*, 1-2 septate with occasional longer ones

up to 48/*. and 3-septate ” ; of another mount ‘ 4 3CM5 x 2-3/*,

2-3 septate”. This seems to connect with the forms described

above and suggests that there is a widely variable species of

Septoria occurring on Populus in both America and Europe.

Septoria Candida (Fckl..?) Sacc. I have not seen, but the des¬

cription indicates that it might readily fall in with the Ameri¬

can forms.

Cercospora geranii Kell. & Sw. Of a specimen collected at

Blue Mounds the following notes were made. Hyphae usually

straight, slightly colored, often toothed, 25-75 x 6-7/*; conidia

hyaline, cylindrical, usually more or less curved, obtuse, becom¬

ing pluriseptate, 100-165 x 4-5/*.

Cercospora subsanguinea Ell. & Evht. is sometimes devoid of

color and the obtuse conidia sometimes divide in the middle. It

appears to be more nearly a Ramula/ria.

Gloeosporium fragariae (Lib.) Mont. My notes of the

measurements of the sporules of the fungus referred to this spe¬

cies range from 12-24 x 4-5/*. It was collected at Spooner.

84 Wisconsin Academy of Sciences , Arts, and Letters.

Gloeosporium rihis (Lib.) Mont. & Desm. As it occurs in Wis¬

consin this usually has the characters of the forma ribis nigri

americana Sacc. The sporules sometimes reach 30 /*, in length.

Gloeosporium tremuloides Ell. & Evht. 2nd suppl. list no. 526

was omitted from the provisional list because of the belief that

the species was founded on imperfectly developed material of

Marssonina castagnei (D. & M.) Magn. which occurs in atypical

forms in Wisconsin. Oudemans proposed the variety monili-

ferae in which the acervuli are amphigenous although more abun¬

dant above. In Wisconsin they are often hypophyllous only and

the sporules are often but 12-15/*, long. Marssonina brunnea (E.

& E.) has been omitted, being considered, perhaps erroneously,

a form of M. castagnei.

Ramularia plantaginis Ell. & Mart. In the description of this

species the spots are said to be minute. Specimens on Plantago

Rugelii collected at Madison in September have spots up to 3 cm.

in diameter. Conidia appear also on the ealyces.

Ramularia alismatis Fautrey. This was reported in the third

supplementary list under the name As cocky t a alismatis (Oud.)

Trail. Dr. R. A. Harper has kindly compared Wisconsin ma¬

terial with the type of Ascockyta alismatis Ell. & Evht. in the

Ellis herbarium and finds them to be the same. The very short

undifferentiated conidia-bearing hyphae makes this an atypical

Ramularia. It is not unlikely that Septoria alismatis Oud. is of

the same character. The spots usually have a slight eminence

in the center as if a pycnidium lay beneath. (See Diedicke,

Ann. Mycol. 10:479).

Ramularia uredinis (Yoss) Sacc. This is the fungus recorded

in the supplementary and 3rd suppl. lists no. 330 under the name

Fusarium uredinum E. & E. The tufts are sometimes pink or

even testaceous. My measurements of the conidia, which are in

branched chains, are from 7-17x3-4^.

Ustilago osmundae Pk. This has been collected on Osmunda

regalis in Washburn and Burnett counties. I have not been able

to follow the author of the species in his reference of it to My-

cosyrinx. (New York State Museum; Report of the Botanist

1911, p. 43). When the fungus is present each frond arising

from the rhizome bears the smut or else is sterile.

Davis — Parasitic Fungi in Wisconsin — I.

85

Ustilago lorentziana Thuem. which occurs at Madison on Hor-

deum jubatum and which was recorded in the 4th supplementary

list seems to have been omitted from the provisional list.

Entyloma linariae Schroet. var. veronicas Wint. The newly

formed spores of this smut were found to germinate readily in

May but to gradually lose the power as the season progressed as

had been found to be the case with E. floerkeae Holw. (2nd

suppl. list, No. 487). The promyeelial spores are usually two

(1-4) in number, 15-20 x 3y.

Material wintered outdoors (May to May) germinated the fol¬

lowing spring in the same manner.

Additional Hosts.

Synchytrium aureum Schroet.

In September, 1912, this was found at Millston, Jackson

county on Lycopus virginicus, Lysimachia terrestris and leaves

of blackberries that I have referred to Rubus hispidus and Ru-

bus villosus. The infection was sufficient to indicate that each

of these plants are normal hosts of the organism in that locality,

Rubus villosus being least affected. No success attended spe¬

cial efforts to find other hosts. In 1913 it was collected at Athel-

stane, Marinette Co., on Rubus hispidus but on no other host.

In 1892 Synchytrium occurred rather abundantly in a bit of

woodland near Berryville on Viola pubescens and Geum cana-

dense and during the same season it was collected at Somers,

but a few miles distant, on Ranunculus recurvatus. The infec¬

tion of the latter was limited and I have not seen it since on

this host. It was' collected again at Berryville in 1894 soon

after which the station was cleared and put under cultivation.

In 1902 a collection on Viola pubescens was made at the Somers

station. In 1907 considerable infection of the same host was

observed at a station intermediate between the other two and

during the same season very limited infection of Prenanthes

alba at this station and of Pedicular is canadensis near Ka-

cine was observed. The infection of the two latter hosts ap¬

peared to be accidental and temporary, the organism failing to

get a permanent foothold. At Millston some of the affected

leaves of Lycopus bore considerable hypertrophies often sur¬

rounded by purple discoloration but usually there was little dis-

86 Wisconsin Academy of Sciences , Arts, and Letters.

tortion of the hosts, even when the sori were numerous and ag¬

gregated, the pressure being into the mesophyl which was some¬

times torn from the epidermis in the area surrounding the gall.

The common factors which make for susceptibility in these

various hosts are not apparent to me.

Septoria astericola Ell. & Evht. on Aster puniceus. In the

specimen on this host the spots become lead color with a dark

border. The largest spots are 1 cm. long. The pycnidia are

epiphyllous, about 80/a in diameter and the sporules 23-33 x 1/a.

Collected at Lake Mills, Oct. 19, 1912.

Gloeosporium saccharinum, Ell. & Evht. Specimens on Acer

spicatum collected at Spooner have circular spots of a pale olive

color with a darker border; the largest sporules are 7x3/a. The

fungus often developes on subcircular spots of a tan color on

Acer Saccharum.

Cercospora caricina Ell. & Dearn. My notes of a specimen on

Cyperus filiculmis collected at Madison, Aug. 12, 1912, are as fol¬

lows: Hyphae 3-8 in a tuft, brown, somewhat nodulose, often

denticulate at the apex, 50-80x3-4y ; conidia hyaline, obclavate-

cylindrical, straight or curved, becoming pluriseptate, 65-lQ0x

3- 4/a. On bracts and culms, spreading from above downward.

Cercospora caricina Ell. & Dearn. is described as having hyphae

15-25 x 3-3 %/a and conidia 34-73 x 3/a, but I have specimens on

Carex in which the hyphae and conidia equal those noted on

Cyperus. Vyperus Houghtonii which was tentatively given as a

host in the 4th suppl. list should not have been omitted from the

provisional list.

Cercospora ceanothi Kell. & Swingle. On Ceanothus ameri-

canus. Madison. In one of the collections on this host the fungus

is particularly well developed, the conidiophores being 20-45x

4- 5/a and the attenuate conidia 80-150 x 4-6/a. A collection made

in the same locality two weeks later agrees with the description

of Cercospora fuliginosa, E. &. E. the conidia being darker,

cylindrical and 30-80 long. It is probable, therefore, that the

descriptions of C. ceanothi Kell. & Swingle and C. fuliginosa Ell.

& Evht. were drawn from different states of the same fungus.

The former is the prior name and the latter is antedated by C.

fuliginosa Ell. & Kell, on Diospyros (1887) for which reason C.

MacClatehieana Sace. & Syd. was substituted.

DaAjis — Parasitic Fungi in Wisconsin — I.

87

Additional Species.

Leptosphaeria folliculata Ell. & Evht. var. oxyspora n.

var. On Garex gracillima. Price Co. Sept. 9, 1911. Differs

from the type in the somewhat narrower asci ( ca . 50x8/a) and

especially in the triseptate acnte ascopores (ca. 15x3/a). On

the comparatively narrow leaves of this host the perithecia are

borne on dead apical areas at the bases of which there is often

evidence of primary spotting and confluence. I am indebted to

Dr. R. A. Harper for comparison of this with the type in the

Ellis herbarium at the New York Botanical Garden. It is not

unlikely that sufficient material would connect these forms with

L. caricicola Fautr. and L. caricina Schroet.

Phyllosticta livida Ell. & Evht. On Quercus macrocarpa,

Millston, Jackson Co. In these specimens the pycnidia are hy-

pophyllous. If they really represent this species the fungus

has a wide distribution in the U. S. previous collections being

reported from California and Florida.

Phyllosticta liatridis n. sp. Spots round, white or sordid,

arid, 1-2 mm. in diameter, usually surrounded by a broad black

border ; pycnidia epiphyllous, prominent, black, about 65/a ;

sporules hyaline, oblong, 2-4 nucleolate, about 10x4/a. On Lia-

tris spicata, Gaslyn, Burnett Co. Aug. 1, 1911. This can hardly

be Phoma minutissima Cke. as that species is described.

Diplodia tjvulariae n. sp. Spots oval to orbicular, white,

thin and arid, usually with a ferruginous border, 8-15x5-10mm. ;

pycnidia mostly epiphyllous, scattered, black, globose, 100-150/a;

sporules elliptical to ovate, brown, uniseptate, 12-20x6-7/a. On

TJvularia (Oakesia) sessilifolia Spooner, Aug. 15, 1911 (type)

and Gaslyn. What is probably imperfect material of this spe¬

cies has been collected at Blue Mounds on TJvularia grandiflora.

It is not unlikely that North Am. Fungi 2153 issued under

the nomen nudum Phyllosticta uvulariae Galloway is of this

character. (See 4th suppl. list under No. 359.) Macroscop-

ically this fungus suggests Phyllosticta cruenta (Fr.) Kx. Oc¬

casional biseptate sporules occur as is to be expected.

Stagonospora intermixta (Cke.) Sacc. Price Co., Oct. 9,

1911. On leaves of Cinna arundinacea. I have not seen an

88 Wisconsin Academy of Sciences , Arts , and Letters.

authentic specimen of this species. The specimens which I have

referred here have depressed-globose pycnidia 40-60/* in di¬

ameter with a round apical pore which is surrounded by a thick

black ring. The long-fusoid sporules are 7 — septate, 40-60x

31/2-5/*.

Septoeia andropogonis, n. sp. Causing narrow elongated

areas of a reddish-yellow color sometimes becoming sordid;

pycnidia epiphyllous, subseriate or scattered, dark brown, de¬

pressed globose, little prominent, 75-100/*; sporules hyaline,

straight or slightly curved, more acute at one end, becoming 2-4

septate, 30-50x2-3/*. On leaves of Andropogon furcatus, Gas-

lyn, Burnett Co. July 31, 1911.

Septoria polita n. sp. Pycnidia scattered, globose, innate,

black, ostiolate, 65-100/*; sporules hyaline, straight or somewhat

curved, truncate to obtusely rounded at each end, becoming 3-5

septate, 35-50x2%-3/*. On Car ex sp. indet. (stellulata? ) Gas-

lyn, Wisconsin, Aug. 4, 1911. This attacks the distal portion

of the very narrow leaves of the host which becomes sere. The

sporules have a very smooth or polished appearance and are not

at all constricted at the septa.

Septoria carpinea (Schw. ?) n. comb. Spots subcircular

to angular, numerous, reddish brown becoming sordid in the

center, somewhat paler below, 1-5 mm. in diameter; pycnidia

epiphyllous, few, scattered, prominent, black, globose, ostiolate,

about 65/*; sporules hyaline, usually curved, frequently arcuate,

pluriguttulate, 25-40x2-3/*. On Carpinns caroliniada, Gaslyn,

Wisconsin, Aug. 8, 1911. It seems quite possible that this is the

fungus called Xyloma by Schweinitz and Depazea by Fries.

Septoria Polymniae Ell. & Evht. The specimens on Polym-

nia canadensis, collected near Somers in 1903, which I hes¬

itatingly refer to this species show suborbicular spots %- 1 cm. in

diameter which become brown above, darker toward the margin.

The pycnidia correspond with those of this species. My notes

of the size of the sporules read 40-45x1 V2-2/*.

Sacidium microspermum (Pk.) n. comb. (Septoria micro-

sperma Pk.) On fallen leaf of Betula alba papyrifera. But¬

ternut, Oct. 8, 1911. Hypophyllous on indefinite brown areas

which show a tendency to extend along the veins; basidia and

Davis — Parasitic Fungi in Wisconsin — I.

89

sporules in a discoid layer 100-150/* broad which is covered by

a chitinoid, p undulate clypeus which becomes irregularly fis¬

sured; sporules straight or allantoid, 6-10 x %-l%/*. North

American Fungi 674 on Betula lent a, collected by Nuttall in

West Virginia, shows faded leaves with circular green areas.

The pycnidia, however, are by no means confined to the green

spots. In the West Virginia specimens also the sporules are

smaller than in the type as described. I assume that it repre¬

sents it in its Sacidium structure. Perhaps this is not distinct

from Leptothyrium betulae Fckl.

(This has since been collected on the same host at Wausaukee.)

A Gloeosporium which has appeared in the greenhouse of the

botanical department at Madison on the leaves of Dendrobium

moschatum causes orbicular arid spots about 1 cm. in diameter

with a dark purple border and elevated margin. The acervuli

are brown, scattered, mostly epiphyllous; the sporules oblong

to ovate-oblong, obtuse at both ends, biguttulate, KKL5x4 /*. Prob¬

ably this is Gloeosporium cinctum B. & C. and perhaps also Gl.

pallidum Karst. & Har. The studies of Shear and Wood, how¬

ever indicate that it is a conidial condition of Glomerella cingu-

lata (Stonem.) S. & V. S. (U. S. Dept, of Agr., B. P. I., Bull.

252).

Colletotrichum helianthi n. sp. Spots definite, orbicular,

olivaceous with a cinereous center and a black margin, paler be¬

low, often confluent, 3-5 mm. in diameter; acervuli very promi¬

nent, one or few on a spot, 50-65/* broad, surrounded by black

rigid bristles 80-150x3-5/* which taper from base to apex ; spor¬

ules hyaline, fusiform to arcuate, nucleolate, acute at each end,

25-35x2%-3%/*. On Helianthus sp. indet. Madison, Wisconsin,

July 7, 1907. This is allied to C. solitarium Ell. & Barth, from

which it differs in the larger bristles and sporules. I found the

specimen in the herbarium of the University of Wisconsin with

the name of the collector not given

Ovularia asperifolii Sacc., var. lappulae n. var. Spots sub-

orbicular, dark brown, %-l cm; conidiophores hypophyllous,

scattered or in tufts of 2—4, hyaline, often toothed, usually 16-

20x3-4/*; conidia in chains which are sometimes branched, hya¬

line, 6-18x3-4^; the lower conidia are cylindrical, acute at each

end, 12-18x3-3%/*, the upper fusoid, 6-12x3-3%/*. Much

9*0 Wisconsin Academy of Sciences , Arts , and Letters.

longer hyphae ( ca . 50/*) have been observed bearing conidia

singly and laterally. On Lappula virginiana. Somers, Ra¬

cine and Blue Mounds. While this fungus causes conspicuous

spotting of the leaves the conidia are inconspicuous and evan¬

escent. I have had it under observation for a number of

years expecting at some time1 to secure specimens with a more

profuse development of conidia. A specimen collected at Blue

Mounds, July 13, 1912, is taken as the type. A more recent col¬

lection made at Potosi bears conidia up to 30/x in length. The

Wisconsin fungus seems to be closely allied to var. symphyti-

tuberosi, Allesch. ( Hedwigia , 1894, p. 73.) These specimens

differ from Hermodendron farinosum Bon. as figured (Bot.

Zeit., t. VIII, fig. 9) in the longer and narrower conidia and the

absence of the two guttulae in the lower members of the chain.

During August and September, 1912, there was collected at

Madison on leaves of Ribes americanum a fungus of which the

following notes were made : ‘ ‘ Spots angular, limited by the vein-

lets, often confluent into irregular areas, brown, 2-5 mm. in di¬

ameter; conidiophores hypophyllous in scattered tufts, closely

fasciculate from a prominent sclerotioid base, hyaline, often

toothed, 30-65x2-3/* ; conidia terminal and lateral, hyaline,

cylindrical, abruptly acute or rounded at each end, occasionally

with a median septum, 20-50x3-4 /*. The tufts usually have

more or less of a pink tinge. Large fasciculi have a marked

stilboid appearance. ’ 9 Leaves bearing the fungus were wintered

out of doors and the following May were found to bear heads of

conidia up to 250/* or more in diameter of a vinous purple color

with the conidiophores compacted into blackish stipes usque 150/*

high each springing from the summit of a plectenehymatous

pseudopycnidium. The conidia borne on these heads were hya¬

line, catenulate, fusoid, continuous, 10-18x3^/*. With these

were fasciculi, snow white to purplish, of the mucedine type and

occasional broader ones more tubercularioid in appearance.

Accepting the coremium structure as the climax development

of this fungus I have labeled the specimens Graphiothecium

vinosum n. sp. and as it appears to be at least a facultative par¬

asite have given it a place in this list.

Ramularia calthae Lindr. Specimens having the follow¬

ing characters have been referred to this species. Spots small,

Davis — Parasitic Fungi in Wisconsin ■ — I.

91

angular, immarginate, limited by the veinlets, becoming conflu¬

ent, brown, more abundant near the margin of the leaf ; conidio-

phores epiphyllous, tufted from a stromatoid base, erect, simple,

hyaline, 15-30x114-2//,; conidia similar, sometimes catenulate

12-24xl-l3/2(u. On Caltha palustris. Gaslyn, Burnett Co.,

Aug. 30, 1911.

Cercgsporella exilis n. sp. Spots round to angular, lim¬

ited by the. veinlets, often confluent, brown, 2-5 mm. ; conidio-

phores in small loose tufts which are effused over the lower sur¬

face of the spots, hyaline, continuous, usually subulate, nearly

straight, seldom branched, 1 0-20x2 *4-3 \ conidia cylindrical,

straight, hyaline, continuous or obscurely septate, 20-40x1-2//,.

On Phryma Leptostachya. Madison, Blue Mounds and Devils

Lake, August and September.

Cladosporium paeoniae Pass. On Paeonia (cult.) Madison.

Pending an investigation of the diseases of paeonies in the

United States, which I am informed, is to be made, I use this name

provisionally.

Cladosporium gloeosporioides Atk. On Hypericum virgin-

icum. Grand Rapids (Peltier) and Madison. This forms def¬

inite alutaceous spots on the leaves. When,; however, the host

plants are in a thick overshadowing growth of Carices and other

taller plants the hyphae are borne on indefinite discolored areas.

Frequently all gradations may be seen on a single host; on the

upper leaves, exposed to the sunlight, the hyphae being confined

to definite tan colored spots while on the lower they are borne

on indefinite subolivaceous areas. I find the length of the hyphae

variable ; in some specimens 20-30//,, in others ca. 60/x. Dr. R. A.

Harper has kindly compared this with specimens of Gloeospor-

ium cladosporioides Ell. & Hals, in the Ellis Herbarium and tran¬

scribed the following notes from the inside of one of the enve¬

lopes; “Hyphae 35-40x4//,, fasciculate, nodulose above, hyaline

becoming dark; conidia oblong-elliptical 10-14x4-6 microns”.

Dr. Harper writes :

“The spores seem like those on your material but the fungus

on Halsted’s material seems to be almost if not entirely on the

stem. His host plant, of course, has narrow leaves quite differ¬

ent from yours. I did not get a good preparation of the conidio-

phores; I should think the two might be the same but I am

92 Wisconsin Academy of Sciences , Arts , and Letters.

doubtful. The New Jersey fungus is certainly not as conspicu¬

ous as yours and produces no such leaf spots. ’ ’

Considering the differences in the hosts it seems to me that

there is a variable Cladosporium on Hypericum to forms of

which these two names were applied. If that is the case I would

prefer the later name here used to avoid tautology.

Cercospora fingens n. sp. Spots suborbicular, immargin-

ate, blackish brown, 3-5 mm. ; conidiophores hypophyllous,

olivaceous brown, somewhat crooked, denticulate, thicker and

paler toward the apex, pluriseptate, 130-250x4-6/*; conidia hy¬

aline, pluriseptate with a tendency to break apart at the septa,

somewhat flaccid, tapering upward, 100-215x3-5/*. On Thalic-

trum dasycarpum , Burnett, Washburn and Price Counties,

July-September. On Thalictrum dioicum, Lone Bock, (R. A.

Harper and G. M. Reed) . Because of the long and slender hy-

phae and conidia this resembles, under a hand lens, Phytoph-

thora thalictri Wils. & Davis for which it was mistaken in the

field.

I was at first disposed to refer this to Cercospora aquilegiae

Kell. & Sw. but as no specimens have been collected on Aquil-

egia in Wisconsin, I infer that it is distinct.

Puccinia microsora Koern. Amphi-and teleuto-spores on

Car ex Tuckermani Price County and Carex scabrata, Bayfield.

Coleosporium sonchi-arvensis (Pers.) Lev. II, III, on Sonchus

asper, I on Pinrn sylvestris, Sturgeon Bay. The uredinia were

collected by Mr. J. 0. Sanders, Entomologist of the Wisconsin

Agricultural Experiment Station, who found it to be locally

abundant. The aecia usually appear upon but one of the paired

leaves.

Herbarium of the University of Wisconsin, Madison Wis¬

consin, March, 1913.

Davis — Parasitic Fungi in Wisconsin— II.

93

NOTES ON PARASITIC FUNGI IN WISCONSIN— II.

J. J. Davis.

This communication holds a supplementary relation to A pro¬

visional List of parasitic Fungi in Wisconsin which was pre¬

sented to the Wisconsin Academy of Sciences, Arts and Letters

in March, 1912, and published in its Transactions, volume

XVII pt. 2. After some notes of a miscellaneous character a

list is given of hosts and another of species additional to those

previously reported for Wisconsin. A eommunicaticn of simi¬

lar scope was presented to the Academy in April, 1913, and is

published herewith.

University of Wisconsin Herbarium, Madison, Wisconsin,

April, 1914.

Peronospora viciae (Berk.) D. By. In the provisional list a

question mark was placed after Vicia americana where given as

a host of this mildew. The reason for this is that the conidia of

the downy mildew collected on this host in Wisconsin are longer

than those of P. viciae on Pisum or on foreign species of Vicia

as far as I have examined them. All of the specimens on

native species- of Vicia X have seen are similar to those that

have been collected on V. americana in Wisconsin. 'The Wiscon¬

sin station is near Lake Mills and on the railroad right of way

where the mildew may have been introduced from the plains re¬

gion which seems to be the habitat of this form. Such evidence

as I have seen indicates that the American and European forms

are physiologically distinct. I would suggest that tfie native

form be distinguished as Peronospora viciae var. americana —

with conidia 30(24-36) x 20(17-26) and that Fungi Columbi¬

ana 1836 on Vicia linearis Stockton, Kansas, Bartholomew, be

taken as the type of this variety.

94 Wisconsin Academy of Sciences, Arts, and Letters.

An interrogation point was also placed after Acalypha vir-

ginica where given as a host of Peronospora euphorbiae Feld, in

the provisional list because the quantity collected was insufficient

for determination. Further search has been fruitless save for

one conidiophore.

Plasmopara ribicola Schroet. The young oospores of this

species that I have seen are few, scattered, globular, smooth,

26-33 [M in diameter. The oogonial wall is often symmetrically

thickened on two opposite sides.

Protomyces fuscus Pk. appears to be a race of Plasrm-

para pygmaea (Ung.) Schroet. that produces oospores abund¬

antly but conidia not at all. I have had this form under obser¬

vation for a number of years with reference to the appearance

of conidia. The similarity of the spores to the oospores of Plas¬

mopara pygmaea (Ung.) Schroet. and the presence of antheridia

indicate the character of the fungus. I label it Plasmopara

pygmaea var. fusca (Pk.) although it loses the character of the

genus with the suppression of conidia.

Protomyces andinus Pat. In 1911 this was collected at Butter¬

nut and Madison on Bidens with but few scattered resting spores

and no hypertrophy of the host. Examination of fixed material

of this kind showed the nuclei to be degenerating. I have not

seen it on Bidens since although abundant on Ambrosia.

Sept or ia alnifoUa Ell. & Evht. A hypophyllus Septoria on

Alnus incana collected at Madison is probably of this species but

doubtfully distinct from S. alni Sacc.

Septoria dentariae Pk. It is probable that this may properly

be referred to S. sisymbrii Ellis as was done in the preliminary

list. There is no question, however, as to its being the fungus

that was later described by Peck under this name. Hennings

& Ranojevic have proposed the name Septoria sisymbrii for a

Servian fungus (Ann. Mycol. 10 :390 (1910) which is perhaps

not distinct from the American plant, but I have not seen Kab.

& Bub. Fungi Imp. Exs. 557.

Gloeosporium saccharinum Ell. & Evht. ( Proc. Acad. Nat. Sci.

Phila. 1891, pp. 82-83) appears to have been founded upon ma¬

terial of an unusual character in which the fungus had run

Davis — Parasitic Fungi in Wisconsin — II.

95

riot. The form ordinarily seen shows spots of subcircular out¬

line 5-15 mm. in diameter. They are at first pale-olivaceous or

reddish brown, become alutaceous, fading with age,' the peri¬

pheral portion darker. They are sometimes confluent. The

tissue of the spots finally disintegrates, becomes ragged and falls

away centrifugally. The acervuli appear centrifugally, are

epiphyllous, saucer shaped, 80-1 60/x in diameter, sometimes con¬

fluent.

Juniper us communis depressa was given as a host of Ger co¬

rpora sequoiae juniperi Ell. & Evht. in the 4th supplementary

list but was omitted from the provisional list. Additional speci¬

mens have been collected at Lake Mills by E. M. Gilbert and the

writer.

Puccinia cirsii-lanceolati Schroet. I find in the herbarium

aecia of this rust collected at Blue Mounds.

Puccinia rubigo-vera (D C.) Wint. Secale cereale seems to

have been omitted from the list of hosts of this rust in the provi¬

sional list.

Gy mno sporangium clavariaeforme (Jacq.) D C. Telia have

been collected on J uniperus communis depressa at Merrimack and

Sullivan. They were abundant at the latter station.

Phragmidium occidentale Arth. In the provisional list aecia

and telia were reported. Uredinia have since been collected at

Ellison Bay in the northeastern corner of the state. Previous

collections were made in the northwestern portion.

Melampsoropsis ledicola (Pk.) Arth. Telia of this species

were collected at the same time as those of the following. The

uredinia have not yet been collected in Wisconsin. The record

“I” in the provisional list was founded on a single collection of

Peridermium decolorans Pk. in Yilas county.

Melampsoropsis ledi (Lk.) Arth. Germinating telia were

collected at Sturgeon Bay June 24th, 1913.

Melampsoropsis chiogenis (Diet.) Arth. In N. A . Flora the

type locality of this rust is given as “ Forest City ’ ’ which should

read Forest county. The station is now included in Oneida

county.

96 Wisconsin Academy of Sciences , Arts , and Letters.

Pucciniastrum myrtilli (Schum.) Arth. Specimens bearing

telia were collected at Athelstane. The teliospores were more

abundant in the epidermal cells of the upper surface of the

leaves. Fraser ( Mycologia 5:237, 6:27) finds that the aecia are

borne on the leaves of Tsuga canandensis. The Peridermium

peckii of the provisional list probably belongs to this species.

The Aecidium sp. indet. on Ampliicarpa monoica of the pro¬

visional list is Aecidium falcatae Arth.

Senecio aureus was unintentionally omitted from the enumera¬

tion of hosts of “Aecidium compositarum” in the provisional

list. Collections of aecia on this host have been made at Racine,

Radisson and Merrimack.

According to the inoculation experiments of Fraser ( Mycol¬

ogia 4 :236, 6 :25) the Peridermium balsameum Pk. of the provi¬

sional list is probably the aeeial stage of Uredinopsis. There

seems to be no way at present of determining with which of the

five described species of TJredinopsis that occur in Wisconsin any

particular specimen of the Peridermium is connected. Fraser

has found that Calyptospora goeppertiana Kuehn also produces

a Peridermium on Abies balsamea. This rust probably occurs

also in Wisconsin but has not yet been collected.

Caeoma abietis-canadensis Farl. has been shown by Fraser

. ( Mycologia 5:188, 5:238, 5:27) to be the aeeial form of a

Melampsora on Populus grandidentata which does not produce

aecia on Larix. Probably some, if not all, of the uredinia and

telia collected in Wisconsin on this host are of this race.

Entyloma lineatum (Cke.) Davis. Material that had been win¬

tered out of doors was brought to germination in tap water slide

cultures early in May. The normal germination appears to be in

the sorus, isolated spores seldom germinating. The promycelium

is consequently long (usually 35-50/x) and is flexuose and irreg¬

ularly nodulose, reminding one of the conidiophores of Rami -

laria. The sporidia are borne in apical whorls of 2 to 4, are

fusoid — cylindrical, 7-14x2 y. The whorl of sporidia is detached

intact together with about an equal length of the distal portion

of the promycelium and then rises to the surface of the water in

the currents of which it revolves and moves in a very irregular

manner. This is the same method of detachment that takes place

Davis — Parasitic Fungi in Wisconsin — II.

97

in Entyloma nymphaeae (Gunn.) Setch. (Trans. Wis. Acad. Sci.

Arts & Letters 77:176) and is probably correlated with the

aquatic habit. It seems to be of service in increasing the flotation

of the sporidia and the likelihood of their catching upon the

host. That this method of detachment is not constant, however,

is indicated by the fact that Setchell does not mention it in his

account of the germination of the spores of this species but re¬

fers to the sporidia as falling from the promycelia. (Bot. Gaz.

73:188 [1894].)

Additional Hosts.

Not previously recorded as bearing the fungi mentioned in

Wisconsin.

Peronospora parasitica (Pers.) Tul. — On Arabis hirsuta . Fish

Creek.

Synchytrium aureum Schroet. A few galls on Caltha palustris

apparently caused by this fungus were collected at Wausaukee

in August, 1913, but the material is scanty and immature.

Plasmopara pygmaea (Ung.) Schroet. Conidia and oospores

on Hepatica triloba. Afton.

Peronospora grisea Ung. On Veronica americana. Ellison

Bay.

Sphaerotheca nwrs-uva& (Schw.) B. &. C. on Ribes gracile.

Detroit Harbor.

MicrospJiaera alni (Wallr.) Wint. On Alnus incana. Wau¬

saukee. Perithecia sparse.

Dimer osporium collinsii (Schw.) Thuem. On Amelanchier,

oblongifolia. Merrimack, May 3rd, 1913. (W. N. Steil). On

leaf of preceding year but ascospores not yet formed.

Exoascus confusus Atk. On fruit of Prunus virginiana. Stur¬

geon Bay.

Exoascus insititiae Sadeb. On Prunus pennsylvanica. Stur¬

geon Bay, causing witches brooms.

Exoascus cerasi (Fckl.) Sacc. On Prunus Cerasus (cult.)

Wyalusing.

7— S. A.

98 Wisconsin Academy of Sciences , Arts , and Letters.

Exoascus coerulescens (Mont. & Desna.) Tnl. On Quercus

coccinea. Richland Center. (R. A. Harper & G. M. Reed.)

Taphrina virginica Seym. & Sadeb. On Ostrya virginiana.

Potosi.

I am indebted to Mr. H. G. MacMillan of the Wisconsin Ag¬

ricultural Experiment Station for identification of specimens in

this group.

Stagonospora intermixta (Cke.) Sacc. To this species I have

referred specimens of which the following notes were made. On

elongated light brown dead areas which soon spread over the

whole leaf; pycnidia epiphyllous, scattered, dark brown, glob¬

ose or depressed-globose, 60-100/* in diameter; sporules at first

hyaline and cylindrical becoming acute at one end with a cen¬

tral row of small guttulae, finally septate and tinted, 26-52x3-4/*.

On Phalaris arundinacea. Devils Lake, Wisconsin, August 5th,

1913. The pycnidial wall is usually thin at the base while the

outer portion is thick and blackened.

Sept or ia agrimoniae-eupatorii Bomm. & Rouss. On Agri-

monia gryposepala. Potosi and Glen Haven.

Septoria cacaliae Ell. & Kell. On Cacalia atriplicifolia. Lake

Mills. Oct. 26, 1901.

Septoria silphii Ell. & Evht. On Heliopsis scabra. Madison.

Sporules 26-36 x 1/*. This species was described as having

sporules 35-50 x ly, but in specimens on Silphium perfoliatum

collected at Madison they are but 26-36 x ly. The spots tend to

become white and arid with age.

Entomosporium maculatum Lev. var. cydoniae Sacc. On

Pyrus Aucuparia. Devils Lake. Sporules 20(16-23) x 8(6-10)/*.

Gloeosporium septorioides Sacc. On Quercus rubra. Devils

Lake. In these specimens the sporules have a narrow median di¬

vision of the cytoplasm which is sometimes apparent without

staining.

Gloeosporium robergei Desm. On Ostrya virginiana. Somers,

South Milwaukee and Devils Lake. In all the specimens which

I have collected on this host the cuticle on the upper surface of

the spots is rugose forming white dendritic lines. It is labeled

var. dendriticum in our herbaria.

Davis — Parasitic Fungi in Wisconsin — II.

99

Gloeosporium ribis (Lib.) Mont. & Desm. On Ribes Cynos-

bati. Devil’s Lake.

Marssonina martini (S. & E.) Magn. On Quercus Muhlen-

bergii. Bridgeport. Some of the leaves bear also larger paler

spots usque 2 cm. in diameter with numerous acervuli resembling

those of Gloeosporium nervisequum (Pckl.) Sacc. (Gloeospor¬

ium canadense E. & E.) but the sporules are uniseptate.

Cylindrosporium glyceriae Ell. & Evht. On Glyceria cana¬

densis. Athelstane. The collection on this host bears longer

sporules, an occasional one much longer, than the type.

Cylindrosporium betulae Davis. On Betula alba papyrifera.

Wausaukee. Sporules usque 55 x 3/a. Microeonidia are quite

common in this species seeming to be produced especially when

the stroma is erumpent and naked.

Microstroma juglandis (Bereng.) Sacc. On Cary a glabra .

Potosi.

Septocylindrium ranunculi Pk. On Ranunculus septentrio -

nalis. Madison.

Ramularia occidentalis Ell. & Kell. On Rumex Britannica.

Madison and Athelstane.

Ramularia pratensis Sacc. On Rumex Britannica. Athel¬

stane.

Ramularia desmodii Cke. On Desmodium illinoense. Bridge¬

port.

Ramularia effusa Pk. On Vaccinium pennsylvanicum. Wau¬

saukee. On these specimens the conidia are borne on definite

orbicular spots about 5 mm. in diameter which also bear numer¬

ous black immature pycnidia.

Ramularia veronicae Fckl. On Veronica serpyllifolia. Madi¬

son. This specimen bears conidia 12-18 x 3/a with a median di¬

vision of the cytoplasm.

Ramularia asteris (Sacc.) Barth. On Aster lateriflorus.

Devils Lake.

Fusicladium radiosum (Lib.) Lind. On Populus balsamifera.

Sturgeon Bay. But a single collection has been made on this

host.

100 Wisconsin Academy of Sciences, Arts , and Letters .

Cercospora caricina Ell. & Dearn. On Car ex castanea and

C. intumescens. Wausaukee. On Car ex retrorsa. Detroit Harbor

and Athelstane. In the specimen from] the latter locality the

conidiophores are only 15-25/a long.

Cercospora rhoina Cke. & E1L On Rhus glabra . Madison

and Bridgeport.

Vstilago utriculosa (Nees) Tul. On Polygonum lapathifo -

Hum. Madison (W. N. Steil).

TJromyces halstedii de Toni. On Leersia oryzoides. Wiscon¬

sin river bottom opposite Bridgeport. The only previous col¬

lection of this rust in Wisconsin was a scanty one made by Dr.

Arthur at the dells of the Wisconsin river in 1893 on Leersia

virginica.

TJromyces scirpi Burr. Scirpus validus is given as a host of

this rust in Wisconsin in North American Flora.

TJromyces junci-tenuis Syd. On J uncus Dudleyi. Uredinia

and telia at Wausaukee.

TJromyces proeminens (D C.) Lev. Telia on Euphorbia glyp-

tosperma. Wausaukee; This name is used instead of the later

TJromyces euphorbiae Cke. & Pk. in the wide sense in which that

name was used in the provisional list although it is also the name

which is applied, in the narrow sense, to the particular race

which occurs on this host species.

TJromyces hyperici-f rondo si (Schw.) Arth. Uredinia and a

few telia on Hypericum Kalmianum. Fish Creek.

Puccinia coronata Cda. Aecia on Rhamnus lanceolata. Glen

Haven.

Puccinia andropogonis Schw. An Aecidium on Linaria cana¬

densis in the herbarium of the University of Wisconsin which

was collected at Mazomanie in June, 1908, is perhaps of this

species.

Puccinia cyperi Arth. Uredinia and telia on Cyperus Hough-

tonii. Athelstane.

Puccinia patruelis Arth. Telia on Carex pennsylvanica.

Madison (E. T. Bartholomew) North American TJredinales, 651 .

Davis — Parasitic Fungi in Wisconsin — II.

101

Puccinia curtipes Howe. On Tiarella cordifolia. Devils

Lake.

Phragmidium disciflorum (Tode) Janies. Uredinia and telia

on Rosa acicularis at Ellison Bay are probably Ph. rosae-acicu-

laris Liro if one accepts that as a distinct species.

Melampsoropsis cassandrae (Pk. & Cl.) Arth. Aecia ( Peri -

dermium consimile Arth. & Kern) on Picea canadensis . Wau-

sankee.

Uredinopsis atkinsonii Magn. A collection on Cystopteris

bulbifera from the Wisconsin river bluff opposite Bridgeport I

have referred to this species.

Additional Species.

Not previously reported as occurring in Wisconsin.

Plasmopara viburni Pk. On Viburnum Opulus americanum.

Wausaukee and Athelstane. Conidia and . oospores. Oospores

subepidermal, oogonia globose, usque 48/a in diameter, wall often

irregularly thickened ; oospores globose, 24-37/a in diameter, wall

2-4/a thick, smooth or nearly so.

Asterina cupressina Cke. On Juniperus commums depressa.

Fish Creek.

Ascochyta Wisconsin a n. sp. ad interim. Spots orbicular to

elliptical, gray with a narrow black border and frequently zonate

above, brown with a less definite margin below, 1-3 cm. long;

pycnidia epiphyllous, scattered, brown, prominent, globose to

sublenticular, 85-110/a in diameter ; sporules ovoid to oblong, hy¬

aline, 4-8 x 2%-31/^/a. Some of the longer sporules have a me¬

dian septum and it is probable that well matured specimens

would show this to be an Ascochyta. The affected leaf tissue

seems quite friable and apparently fragments and falls away

piecemeal. On Sambucus canadensis. Devils Lake. August

7, 1913. On examining a specimen of Septoria sambucina Pk.

collected at Racine it was found to bear also an Ascochyta with

102 Wisconsin Academy of Sciences , Arts, and Letters .

sporules 8-10 x 2^-3/x. The pyenidia are epiphyllous on spots

1-2 cm. long which are sordid-arid with a purple border above,

olivaceous below. I use this name for convenience until the re¬

lationship of the fungus to the various species that have been

described on Caprifoliaceae is known.

Ascochyta caulicola Laubert ( Ascochyta lethalis Ell. & Barth.)

On living stems of Melilotus alba. Madison (A. H. Gilbert).

Stagonospora paludosa (Sacc. & Speg.) Sacc. On Carex re -

trorsa . Athelstane.

Septoria betulicola Pk. The common Septoria on Betula in

Wisconsin, first manifests itself by the formation of small

(l-2mm.) scattered, angular, intervenular, black brown spots

which are lighter colored below. These spots become surrounded

by an indefinite yellow discoloration which later becomes of a

more or less reddish brown above and light brown or buff below.

These run together into indefinite areas usually 1-2 cm. in di¬

ameter. On the lower surface of these areas the usually few and

scattered pyenidia are borne. These are subepidermal, globose,

thick-walled, about 100/* in diameter. The sporules are straight

to strongly curved, spuriously pluriseptate, 40-60x11/2-2/*.

This is the form that is usually distributed under the name Sep¬

toria betulicola Pk. although North American Fungi 2nd series,

2166 which resembles it was issued as Septoria betulae (Lib.).

Other specimens show smaller (ca. 5mm.) darker, more defi¬

nitely limited areas which become cinereous above. Septoria

betulicola apparently has not been reported in any of the Wis¬

consin lists. The characters of the sporules seem to ally this

with Septoria betulina Pass. Septoria betulae (Lib.) West,

was reported in the supplementary list 402a. The specimen

upon which this record was based (Three Lakes, June 25th,

1892) bears circular light yellowish brown spots 1-2 mm. in di¬

ameter with a definite dark brown border. The pyenidia are

epiphyllous but visible below, globose, thick-walled, about 80^ in

diameter; the sporules straight or curved, triseptate, 30-40x2/*

Fungi Columbiani 1586 on Betula Occident alis, collected in Ore¬

gon and issued as Septoria betulicola Pk. seems to differ from

this only in the more irregular spots and the much paler and less

distinct border.

Davis — Parasitic Fungi in Wisconsin — 11.

103

Septoria alni Sacc. On Alnus incana. Wausaukee. Re¬

ferred to this species because of the short sporules, the longest of

which attain 40//..

Septoria hepaticae Desm. On Hepatica acutiloba. Glen Hav¬

en. I have collected this but once when it occurred in connection

with Protomyces fuscus Pk.

Septoria cassiaecola Kell. & Swingle. On foliage leaves of

Cassia chamaecrista. Glen Haven. Amphigenous on interven-

ular areas of the leaflets which are not at first discolored but

which become brown.

Septoria senecionis-aurei n. sp. ad interim. On irregular

indefinite grey portions of large brown areas of the radical leaves.

Pycnidia epiphyllous, scattered, brown black, spherical, with a

distinct cellular wall, 55-65//, in diameter; sporules hyaline,

straight, 16-26 x 1/x. On Senecio aureus. Devils Lake, Septem¬

ber 1, 1913. I use this name for convenience until the relation

of the fungus to previously described forms such as Septoria

senecionis-sylvatici Syd. and S. adenocauli E. & E. becomes

known.

Gloeosporium cylindrospermum (Bon.) Sacc. On Alnus in¬

cana. Madison.

Marssonina neilliae (Hark.) Magn. On Physocarpus opulifol-

ius. Wausaukee. Macroscopically this resembles Entomospor-

ium. Marssonina coronaria (Ell. & Davis) is similar and doubt¬

less closely related.

Marssonina baptisiae (E. & E.) On Baptisia leucantha.

Bridgeport. The globose acervuli are not subcuticular, as one

might infer from the description, but subepidermal or innate.

Sporules as long as 33//. were observed. Septation of the spor¬

ules seems doubtful.

Marssonina rhabdospora (E. & E.) Magn. A collection on

leaves of Populus grandidentata made on the bank of the Wis¬

consin river opposite Bridgeport September 18, 1913, bears

hypophyllous subcuticular acervuli which are applanate to

hemispherical, 75-150// in diameter. The affected leaf areas

are first yellow, then brown, then indefinite sordid spots ap¬

pear which become determinate, orbicular, arid, zonate, 3-5 mm.

104 Wisconsin Academy of Sciences, Arts, and Letters.

in diameter with a narrow dark margin. In these spots the leaf

parenchyma separates from the venules and probably falls away

leaving the venular network. The zonate spots look much like

the work of leaf miners, the dark line# suggesting burrows con¬

taining excreta. From the closely massed, erect, straight hyphae

of the acervuli are abstricted hyaline filiform sporules 7-11 x 2 //.

Fungi Columbiani 1587 (on Populus tremuloides, New field, N.

J. J. B. Ellis.) issued under the name Septoria musiva Pk.

bears similar but somewhat larger spots (1-2 cm.) and sporules

18-30x2-3//, uniseptate. I am considering the Wisconsin collection

to be a microconidial state of this which I refer to Marssonina

rhabdospora E. & E.

Since this was written collections on Populus grandidentata

have been made at Phlox and Neopit. The following notes were

made from the latter: Spots circular, alutaceous shading out¬

ward into reddish brown and with a darker margin, the upper

surface darker than the lower, 1-4 mm. in diameter, sometimes

confluent; acervuli hypophyllous, usually few; sporules gener¬

ally straight, uniseptate, 18-33 x 2-3 //. In the Phlox collection

the spots are more numerous, rather more angular and more

frequently confluent.

Cylindrosporium vermiforme n. sp. Spots amphigenous, sub-

circular to irregular, immarginate, brown, 5-15 mm. in diam¬

eter ; acervuli epiphyllous, scattered, subcuticular, flat, 40-60 // in

diameter; sporules, hyaline, vermiform, curved, sigmoid or

flexuose, pluriseptate, 150-250x4-5//. On leaves of Alnus in-

cana Devils Lake, Wisconsin. The sporules suggest eel worms

in appearance. Specimens in the herbarium of the Uni¬

versity of Wisconsin collected at Devils Lake, August 15th,

1906, by R. A. Harper show many of the sporules pro¬

vided with a rostrum, 10-42x1-1%//, at the apex. A collec¬

tion made in August, 1913, does not show the “ rostrum’ 9

but one made September 1, 1913, showed some of the spor¬

ules so provided. Living sporules, germinating in water, in

addition to the lateral germ tubes, put forth one from the apex

so like the ‘ 1 rostrum’ ’ that I infer that the beak is a germ

tube put forth while the sporule is still in situ. Some spor¬

ules bearing a beak put forth in water a second tube alongside

the rostrum and similar to it. Because of the large, erumpent,

fasciculate sporules this might be referred to Hyphales.

Davis — Parasitic Fungi in Wisconsin — 11. 105

Ascochyta saniculae n. sp. On indefinite, discolored, more

or less mottled areas which may include the entire leaf ; pycnidia

scattered, innate, globose to lenticular, thin walled, light reddish

brown with a round apical pore surrounded by a dark ring,

100-170/* in diameter; sporules hyaline, cylindrical, usually

straight, 4-guttulate, 20-30 x 4-6/*. On leaves of Sanicula

marilandica. Grant County, Wisconsin, September 19th, 1913.

The pycnidia are very inconspicuous. They are most readily

seen by transmitted light when they show as translucent points.

Cylindrosporium skepherdiae Sacc. (Ann. Mycol. 11 :551.

1913). To this species, founded upon material collected at

Field, B. C. by Dearness, I am referring specimens from which

the following notes were made. The spots are circular, reddish

brown, concentrically rugose, 3-5 mm. in diameter. They have

a greenish border and are seated upon an indefinite yellowish

area. The epiphyllous pycnidia ( ?) are aggregated in the cen¬

tral portion of the spot, are soon erumpent, and widely open,

the white mass of sporules within being visible under a hand

lens. The sporules are hyaline, oblong-cylindrical, obtuse, pluri-

septate, 18-40 x 3-4/*. Collected on ShepJierdia canadensis at

two stations near Ellison Bay and at Detroit Harbor. I have not

seen Septoria argyraeae Sacc. which I suspect is not very differ¬

ent.

Ramularxa fraxinea n. sp. Spots none, the fungus appearing

as small snow white patches on the lower surface of the leaves ;

conidiophores densely clustered on a more or less hemispherical

stroma, hyaline, sometimes bulbous at base, 10-20 x 3-4/* ; coni-

dia apical, hyaline, cylindrical, obtuse at both ends, distal %-

% more or less strongly curved, becoming 1-4 septate, 40-80 x 4

-5/*. On languishing leaves of Fraxinus pennsylvanica. Bridge¬

port, Wisconsin, September 17, 1913. The gross appearance sug¬

gests a light development of Microstroma and the shape of the

conidia a hockey stick. Perhaps this should be referred to

Fusarium. Since collected on Fraxinus pennsylvanica lanceo-

lata at Maiden Rock.

Cercosporella nivea EU. & Barth. On Solidago undigulata.

Athelstane. The specimen referred here bears conidia usque

118 x 3-4/* on short conidiophores. The affected leaf areas are

106 Wisconsin Academy of Sciences , Arts , and Letters.

not at first discolored but later become dead and brown and the

air spaces contain hyaline mycelium and immature pycnidia or

perithecia.

Cercospora echinochloae, n. sp. Spots elongate-linear red¬

dish brown, becoming arid in the center ; conidiophores hypophyl-

lous in small tufts, brownish tinted, continuous, straight or bent,

entire or denticulate or oblique at the apex, 10-26 x 4/a ; conidia,

hyaline, straight or curved, cylindrical to obclavate-cylindrical,

distinctly 1-7 septate, 23-53 x 3-4/a. On leaves of Eckinockloa

Crus-galli. Devils Lake, Wisconsin, August and September

1, 1913.

Cercospora passaloroides Wint. Poor specimens of this fungus

on leaves of Amorpha fruticosa were collected on the bank of the

Wisconsin river opposite Bridgeport. Better specimens have

been collected at Trempealeau.

Pusarium carpineum n. sp. Hypophyllous on indefinite areas

which often follow the principal veins and which come to have

a wilted appearance and finally assume a lethal brown below and

almost black above ; sporodochia superficial, convex to subhemis-

pherical, 25^0 /a in diameter, composed of globose cells 7-8^ in

diameter which become flask shaped and 12-15/a long; conidia

borne singly on the apex of these conidiophores, hyaline, cylin¬

drical, obtuse at both ends, curved, biseptate, 35-50 x 3-4/a. On

Carpinus caroliniana, Wyalusing, June 12, 1913.

TJromyces graminicola Burr. Telia on Panicum virgatum.

Madison. On the railroad right of way. Perhaps adventive.

Puccinia melicae (Erikss.) Syd. Uredinia on Melica striata.

Vilas county. Treboux states as the result of inoculation experi¬

ment that this is one of the forms of crown rust (Ann. My col,

72:5:483 [1914]).

Puccinia gigantispora Bubak. Aecia and telia on Anemone

cylindrica or virginiana. Glen Haven.

Puccinia cickorii (DC.) Bell. Uredinia and telia on Cickorium

Intybus. Madison, (E. T. Bartholomew) Fungi Columbiana

3933.

Davis — Parasitic Fungi in Wisconsin — II. 107

Gy mno sporangium corniculans Kern. Galls on Juniperus

horizontalis on which were some dried teliospores agreeing with

those of this species were collected on the lake Michigan beach

east of Sturgeon Bay in June.

Hyalopsora aspidiotus Pk. Uredinia on Phegopteris Dryop-

teris. Detroit Harbor.

Melampsora farlowii (Arth.) n. comb. (Necium farlowii Arth.)

On young twigs and leaves of Tsuga canadensis. Detroit Har¬

bor.

Melampsora arctica Rostr. Aecia on leaves of Abies balsamea

supposed to be of this species have been collected at Brule (E.

M. Gilbert) Fish Creek and Ellison Bay. The reference to this

species is based on the inoculation experiments of Fraser (Myco-

logia 4:187, 5:238). The uredina and telia have not yet been

recognized in Wisconsin.

Aecidium xanthoxyli Pk. On Zanthoxylum americanum.

Mazomanie (R. A. Harper & G. M. Reed) Glen Haven.

Aecidium proserpinacae B. & C. On Proserpinaca palustris.

Detroit Harbor. Abundant on the marshy border of a pond on

Washington Island.

108 Wisconsin Academy of Sciences, Arts , and Letters.

INDEX TO HOSTS MENTIONED IN “NOTES” I AND II.

Davis — Parasitic Fungi in Wisconsin — II.

109

Prunus Cerasus . .

Prunus pennsylranica .

Prunus virginiana . .

Pyrus Aucuparia .

Quercus coccinia .

Quercus macrocarpa ..... - -

Quercus Muhlenbergii .

Quercus rubra .

Ranunculus recurvatus .

Ranunculus septentrionalis . . .

Rhamnus lanceolata .

Rhus glabra .

Ribes amerlcanum .

Ribes Cynosbati .

Ribes gracile . .

Rosa acicularis . . . .

Rubus hispidus .

Rubus villosus .

Rumex Britannica . . . . .

Sambucus canadensis .

Sanicula marilandica .

Pago

Scirpus validus . . . 100

Secale cereale . 95

Senecio aureus . 96, 103

Shepherdia canadensis . 105

Silphium perfoliatum . 98

Solidago uniligulata . . 105

Sonchus asper . 92

Thalictrum dasycarpum . 92

Thalictrum dioicum . 92

Tiarella cordifolia . 101

Tsuga canadensis . . . 96, 107

TTvulari grandiflora . 87

Vacinium pennsylvanicum . 99

Veronica americana . . 97

Veronica serpyllifolia . 99

Viburnum Opulus . 101

Vicia . 93

Vicia americana . . 98

Vicia linearis . 93

Viola pubescens . 85

Zanthoxylum americanum . 107

Page

97

79, 97

98

87

99

98

85

99

100

90

99

97

101

85

99

101

105

THE RELATION OF THE CORPUS CHRISTI PROCESSION

TO THE CORPUS CHRISTI PLAY IN ENGLAND

Merle Pierson

Introduction

One of the theories concerning the rise of the vernacular drama

in England traces the development of the Corpus Christi play

from the Corpus Christi procession. The supposed evolution

of the one from the other has been described by Mr. Davidson

(Studies in the English Mystery Plays 9 p. 93) as follows:

It seems, then, that shortly after the confirmation of Corpus Christi

in 1318, pageants of the Biblical story were introduced in conjunction

with the banners of the crafts. These at first were mute mysteries

expressed by action. Indeed, connected pantomimic action would seem

impossible in a moving procession; therefore this custom may be older

than the spoken drama. In a short time, however, spoken drama,

. . . became an established custom in England. A spoken drama

necessitated frequent halts by the procession, as it was impossible to

act satisfactorily in motion. These halts prolonged the procession be¬

yond reasonable limit, and were avoided by transferring the pageants

to the rear of the procession. A division of the procession immediately

arose through the slower movement of the pageants, but the plays,

though much belated, followed the traditional course of the procession

through the city.

On the basis of this statement five stages in the development

of the Corpus Christi pageant, from the Corpus Christi proces¬

sion might be assumed:

I. Crafts merely marching in the procession.

II. Crafts with banners in the procession.

III. Mute Mysteries.

YL Spoken drama in the procession.

V. Separation of the plays from the procession.

Pierson — The Corpus Christi Procession.

Ill

Ob the authority of Mr. Spencer (Corpus Christi Pageants in

England, p. 81) one more stage might be added:

VI. Pageant wagons and actors in the procession after the

separation from the plays.

The evidence given by Mr. Davidson and by Mr. Spencer for

each of these stages is, however, too meagre to be conclusive. It

was in the hope of discovering additional material that I under¬

took this study.

To be definitive, the illustrations must show us the process as it

worked out in some one place. It will not do to say that since

in Durham, the crafts with banners marched in the Corpus

Christi procession, since in Prance, the crafts acted mute mys¬

teries, and since in Beverley the procession and the plays were

separated, therefore the development followed the consecutive

stages I, II, III, IV, V, VI. If, however, one should find that

in Coventry every stage is represented in the correct order, he

might assume that the theory was substantiated, at least for that

one case. For each town, therefore, I have grouped together in

a chronological order all the material on the Corpus Christi play

and the Corpus Christi procession. Except in unimportant en¬

tries, the original language of the illustrations is given. In the

first column to the left will be found the date to which the evi¬

dence refers; in the second narrow column, the authority (full

titles are given in the bibliography). In the wide column at

the left, is the evidence itself; in the wide right-hand column,

a comment on the stage represented.

Aberdeen

DOCUMENTS.

Content

“The play is men¬

tioned in a regulation of

1440.”

“The samyn dai, the

consale and brethryn of

gilde beand present for

the tym, has consentit

and ordanit the aider-

man to mak the expen-

sis and costis — of the

play to be plait in the

fest of Corpus Xristi

nixttoeum.”

Interpretation

“It seems that on Cor •

pus Christi day after

the procession a play

was usually performed

on Windmill-hill. The

play is mentioned in a

regulation of 1440 and

again in 1479, but it

probably changed from

year to year, and was

under the care of the

Abbot of Bonaccord. It

certainly was not the

charge of the gilds.”

Davidson, p. 97. Stage I.

112 Wisconsin Academy of Sciences, Arts, and Letters .

Aberdeen

Date Document

1512. Register, p. 442.

N

1530. Register, p. 449.

1531. Register, p. 450.

1533. Chambers, II, 332

1538. Council Register,

p. 452.

1546. Chambers, II, 332

1553. Register, p. 456.

DOCUMENTS— Continued.

Content

“Every Craft — s all

have a pair of torcheiss,

— to decoir and worschip

the sacrament one Cor¬

pus Xti day.”

“The craftismen of

this burgh sail, in thair

best arraye, keep and

decoir the processionls

on XX i [Corpus Christi]

day and Candelmas day

— every craft with thair

avin banar, with the

aimes of their craft

thairin, with thair peg-

ane. — And every craft

in the said processionis

pall furneiss thair pe-

gane and banar honest-

lie as effers, conforme

to the auld statut maid

in the yeir of Godjajvc

tene yers.”

“The Craftismen

— keipe and decoir the

procession on Corpus

Christi dais, and Candil-

mes day — Every craft

with thair Awin banar,

with the armes of thair

craft therein. — E very

ane of the said craftis,

in the Candilmes pro-

cessioun, shall furneiss

thair pageane, conforme

to the auld statut, maid

in the yeir of God jai

vc and X yeris.”

“‘The craftismen —

sail — keip and decoir,

the processionis on XXi

day and Candelmes day

— every craft with thair

avin banar — with their

pegane.’ ”

Hammermen complain

that “ameraris” usurp

their place in Corpus

Christi procession.

“Litsters ordered to

‘haue thar banar and

Pagane, as uther craftis

of the said Burgh hes,

ilk yeir, on Corpus Xhri

day, and Candilmess

dayis processiounis.’ ”

Smiths convicted of

“refusing Contempur-

indlie to gang in ordour

in the processioun of

Corpus Xris day.”

Interpretation

‘Pegane’ probably re¬

fers to the pageant

wagon for in the regu¬

lations for the Candle¬

mas procession (1505/6)

is this phrase “pa¬

geant that they fur-

nyss to keip their geir.”

Chambers, II, 331.

Stage VI, Stage II.

The furnishing of the

pageants probably re¬

fers only to the Candel¬

mas procession.

Since by 1531, the Cor¬

pus Christi plays had

been established, the

carrying of the banners

represents a survival

from stage II, rather

than stage II itself.

Stage I.

Stage VI, Stage II.

Stage I.

Pierson — The Corpus Christi Procession.

113

A berdeen

DOCUMENTS— Continued.

Date

Document

1554. Register, p. 457.

See also Cham¬

bers, II, 332.

1556. Chambers, II, 333

Davidson, note, p. 95.

Content

“Williame Robertsone,

dekin of the smythis,

comperit in judgement,

— thai war in vse of

ganging be thame self-

fis in the said proces-

sione, under thair awin

baner, hindmaist and

nixt the Sacrament, and

the said wrychtis ma-

souns, cowperis, and

sklaiteris to proceid to-

gidder befoir thame,

under ane baner and

pegane, separat fra the

said smythis.”

“Order for observance

of statute as to Corpus

Christi procession.”

Interpretation

Stage YI, Stage II.

“On Corpus Christi

day the procession was

under direction of the

Abbot of Bon-Accord,

later under that of

T3 y— v i n TJrtArl

Conclusion: Since by the first reference in 1440 spoken

drama had already been established in Aberdeen, the succeeding

references to the carrying of banners and the drawing of pag¬

eant wagons in the procession indicate merely a possible earlier

connection between the procession and the plays. Certainly

the gilds (contrary to Mr. Davidson’s; statement, p. 97) were

concerned with both plays and procession (see entry 1479.) Be¬

cause the pageant wagons were drawn in the procession, it is

not necessary that the plays should have originated in it. The

plays may have developed separately and have become at¬

tached to the Corpus Christi festival. The wagons might then

have been sent in the procession to add to its splendor or to an¬

nounce the plays. Obviously, material earlier than 1440, show¬

ing that the procession was flourishing before the first spoken

drama, must be found to prove the validity of the theory.

8— -S. A.

114 Wisconsin Academy of Sciences, Arts, and Letters .

Beverley.

DOCUMENTS.

Date Document

1377. Hist. Mss., p. 65.

Leach (2), p. 209.

Chambers, II,

339. Bev. Town

Doc., p. 45.

1390. Hist. Mss., p. 66.

Chambers, II,

339. Leach (2),

p.208. Bev. Town

Doc., p. 33.

1391. Hist. Mss., p. 66.

Leach (2), p. 209.

Chambers, II,

339.

1392. Hist. Mss., p. 66.

Bev. Town Doc.,

p. 36.

2. Leach (2), 209.

1409. Hist. Mss., p. 89.

Bev. Town Doc.,

p. 40.

Content

“Also, A. D. 1377, they

agreed in the Guild hall

that all tailors of Bev¬

erly should be present

at the account of the

expenses incurred by

the pageant of the play

of Corpus Christi.”

38 specified crafts to

“have their plays and

pageants (‘pagentes’)

prepared hereafter

every day in the Feast

of Corpus Christi, in

manner and form ac¬

cording to the ancient

custom of the town of

Beverley.”

John of Erghes, hay-

rer, before 12 Keepers

of town “undertook for

himself and his fellows

of the same craft to

play a certain play

called Paradise suffi¬

ciently, viz., every year

on the feast of Corpus

Christi when other

craftsmen of the same

town play, during the

life of the said John of

Erghes, at his own

cost.”

1. Smiths pay penalty

for not playing their

play of Ascension on

Corpus Christi day.

2. Play under penalty.

“And the said gold¬

smiths shall yearly

maintain — a torch in

the procession on Cor¬

pus Christi day forever.”

Interpretation

Stage V.

Stage I.

Stage V.

Crafts for play in¬

clude trades not joined

before 1416. (note in

Hist. Mss., p. 99.)

1410-11. Hist. Mss., p. 97.

1411 ( ?). Hist. Mss., p.

99.

“Every master of the

craft of coupers who

shall newly occupy shall

pay on his entrance to

the use of the craft and

costs of Corpus Christi

play 2s.

Ordinances of Bowers

and Fletchers under

date of 1411 provide for

play of “Fleyhg into Egip

and for play of Habra-

ham and Isaak.”

Pierson — The Corpus Christi Procession.

115

Beverley.

Date Document

1411. Hist. Mss., p. 67.

Chambers, II,

341. Leach (2),

211.

Bev. Town Doc., p.

34.

1413. Hist. Mss., p. 98.

1414. 1. Chambers, II,

339.

2. Leach (2),

p. 211.

1416. Hist. Mss., p. 101.

1420. Leach (2), p. 208.

1423. Hist. Mss., p. 160.

Leach (2), p. 216.

1423. Hist. Mss., p. 160.

Leach (2), p. 214.

1423. Hist. Mss., p. 160.

Chambers, II, 339.

DOCUMENTS— Continued.

Content

Those of the worthier

sort, though they had

not done so before,

should on Corpus Christi

day erect a pageant,

and support it at their

own cost, and cause a

play to be played hon¬

ourably and fittingly.”

“The Bowyers and

Fletchers to “play or

cause to be played, a

certain pageant on Cor¬

pus Christi Day of Ab¬

raham and Isaac,’ when

the community on St.

Mark’s Day consent that

the pageants generally

shall be played.

1. Barbers’ ordinances

require members to pay

certain sums toward

play.

2. According to ordin¬

ances of barbers, codi¬

fied or written down in

1414, their play was the

Baptism of Christ by St.

John.

The tanners to “sus

teyn and uphold forever

— two torches of wax

yerly and forever to be

born in procession in

the Feast of Corporis

Christi.”

According to ordin-

a n c e s of Carpenters’

Gild the Resurrection

was their play.

Fines for neglecting

pageants on Corpus

Christi day.

“Expenses of the

Twelve Keepers on Cor¬

pus Christi day in gov¬

erning all the pageants

passing through the

whole town.”

“To Master Thomas

Bynham, Frier Preacher,

for making! and com¬

posing the banns (Tes

banes’) before the Cor¬

pus Christi play pro¬

claimed through the

whole town, 4 May, 6S.

8d. To the waits of the

town, on the morrow of

Ascension Day, riding

with the said proclama¬

tion of Corpus Christi

through the whole

town, 20d.”

Interpretation

Stage V.

The procession seems

to have been a settled

yearly thing; it was

necessary to vote on St.

Mark’s Day to have the

plays.

Stage V.

Stage I.

Stage V.

116 Wisconsin Academy of Sciences , Arts , and Letters.

Pierson — The Corpus Christi Procession .

117

Beverley.

DOCUMENTS— Continued.

Date Document

1441. Hist. Mss., p. 100.

1442. Hist. Mss., p. 129.

1446. Hist. Mss., p. 131,

132.

1449. Hist. Mss., 133.

1449. Leach (2) p. 214.

Chambers, II, 341.

1460. Hist. Mss., 134.

Leach (2), p. 216.

Content

Sadlers to play the

pageant of the Creation

of the World yearly on

Corpus Christi day,

whenever the 12 keepers

of the town shall order

plays.

“Rob. Peterer et Th.

Cowton moniti sunt ad

exponendum in gratiam

communitatis xls; pro

eo quod ipsi non habue-

runt lumen suum artis

Piscatorum in proces-

sione die Corpris Christi

per festum ad Vincula

Sancti Petri prox.”

“ ‘Uxor Egidii Bokeler,

smyth’, promised to pay

‘solutio xld. annuatim

Pistoribus quamdiu oc-

cupaverit, xxd. ad ex¬

pen sas castelli Pistorum

et luminis, et ad ex-

pensas et costagia pag-

endae cum ludi contig-

erit xxd.’ ”

“Ordinatum est per

decern de xij — quod pag-

endae ludi Corporis

Christi assignentur in

forma subscripta delud-

endae hoc anno: viz., ad

Barras Borialis; juxta

Bulryng; inter Johan-

nem Skipwith et Rober-

tum Couke in Alta Via;

apud Crossebrig; apud

Fisch-m a r k e t: apud

Mynstir bowe; et ad

Torrentum.

Expenses of Govern¬

ors, common clerk, and

sergeant governing the

pageants of the town on

Corpus Christi through

the whole day. “The

play lasted only one day,

and was given in 1449 at

six stations.” Cham¬

bers, II, 341.

Certain fishers “moniti

sunt hie ad exponendum

[etc.] xls. quia non ha-

buerunt pagendam suam

deludendam die Corporis

Christi hoc anno secun¬

dum consuetudinem vil-

lae — et transgr e s s i o

pardonatur graciose sub

hac condicione, quod

praedicti Piscatores pa¬

gendam suam facient

competenter in omnibus

citra diem Dominicam in

Ramis Palmarum proxi-

mum futuram post da¬

tum praesentium.” |

Interpretation

Stage V.

118 Wisconsin Academy of Sciences , Arts , and Letters.

Beverley.

DOCUMENTS— Continued.

Date Document

1450. Hist. Mss., 135.

1450. Leach (2), p. 214.

1451. Leach (2), p. 217.

1452. Hist. Mss., p. 136.

1452. Chambers, II, 340

Leach (2), p. 216.

1455. Hist. Mss., p. 89.

1466. Leach (2), p. 216.

Content

Places for playing;

Corpus Christi play as¬

signed. “In primis ad

Barras Boriales. Item

apud Bulryng. Item ad

domum Johannis Skip-

with. Item deinde apud

Fischmarket. Item in¬

ter cer— (?) et eam-

panam. Item ad monas-

terium. Item ad Tor-

rentem.”

Expenses of Twelve

Governors governing pa¬

geants. Governors con¬

tribute to craft of Skin¬

ners for their play on

Corpus Christi day.

Alderman of Skinners

paid fine because he did

not produce his play on

Corpus Christi day.

“Porters and Crelers

inferius nominati moniti

sunt praedicto die et

anno quod habeant j

pagendam de novo fac-

tam ad ludendum super

die Corporis Christi prox¬

imo futuro post datum

praesentium seu festum

annunc. B. M. V. prox.

fut. sub poena forisfac-

turae xls. ad usum com-

munitatis. Rob. Thorn-

s k e w, aldermannus,

monitus est hie xvj. die

Jun. ad exponendum vjs.

viijd. eo quod lusores

artis carpentariorum ne-

sciebant ludum suum

die Corporis Christi con¬

tra poenam proclama¬

tion^ communis cam-

panatoris.”

Fine on Henry Cow-

per for not knowing his

part in play (“nesciebat

ludum suum’ ”) on Cor¬

pus Christi day.

“ ‘Payntners, gold-

smyths, masons and gla-

sears.’ — “To be one bro¬

therhood ‘and have

yearly a pageant of

Three Kings de Ce-

lane.’ ”

William Hoseham

warned to put down 40s.

“because the players of

the pageant of the Dy¬

ers’ craft were not

ready to play their pa¬

geant in the first place

at the North Bar.”

Interpretation

Stage V.

Stage Y.

Stage V.

Pierson— The Corpus Christi Procession . 119

120 Wisconsin Academy of Sciences , Arts , and Letters,

Beverley.

Date Document

1485. Hist. Mss., p. 103.

1491. Hist. Mss., p. 103.

1493. Hist. Mss., p. 100.

1493. Bev. Town Doc.,

p. 99, p. 101.

1493. Chambers, II, 340.

1494. Hist. Mss., p. 101.

1498. Hist. Mss., p. 68.

Bev. Town Doc.,

p. 68.

DOCUMENTS— Continued.

Content

to contribute to the

playing and mainten¬

ance of the said pa¬

geant have refused and

refuse, to do so; There¬

fore they made the or¬

dinances — underwrit¬

ten —

First, that all cutlers,

Furberors, Plumbers,

Braziers, Cardmakers.

and Pewterers ( icritten

in a different hand “and

Pynners”) in the liberty

of Beverly there should

be one Brotherhood, and

that they and each of

them should support the

charge of the said pa¬

geant, and should per¬

form and cause the

same to be played for

ever.”

f. 79 headed — “Their

Play the ‘Redemption of

Adam and Eve, called

le Coke Pageant.’ ”

Millers’ Ordinances,

1491. “Pageant of the

Resurrection of Laz¬

arus.”

“Every maker of ‘lade

sadvll panels’ to pay to

maintenance of play and

light 8 d. a year.”

Drapers to play

‘Dooming Pilate’ when¬

ever plays are given.

Regulations on support

of Corpus Christi play.

The play of the Drap¬

ers and Mercers divided,

Drapers taking ‘Demyng

Pylate’ and Mercers

‘Blak Herod.’

Journeymen to pay

toward expenses of Cor¬

pus Christi play when it

is played.

“ ‘Allso it is ordande by

the forsayde xij Gover¬

nors — that the forsayde

xij for tyme beyng shall

go yerly in processyon

on. Corpus Christi day, or

on the morue after, as

itt shall happen, afore

all the aldermen; and

evere man of the other

two bynks [benches]

to go with there aider-

man of ther occupacyon

in thar clothyng belong-

yng to thar brodyr-

hed.’ ”

Interpretation

Stage V.

Stage I. Note that

the day of the proces¬

sion is variable. This

suggests that there was 1

no connection between

the plays given on Cor¬

pus Christi day and the

Corpus Christi proces¬

sion.

Pierson — The Corpus Christi Procession.

121

Beverley.

DOCUMENTS— Continued.

Mr. Davidson suggested that the ‘ ‘ order of the Gilds is a mat¬

ter of importance to us, as the earliest order of the gilds in the

craft plays was doubtless the same as in the procession” (p. 91).

I have arranged in tabular form the list of crafts ( in order) in

the processions for 1430 and 1498, and compared them with a

list of crafts (in order) for the plays of 1520. ' The list of 1520

will serve for the hundred years between 1420 and 1520 since the

same crafts seem to have given the same plays for the whole

period.

1. The play of the Smiths in 1392 and in 1520 is the same.

2. The play of the Bowers in 1411 ( ?), 1413, and in 1520 is

the same.

3. The play of the Barbers in 1414 and in 1520 is the same.

4. The play of the Carpenters in 1420 and in 1520 is the

same.

5. The play of the Bakers in 1428 and in 1520 is the same.

6. The play of the Sadlers in 1441 and in 1520 is the same.

7. The play of the Goldsmiths (Painters, Masons, etc.) in

1455 and in 1520 is the same.

8. The play of the Cooks in 1485 and in 1520 is the same.

9. The play of the Millers in 1491 and in 1520 is the same.

10. The play of the Drapers in 1493 and in 1520 is the same.

11. The play of the Mercers in 1493 and in 1520 is the same.

BevdrUy. DOCUMENTS— Continued,

122 Wisconsin Academy of Sciences, Arts, and Letters.

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Pierson — The Corpus Christi Procession .

123

Conclusion : The order of the gilds in the craft plays is not

the same as in the procession. Moreover, only stages I and V

are represented at Beverley. The length of the plays (see entry

1449) would preclude any connection with the procession.

Therefore, the procession and the plays from 1377 on, seem to

have been distinct.

Bungay.

DOCUMENTS.

Date Document

1514. Chambers, II, 343.

Collectanea (L’Es-

trange editor),

p. 272. (original

document in lat¬

ter).

Content

In 1514 certain per¬

sons “ ‘brake and threw

down five pageants of

the said inhabitants that

is to saye, hevyn pagent,

the pagent of all the

world, Paradyse pagent,

Bethelem pagent and

helle pagent, the whyche

wer ever wont tofore to

be caryed abowt the

seyd town upon the seyd

daye in the honor of the

blessyd Sacrement.’ ”.

Interpretation

“There were pageants

also in the Corpus

Christi processions at

Bungay and at Bury St.

Edmunds, but the no¬

tices are too fragmen¬

tary to permit of more

than a conjecture as to

whether they were ac¬

companied by plays.”

Chambers, II, 162.

In the absence of

further material, I

should assign this ref¬

erence to Stage III.

Conclusion : The material is too fragmentary to be conclusive.

Bury St. Edmunds.

DOCUMENTS.

Date Document

Content

Interpretation

1477.

Chambers, II, 343.

Certain fines are to go

to “ ‘the sustentacione

and m,ayntenaunce of

the payent of the As-

sencione of oure Lord

God and of the yiftys of

the Holy Gost, as yt

hath be customed of

olde tyme owte of

mynde yeerly to be had

to the wurschepe of

God, amongge other

payenttes in the proces-

sione in the feste of

Corpus Xri.’ ”

Notice Chamber’s com¬

ment under Bungay

above.

This may refer to the

drawing of the pageant-

wagons in the proces¬

sion (Stage VI), to

Stage III, or to the pro¬

cessional nature of the

cycle.

Conclusion : The material for Bury St. Edmunds is too frag¬

mentary to be conclusive.

124 Wisconsin Academy of Sciences , Arts , and Letters.

Canterbury.

DOCUMENTS.

Date Document

1490. Hist. Mss., IX, pt.

1, p. 174.

1525. Archseologia Can-

tiana, XVIX, 85.

1527. Arc. Cant., XVII,

88.

1545. Arc. Cant., XVII,

107.

1546. Arc. Cant., XVII,

109.

1547-8. Arc. Cant.,

XVII, 111.

Content

“before this tyme ther

hath bene, by the most

honourable and wor-

shipfull the Cite of

Canterbury — a play

called Corpus Xpi Play —

of late daies it hath bene

left and laide apart.' — -

Wherfore it is enacted

— that from hensforth

every craft — being not

corporate for their non

sufficience of their

crafts, be associate to

some crafte moste ne-

dynge support, yf they

will not labour to be

corporate within them

selfe. — And yf eny suche

crafte — wille not make

suit to the Burgemote

for the reformacion of

the premisses by the

seide feste (of St. Mich¬

ael next ensuing)” they

shall pay a fine and be

subject to punishment.

“Item for a calues

hede flaggis and thredde

at Corpus Christi day

for ryngaris — vijd.”

Same as 1525.

“Item for flaggis bred

and drynke on Corpus

Christi day — ijd.”

For “flaggis” on Cor¬

pus Christi dajr.

Same as 1546.

Interpretation

Stage V.

This and the follow¬

ing citations refer to

the procession as di¬

rected by a church.

Conclusion : The references at Canterbury are too few to be

conclusive.

Pierson — The Corpus Christi Procession ,

125

Chester .

Date Document

1358. Morris, p. 405.

1462. Chambers, II, 352.

Morris, p. 316.

1471. Morris, p. 316.

1520. Morris, p. 316.

1520. Morris, p. 349.

1544-? Chambers, II,

350.

? Morris, p. 309.

Pre-Reformation.

Banns.

DOCUMENTS.

Content

Trades go in Corpus

Christi procession with

candles.

“Baker’s charter re¬

fers to their ‘play and

light of Corpus Christi.’ ”

Half of certain fines

of sadlers are to go to

the support of pageant,

light, and play on the

Festival of Corpus

Christi.

“ ‘The Stuards of the

Founders and Pewters

agree with the Stew¬

ards of the Smiths to

here and draw the

Whitson Playe and Cor¬

pus Christi light.’ ”

Certain gilds to have

Corpus Christi light.

“ ‘On Corpus Xpi day

the Colliges and prestys

bryng forth a play on

the assentement of the

Maire.’ ”

“There will be a ‘sol-

empne procession’ with

the sacrament on Cor¬

pus Christi day from

‘Saynt Maries on the

Hill’ to ‘Saynt John’s,”

together with ‘a play

sett forth by the clergye

In honor of the fest.’ ”

“Also maister maire

of this Citie with all his

bretheryn accordingly

A solempne procession

ordent hath he

To be done to the best

Appon the day of Cor¬

pus Christi;

The blessed sacrament

carried shall be

And a play sett forth

by the clergye

In honor of the fest

Many torches there may

you see

Marchaunts and craftys

of this citie,

By order passing in

their degree.

A goadly sight that day

They come from Saynt

Maries on the hill

The Church of Saynt

John untill

And there the sacra¬

ment leve they will.

The south [sooth] as I

you say.’ ”

Interpretation

Stage I.

Stage V.

The play was prob¬

ably transferred to

Whitsuntide to avoid a

clash with the proces¬

sion.

Chambers, II, 353.

Note that this is not a

craft play.

Note that this is not a

craft play.

126 Wisconsin Academy of Sciences , Arts, and Letters.

Chester

DOCUMENTS— Continued.

Conclusion : The data of composition of the Chester plays,

originally given on Corpus Christi Day, is about 1327- (See

Chambers, Vol. 2). The earliest reference to the Corpus

Christi procession in England is at Ipswich in 1325 (Chambers

II, 371). Clement V. at the Council of Vienne (1311) confirmed

the bull of Pope Urban IV (1264) concerning the festival

(Friedburg, Vol. II, Col. 1174-1177). By 1318, the feast was

celebrated in almost every church in France. The feast was en¬

joined on Canterbury in 1332. The Exeter Ordinale speaks of

Corpus Christi as a novelty in 1337. Therefore, it seems reason¬

able to suppose that the procession was not instituted in Chester

much before 1325. If the date of the plays is + 1327, the de¬

velopment from procession to play (if there was one) must have

been very rapid. Moreover, the Chester Whitsun plays took

three consecutive days. The plays on Corpus Christi day could

not have been much shorter, and therefore could not have been

given during the procession. Plays and procession at Chester,

though given on the same day, seem to have had no connection.

Pierson — The Corpus Christi Procession.

127

DOCUMENTS.

Coventry.

128 Wisconsin Academy of Sciences, Arts, and Letters .

Coventry.

DOCUMENTS— Continued.

Date

Document

1444. Sharp, p. 171.

1445„ Leet Book, p. 220.

1447. Leet Book, p. 231.

1448. Sharp, Diss., p. 163

1452. Sharp, Diss., p. 163.

Content

Procession had been

held time out of mind.

Certain crafts to pro¬

vide wax for the pro¬

cession.

“Pur le Ridyng on

Corpus Christi day and

for Watche on Midsomer

even.

The furst craft, ffysh-

ers and Cokes. Baxsters

and Milners. Bochers,

Whittawers and Glou-

ers. Pynners, Tylers and

Wrightes. Skynners.

Barkers. Coruisers.

Smythes. Weuers. Wir-

drawers. Cardmakers,

Sadelers, Peyntours and

Mason[s]. Gurdelers.

Taylours, Walkers and

Sherman. Deysters, Dra¬

pers. Mercers.”

Order that riding as

from ancient times be

kept up.

Payment for bearing

of torches on Corpus

Christi day — Carpenters’

Accounts.

Same

1 448

entry for

Interpretation

Stage I.

1452. Sharp, Diss., p. 79.

It was ordained that

the “Wryghtes Crafte of

Coventre schall paye to

the Pageant Xs upon

Whytsonday or else by

Corpus XPi daye uppon

the payne of XXs halfe

to the Meyor & halfe to

the Crafte & they to

have no more to doo

wythe the Pageant but

payeing there Xs.”

Contract for the rule'

of the pageant.

Same as entry for

1448.

Queen came to see the

plays.

Every craft that has a

pageant shall make that

pageant ready upon

penalty.

1456 in Poole, p. 44.

Stage V.

Pierson — The Corpus Christi Procession .

129

Coventry.

DOCUMENTS— Continued.

Date Document

1459. Sharp, Dias., p. 160

1461. Sharp, Dias., p. 79.

1463. Sharp, Dias., p. 160.

1469. Sharp, Disa., p. 21.

1473. Sharp, Disa., p. 77.

1476. Sharp, Dias., p. 164.

1477. Sharp, Diss., p. 164.

1481. Sharp, Disa., p. 15.

Chambers, II, 359.

1485. Harris, Coventry,

p. 288.

1487. Sharp, Diss., p. 164.

1487. Harris, p. 288.

Content

From Trinity Guild

accounts — “Exp’s fact in

festo Corpis XPi viz. ad

iiijor Torchberers ad

portend iiijor Tortices p

tempus p cessional circa

le Cowpe in quo continet

Corp dni.”

From Carpenters’ Ac¬

counts — “payd to pyn-

ners & tylers for the

page’t.

“Itm to iiij torchber¬

ers in festo corp is XPi.’’

“It’ for iiij Jaked men

about the parent.”

“R’ Joh’e Thrumpton

& Thoma Colyns custod-

ibz de m cers p reddit de

pagent house.”

“It ffor hors hyre to

Herod.” Smiths’ A c-

counts.

“ ‘Hit is ordened at

this p’ sent leetq that

ev-’y Crafte wt in this

Cite com wt their pa-

geaunts accordyng as

hit haith byn of olde

tyme and to com wt

their p’ cessions & ri-

dyngs also when the

byn required by the

Meir for the worship of

the Cite in peyne of

Xli.’ ”

Payment for bearing

of torches on Corpus

Christi day.

Contract for the rule

of a pageant.

Richard III saw the

plays.

Payment to torch-

bearers and to minstrel

on Corpus Christi day.

Carpenters’ accounts.

Henry YII saw the

plays.

Interpretation

Stage V.

Stage I.

Stage V.

Stage VI. It appears

that the person who

played the part “of

Herod in the Smiths’

pageant joined the Pro¬

cession.” Sharp, p. 164.

The procession and

the plays appear to

have been separate in

1476.

Stage I.

Stage V.

Stage V. Mr. Cham¬

bers, (II, 358) and Mr.

Poole, (p. 44) assign

this item to 1486.

Stage Y.

9— S. A.

130 Wisconsin Academy of Sciences , Arts , and Letters ,

Coventry.

DOCUMENTS— Continued.

Date Document

1492. Sharp, Diss ., p. 9.

1493. Harris, p. 288.

Chambers, II. 358.

1494. Sharp, Diss., p. 81.

1494. Sharp, Diss., p. 163.

1494. Sharp, Diss., p. 79.

1494. Harris, p. 292.

1494. Poole, p. 40.

1494. Sharp, Diss., p. 10.

1496. Sharp, Diss., p. 10.

1497. Sharp. Diss., p. 20,

note.

Content

Chandlers and Cooks

are to contribute to the

support of the Smiths’

pageant.

Henry VII saw plays.

Butchers agree to help

“Whittawers” to sup¬

port their pageant.

Mention of torch bear¬

ers on Corpus Christi

day.

Dyers’ Accounts.

Carpenters ask for

help since they are

charged “with a Pag-

iant, kepyng wacehes ”,

etc.

Dyers refuse to have

pageant.

Mayor and council to

help those overburdened

with pageants by unions

of crafts.

Inhabitants of Gosse-

ford Street ask to have

pageants stand at the

place there.

Cardmakers ask that

the craft of Skinners

and Barkers may pay

towards the charge of

their Pageants.

“Wrights,” “Tylers,”

and “Pynners” ask for

help in supporting pag¬

eant.

“Cappers” and “Pull¬

ers” are to help “Gird-

lers” in supporting

their pageant.

“ ‘It’m for the horss-

yng of the padgeant.’ ”

Interpretation

Stage VI. See com¬

ment on entry 1476. This

does not necessarily re¬

fer to the Corpus Christi

procession. These two

references to Herod’s

riding in the procession

are the only ones of

their kind that have

been found.

Note that this does

not necessarily mean

that the plays developed

from the procession.

Stage V.

Mr. Poole, p. 44 gives

this date as 1492.

Stage V.

Stage I.

Stage V.

1489. Sharp, Diss., p. 167.

Craig, p. XVII.

Chambers, II, 363.

“It’ payd ffor Aroddes

garment peynttyng pt

he went a p’ssayon in.”

Pierson — The Corpus Christi Procession. 181

Coventry .

DOCUMENTS— Continued.

132 Wisconsin Academy of Sciences, Arts, and Letters.

Coventry.

DOCUMENTS— Continued.

Pierson — The Corpus Christi Procession,

133

DOCUMENTS— Continued.

Coventry.

Conclusion : The plays are first mentioned in 1392 ; the pro¬

cession in 1348, when it was apparently under the control of the

Corpus Christi Gild. There is no indication that between 1348

134 Wisconsin Academy of Sciences, Arts, and Letters.

and 1392 the plays were part of the procession, either as mute

mysteries or as spoken drama. The length of the plays makes

it probable that after 1392 at least, they were distinct from the

procession. The puzzling entries in 1476 and 1489 may mean

(and this would substantiate Mr. Spencer’s theory) that, after

the separation of the plays from the procession, characters from

the pageants continued to ride in the procession. The relation¬

ship, however, may be no more than external. The two items

are exceptional. I have interpreted the entries for 1501 and

1539 as follows: At Coventry (as at Chester the “prestes” and

“ colleges”), the religious gilds performed in the procession a

play that had no connection with the craft plays. To conclude —

while we have at Coventry stages I, V, and VI, there is no evi¬

dence that the development followed the consecutive stages I, II,

III, IV, V.

DOCUMENTS.

Dublin.

Date Document Content

1498. Chambers, II, 363. “The Chain Book of

the City contains the

following- memorandum,

— Corpus Christi day a

pagentis: — “The pagen-

tis of Corpus Christi

day, made by an olde

law and confermed by

a semble — :

‘Glovers: Adam and

Eve, with an angill fol-

lowyng berryng a

swerde. Peyn XLs.

‘Corvisers: Caym and

Abell, with an auter

ard the offerance. Peyn

XL. s. ‘Maryners, Vyn-

ters, Shipcarpynderis,

and Samountakers: Noe,

with his shipp, appara-

lid accordyng. Peyn,

XL. s.

‘Wevefs: Abraham

[and] Ysaak, with ther

auter and a lamb and

ther offerance. Peyn,

XL. s.

‘Smythes, Shermen,

Bakers, Sclateris, Cokes

and Masonys: Pharo,

with his hoste. ‘Skyn-

ners, House-Carpynders,

and Tanners, and Brow¬

ders: for the body of

the Carnell and Oure

Lady and her ehil[d]e

well a,pereled, with Jo-

Interpretation

“These pageants, — ap¬

pear from their irregu¬

lar order, to be only

dumb-show accompani¬

ments of a procession.”

Chambers, II, 365.

Stage III, IV, or V?

Pi.'rs(/n — The Corpus Christi Procession.

135

DOCUMENTS— Continued.

Dublin.

Date

Document

1569. Chambers, II, 365.

Content

seph to lede the Cam-

ell, and Moyses with the

children of Israeli, and

the Portors to berr the

Camell. Peyn, XL. s.

and Steyners and Peyn-

tors to peynte the hede

of the Camell. (Peyn,)

XL. s.

‘[Goldsmy] this: The

three kynges of Col¬

ly nn; ridyng worsliup-

fully, with the offer-

ance, with a sterr afor

them.

‘[Hoopers]: The shep-

[er]dis, with an Angill

syngyng Glorea in ex-

celsis Deo. —

‘Corpus Christi yild:

Criste in his Passioun,

with three Maries, and

angilis berring serges

of wex in ther hands. —

‘Taylors: Pilate, with

his fellaship, and his

lady and his kynghtes,

well beseyne. —

‘Barbors: An[nas] and

Caiaphas, well araied

acordyng. —

‘Courteours: Arthure,

with [his] knightes —

‘Fisshers: The Twelve

Apostelis. —

‘Marchauntes: The

Prophetis —

‘Bouchers: tormen-

tours, with ther gar-

mentis well and clenly

peynted. —

‘The Maire of the

Bulring and bachelers

of the same: The Nine

Worthies ridyng wor-

shupfully, with ther fol¬

lowers accordyng. —

‘The Hagardmen and

the husbandmen to berr

the dragoun and to re-

paire the dragoun a

Seint Georges day and

Corpus Christi day.’ ”

“In 1569 the crafts

were directed to keep

the same order in the

Shrove Tuesday ball

riding ( — ) ‘as they are

appointed to go with

their pageants on Cor¬

pus Christi daye by the

Chayne Booke.’ ”

Interpretation

Conclusion : From these two references it is difficult to judge

of the nature of the “pagentis.” Since the Corpus Christi pro¬

cession is not mentioned, I am inclined to think that the quota¬

tions describe spoken drama.

136 Wisconsin Academy of Sciences, Arts, and Letters.

H&reford.

DOCUMENTS.

Date. Document.

[1500 to 1520?] Hist.

Mss., XIII, pt. 4,

p. 304.

1503. Hist. Mss., XIII,

pt. 4, p. 288, 9.

1549. Hist. Mss., XIII,

part 4, p. 305.

Devlin, p. 65.

Content.

“ ‘To the ryght wor-

shypeful mayer of the

cytey of Herefford and

to hys bretherne. She-

wythe unto your good

mastershippes your um-

ble orators the persons

subscribed beyng jour-

nemen of th’ occupacion

of corvesers within this

cytey have obtayned of

your mastershippes pre¬

decessors mayers and

aldermen of the seyd

cytey a composysyon

whereby your sayd ora¬

tors were bound to

bryng furth certen

torches in the proces¬

sion on the day of Cor¬

pus Christi yerelye’ ” — .

“ ‘The paiants for the

procession of Corpus

Christi

Furst, Glovers — Adam,

Eve. (Jayne and Abell

(.erased).

Eldest Seriant — Cayne,

Abell, and Moysey, Aron,

Carpenters — Noye ship.

Chaundelers — Abra¬

ham, Isack, Moysey.

Bakers — Knyghtes in

harnes.

Journeymen Cappers

— Seynt Keterina with

tres(?) tormentors!”

“ ‘fforasmuche as ther

was before thys tyme

Dyvers corporacions of

Artiffycers, craftes, and

occupacions in the sayd

cytty who were bounde

by the grante of their

corporacions yerelye to

bryng fforth and sett

forward Dyvers pa-

geauntts of Auncyentt

historyes in the proces¬

sions in the sayde cytty

upon the Day and ffeast

of Corpus Xti [ — ],

which nowe ys and are

omytted and surceased,

whereof it ys Agreed,

condescended, and gran¬

ted that all and everye

of the sayd craftes and

corporacions shall in

stede and place of the

settynge fforthe of the

sayd pageauntts on the

sayd daye or ffeast of

Corpes Xti, yerely con-

sente to pay att the

ffeaste of the Annuncy-

acion — one Annuyte — to

the vse — of the sayd

ctey.’ ”

Interpretation.

Stage I.

“The 1503 list seems

to concern a dumb show

only.” Chambers, II,

368.

Stage III.

Stage III or Stage IV.

Pierson — The Corpus Christi Procession. 137

Conclusion: At Hereford, to judge from the material we

have, certain mute mysteries were performed in the course of

the Corpus Christi procession through the city. The quotations

are too late, however, to help the theory very strongly.

DOCUMENTS.

Ipswich.

138 Wisconsin Academy of Sciences , Arts, and Letters.

DOCUMENTS— Continued.

Ipswich.

Conclusion: Until the meaning of the word ‘ e pageantes 1 1 is

determined, one can not tell whether the plays were performed

during the procession, or whether the pageant wagons were

merely drawn in the procession and the plays performed later

in the day.

Pierson — The Corpus Christi Procession ,

139

DOCUMENTS,

King’s Lynn.

140 Wisconsin Academy of Sciences, Arts, and Letters.

Lincoln.

DOCUMENTS— Continued.

Date Document

1615. Spencer, p. 81.

1518. Spencer, p. 62.

1664-5. Chambers, II,

379.

Content

“At Lincoln in 1515

the players not only

were required to go in

character in the proces¬

sion, but constables

were stationed ‘to wait

upon the array in pro¬

cession, both to keep the

people from the array,

and also to take heed of

such as wear garments

in the same.’ ”

“In the early years of

the Corpus Christi festi¬

val, when the proces¬

sion and the plays were

all one, the ceremonies

of the day seem to have

begun at an early hour

in the morning-. — What

the exact hour was in

the earliest years of the

procession we do not

know. — At Lincoln in

1518 it was at seven

o’clock in the morning.”

In 1554 and 1555 “ ‘it

was ordered that St.

Anne’s Gild with Corpus

Christi Play shall be

brought forth’ and

played this year.’ ”

Interpretation

1515, 27 July. — “It is

agreed that whereas di¬

vers garments and other

‘heriorments’ are yearly

borrowed in the country

for the arraying of the

pageants of St. Anne’s

Guild, but now the

knights and gentlemen

are afraid with the pla¬

gue so that the ‘grace-

man’ cannot borrow

such garments, every

alderman shall prepare

and set forth in the said

array two good gowns

and every sheriff and

every chamberlain a

gown, and the persons

with them shall wear

the same. And the con¬

stables are ordered to

wait upon the array in

procession, both to keep

the people from the ar¬

ray, and also to take

heed of such as wear

garments in the same.”

Hist. Mss., XIV, pt. 3,

p. 25.

1518, 16 June. “Ord¬

ered that every aider-

man shall send forth a

servant with a torch to

be lighted in the pro¬

cession with a rochet

upon him about the Sac¬

rament, — And also send

forth one person with a

gown upon his back to

go in the procession.

That every constable

shall wait on the pro¬

cession on St. Anne’s

day by 7 of the Clock,

upon pain of forfeiture

of 12 d.” Same, p. 26.

I do not believe that

the two entries for 1515

and 1518 refer to the

Corpus Christi proces¬

sion for these reasons:

1. The procession is

evidently the procession

on St. Anne’s day.

2. The dates of the

enactments (June, July)

are more appropriate

for St. Anne’s day than

for Corpus Christi day.

Pierson — The Corpus Christi Procession . 141

Conclusion : The plays and the procession at Lincoln were evi¬

dently on different days. After ruling out the quotations for

1515 and 1518, one has little material on which to base any con¬

clusion. Between 1328 and 1478, the play may have been given

on Corpus Christi Day. In the sixteenth century the plays were

certainly on St. Anne’s day. Mr. Leach (in A Miscellany Pre¬

sented to Dr. Furnivall, p. 226) notes that the play of St. Anne

did not differ much from the Corpus Christi plays. The close

relation between the St. Anne procession and the play of St.

Anne may then be an argument for a similar close relation be¬

tween the Corpus Christi procession and the Corpus Christi

plays.

DOCUMENTS.

London.

142 Wisconsin Academy of Sciences , Arts , and Letters.

London.

DOCUMENTS— Continued.

Date Document

1477- 9. Littlehales,

p. 81.

1478- 81. Littlehales,

p. 100.

1487-88. Littlehales,

p. 131.

1489-90. Littlehales,

p. 149.

1491-2. Littlehales,

p. 173.

1519-20. Littlehales,

p. 305.

1523-24. Littlehales,

p. 322.

1524-25. Littlehales,

p. 330.

1648. Nichols, Grey

Friars Chronicle.

1554. Same, p. 89.

1557. L. T. Smith, York

Plays, p. LXIV.

Content

Garlands for proces¬

sion.

Flags, garlands,

torches for procession.

Laten bells for cana-

pye on Corpus Christi

day.

Children for the pro¬

cession.

Roses for the proces¬

sion.

Garlands for the pro¬

cession.

Garlands for crosses

and chair and for

strangers who bore

“eopis”.

Bearing of crosses on

Corpus Christi day.

“This same yere was

put downe alle goyng

abrode of processyons, —

and the skynners’ pro-

cessyon on Corpus

Christi day.”

“Item the XXIIIj day

of May was Corpus

Christi day that some

kepte holy day and some

wolde not, and there

was a joyner — he was in

Smythfelde when the

procession of sent

Pulchers came by hym,

and he wold a tane the

sacrament from the

prest.”

Passion of Christ giv¬

en on Corpus Christi

day.

Interpretation

See comment on pre¬

ceding.

See comment on pre¬

ceding.

See comment on pre¬

ceding.

See comment on pre¬

ceding.

See comment on pre¬

ceding.

See comment on pre¬

ceding.

See comment on pre¬

ceding.

See comment on pre¬

ceding.

See comment on pre¬

ceding.

See comment on pre¬

ceding.

Conclusion : There is only one mention of a play on Corpus

Christi day ; so that it is difficult to connect it with the proces¬

sion, which seems to have been in charge of the individual

churches, and of a religious gild, not of the crafts.

143

Pierson — The Corpus Chrisii Procession .

Newcastle.

DOCUMENTS.

Date Document

1426-7. Chambers, II,

385.

1436. Brand, Newcastle,

II, 370.

1437. Brand, Newcastle,

II, 370, note.

1442. Brand, Newcastle,

II, 370.

1451. Brand, Newcastle,

II, 370.

1454. Chambers, II, 424.

1459. Brand, Newcastle,

II, 370.

1477. Brand, Newcastle,

II, 370.

1479-80. Surtees, Vol. 93,

p. 4.

20 ED. IV.

Brand, History and An¬

tiquities of Neivcastle, II,

371.

Content

Corpus Christi plays

first mentioned.

Corpus Christi play

mentioned in ordi¬

naries of smiths and

glovers.

Skinners’ ordinary

mentions plays.

Plays mentioned in

ordinary of barbers.

Plays mentioned in

ordinary of slaters.

Creation of Adam and

Flight into Egypt played

by Bricklayers and

Plasterers.

Plays mentioned in

ordinary of sadlers.

Plays mentioned in

ordinary of fullers and

dyers.

“Also it is asentit,

— by the said Felleship,

— t hat wppon Corpus

Christi Day yerly, in

honoryng and worship-

pyng of the solemp pro¬

cession, every man of

the said Felleship beyng

within the tranches of

4his town the said day

as it shall fall, shalle

apper in the [Beer

interlined ] Marcoth by

VIj of clok in the morn-

yng. — Also that thair

be a rowll mayd of all

the names of the same

Fellowship, for the said

procession, and accord -

yng to that rowll callyd

by the Clark, the lattast

mayd burges to go for-

mest in the procession —

Provyded always that

all those of the said Fel¬

leship that shalbe Mair,

Shereff, and aldermen,

with thaire officers and

servandes, than beyng,

attend wppon the holy

sacramente. Provydet

also, that all those of

the said Felleship as

beyn maires, shereffs,

and aldermen in yerys

by passyt, shall go prin-

cypall in the sayd sol¬

emp procession, accord-

yng as they war chossen

into the sayd officese.”

Interpretation

Stage V.

The plays were ap¬

parently processional.

The pageants may first

have taken part in the

Corpus Christi proces¬

sion proper and after¬

wards have gathered in

:a field.

Chambers, II, 385.

“The difficulty seems

to have been solved at

Newcastle by sending

the pageants around

with the procession in

the early morning and

deferring the actual

plays until the after¬

noon.” Chambers, I,

162.

Stage I.

144 Wisconsin Academy of Sciences, Arts, and Letters.

Newcastle.

DOCUMENTS— Continued.

Conclusion : The plays and the procession at Newcastle seem

to have been distinct. The items for 1561 and 1568 may repre¬

sent stage VI, but more probably they refer to the processional

nature of the plays.

Pierson — The Corpus Christi Procession ,

145

Norwich.

Date Document

1478. Chambers, II, 386.

1489. Waterhouse,

p. XXX-XXXI.

Chambers II, 389.

1527. Waterhouse,

p. XXIX.

1534. Chambers, II, 387.

1535. Waterhouse,

p. XXXI.

1536. Waterhouse,

p, XXXI.

DOCUMENTS.

Content

J. Whetley writes

from Norwich on Corpus

Christi day " ‘at hys be-

yng ther that daye ther

was never no man that

playd Herrod in Corpus

Christi play better and

more agreable to his

pageaunt than he dud.’ ”

In 1849 it was ordered

that the thirty-one

guilds of the town, on

Corpus Christi Day,

should go in procession

before the pageants

“ ‘ad Capell, in Campis

Norwici, modo sequi.’ ”

The procession was ar¬

ranged in the following

order: the thirty-one

guilds, the pageants, the

Shreves clothing, Mr.

Shreve, the Mair’s cloth¬

ing, Maister Mair and

Maister Aldermen with

bokes or beads in their

hands.

“For some time previ¬

ous to 1527, the St.

Luke’s Guild, consisting

of the pewterers, bra¬

ziers, plumbers, bell-

founders, glaziers, stey-

ners and several other

crafts, had apparently

become responsible for

the entire management

of, and outlay in con¬

nection with, the Cor¬

pus Christi plays; and

in tfhat year, finding

themselves, as a result

of this, almost in a

bankrupt condition, they

petitioned the corpora¬

tion to divide the re¬

sponsibility and expense

among the various

guilds.”

4 surveyors of pageant

of grocers chosen. As¬

sessment made on Gro¬

cers “for the pageant

and the Corpus Christi

procession.”

Performance of Cor¬

pus Christi play.

Performance of Cor¬

pus Christi play.

Interpretation

Stage V.

Note in Waterhouse p.

XXIX: The pageants

are “referred to always

as the procession.”

There is no proof that

by procession is meant

pageants.

In 1527 St. Luke’s

guild urges that “ ‘where

of longtime paste the

said Guylde of Seynt

Luke yerely till nowe

hath ben used to be

kept and holden within

the kcitie aforesaid upon

the Mundaye in pente

eoste weke at which

daye and the daye next

ensuying many and di¬

vers disgisyngs and pa-

geaunts — that every oc-

cupacion wythyn the

seyd citye maye yerly

at the said procession

upon the Mondaye in

Pentecost weke sette

“forth one pageaunt.’ ”

Chambers, II, 387.

Mr. Waterhouse has

evidently misinterpreted

the passage.

Stage I and Stage V.

The procession and the

plays were apparently

separate.

Stage V.

10— S. A.

146 Wisconsin Academy of Sciences , Arts , and Letters .

Norwich.

DOCUMENTS— Continued.

Date Document

1538. Waterhouse,

p. XXXI.

1539. Waterhouse,

p. XXXI.

1540. Waterhouse,

p. XXXI.

1541. Waterhouse,

p. XXXI.

1542. Waterhouse,

p. XXXI.

1546. Waterhouse,

p. XXXIII, note.

1546. Waterhouse,

p. XXXI.

1556. Chambers, II, 385.

1558. Chambers, II, 389.

Content

Performance of Cor¬

pus Christi play.

Performance of Cor¬

pus Christi play.

Surveyors apparently

^contracted for the per¬

formance of the plays.

Plays apparently per¬

formed.

Plays apparently per¬

formed.

“ ‘Accordyngly were

chosen 4 Aldermen & 8

Comyners — ; 2 Warde-

yns & 2 Surveyors for

settyng forth pe Pro¬

cession on Corpus Xi

day, & for pe Pageant

yf it go forth pe next

year.’ ”

After 1546 p lays

waned.

From 1556 on “ ‘Gryf-

fon,’ ‘Angell,' and Pen-

don’ of the Corpus

Christi procession, with

flowers, grocery, and

fruit ‘to garnish ye tre

wth’ &., appear alone

in the accounts.” (Gro¬

cers’)

Corpus Christi pro¬

cession mentioned in

grocers’ records until

1558. “They seem to

have been represented

by the ‘griffon’ from the

top of their pageant, a

banner with their arms,

a crowned angel, and an

emblematic tree ‘of fruit,

and grocery.’ ”

Interpretation

Stage V.

Stage I and Stage V.

The procession and the

plays seem to have been

distinct.

Stage II. It does not

appear that the pa¬

geants had any connec¬

tion with this proces¬

sion.

Does emblematic tree

indicate a play in the

procession?

Conclusion : The plays and the procession during the period

covered by the available documents seem to have been distinct.

Both statements : — that plays were performed in the procession*

and that the pageant wagons were drawn in it, — are mere con¬

jectures.

Pierson — The Corpus Christi Procession. 147

148 Wisconsin Academy of Sciences, Arts, and Letters .

Shrewsbury.

DOCUMENTS— Continued.

Date Document

1546-7. Hist. Mss., XV,

pt. 10.

? Owen, p. 64.

? Salopian, p. 11.

Sharp, p. 171.

Owen, p 65.

? Hist. Mss., XV, pt.

10, p. 10.

Content

“Pro vino et tortis da-

tis ballivis et associatis

suis in festo Corporis

Christi euntibus in pro-

cessione.”

“Preceded by their

Masters and Wardens,

and graced with col¬

ours” the companies “at¬

tended the Bailiffs and

members of the Corpor¬

ation, who with the

Canons of St. Chad and

St. Mary, the Friars of

the three convents, and

the Parochial Clergy,

followed the holy Sac¬

rament” on Corpus

Christi day.

The Tailors had at

one time in the proces¬

sion “Adam and Eve,

before whom a large

bough was borne, from

which an apple was oc¬

casionally plucked; and

two knights with drawn

swords.

Procession of crafts

with emblematical de¬

vices.

“ ‘Ordinacio proces-

cionis artificum ville

Salopie in festo Corporis

Christi.’ The companies

were in the following

order: Molendinarii, Pis-

tores, Piscatores, Coci,

Carnifices, B a r c a r i i,

(Tanners), Cordewena-

rii, Fabri, Celarii, Car¬

pentaria Flechers, Cow-

pers and Bowers, Tex-

tores, (T)onsarii cum

Barbitonsoribus [Ci]ro-

tecarii, [Scijssores.”

Interpretation

Stage I.

If this statement is

authentic, it refers to

Stage III.

Before Reformation,

tableaux were usually of

a biblical or ecclesiasti¬

cal nature; after of

mythological or histori-

Hibbert, p. 117.

Stage I.

Conclusion: There is no evidence that the procession ever

developed beyond stage II or at most stage III. I found no ref¬

erence to spoken drama on Corpus Christi day.

Pierson— The Corpus Christi Procession,

149

DOCUMENTS.

York.

150 Wisconsin Academy of Sciences, Arts, and Letters.

York.

DOCUMENTS— Continued.

Pierson — The Corpus Christi Procession .

1*1

York.

D O CUME NT S— Continued.

Date Document

1426. Sharp, Disserta¬

tion, pp. 133, 134.

1428. Antiquary, XI, 108

1430-1440 .

1431. Register of Guild

of Corpus

Christi, p. 251.

252.

1443. Antiquary, XI, 108

1475. Surtees, CXX, 134

1477. Surtees, CXX, 134

Content

“ ‘Whereas for a long

course of time the arti¬

ficers and tradesmen of

the city of York have, at

their own expence, acted

plays; and particularly

a certain sumptuous

play, exhibited in sev¬

eral pageants, wherein

the history of the old

and new testament in

divers places of the said

city, in the feast of

Corporis Christi, by a sol¬

emn procession is repre¬

sented in reverence to

the sacrament of the

body of Christ. Begin¬

ning first at the great

gates of the holy Trin¬

ity in York, and so going

in procession to and into

the Cathedral Church of

the same; and after¬

wards to the hospital of

St. Leonard, in York, leav¬

ing the aforesaid sac¬

rament in that place/ ”

Friar Melton induces

the people to have the

play on one day and the

procession on the sec¬

ond.

Smiths charge mar¬

shals with not paying

their pageant silver.

Finally agreed that they

shall “of thair bather

costages bryng furthe

pair bather playes, and

uphold thair torches in

r>e procession of Corpus

Xpi day.”

Register .

“Agreement between

the mayor and citizens

of York and the keep¬

ers of the Guild [of Cor¬

pus Christi] about car¬

rying the shrine in the

annual procession on the

feast of Corpus Christi.”

Same as 1428.

Those who are to help

girdlers bring out pa¬

geant mentioned.

Certain fines in craft

of cutlers to go to the

support of their pa¬

geant.

Interpretation

Stage IV(?)

If the word procession,

refers to the Corpus

Christi procession and

not to the processional

nature of the cycle, we

have here very definite

evidence for stage IV of

the Davidson-Spencer

theory.

The courses of the

procession and of the

plays are suspiciously

alike. See entries for

1399. After 1426 the

plays were separated.

Stage V.

152 Wisconsin Academy of Sciences , Arts , and Letters,

York.

DOCUMENTS— Continued.

Date

Document

1478. Davies, (18 Ed.

IV), p. 75 ff.

Content

1478. Davies, p. 18, 63,

65. (18 Ed. IV.)

1479-1480. Surtees, CXX.

135.

1483-1484. Hist. Mss., I,

108.

1485. Surtees, CXX, 186

1490. Surtees, CXX, 201

Same, p. 202.

1493. L. T. Smith, York

Plays, p. XLI.

1493. Davies, p. 257.

“And in expenses in¬

curred this year by the

mayor, aldermen, and*?

many others of the

council of the chamber

at the Feast of Corpus

Christi, seeing- and di¬

recting the play in the

house of Nicholas

Bewyk, according to

custom — and 3s. 4d. paid

to one preaching and

delivering a sermon on

the morrow of the said

feast, in the cathedral

church of St. Peter of

York, after the celebra¬

tion of the procession.”

Paid for banner for

Corpus Christi play.

Paid for repair of ban¬

ners of Corpus Christi

play.

Certain fines in crafts

of cutlers and blade-

smiths to go to support

of their pageants.

Innholders contract

for a space of eight

years following to bring

forth yearly their pa¬

geant of the Coronation

of Our Lady.

Those who shall con¬

tribute to pageant of

girdlers enumerated.

Certain fines in craft

or ironmongers to go

to support of pageant.

Those who are to as¬

sist ironmongers in sup¬

porting pageant named.

The pageant of the

ironmongers needs re¬

pairs.

All the masters of the

craft of the “Spuriers

and Lorymers” “ ‘shall

attend vppon yer pai-

aunt’ ” from the begin¬

ning of the play to the

end.

Award to craft of

cordwainers that “ ‘when

the procession were sol-

empnely done the mor-

owe next after Corpus

Xpi day, to bere their

torches- honestly made

and lighted with the

craft of the weavers

and going of the weav¬

ers’ left handes, as had

been there afore acus-

tomed.’ ”

Interpretation

Stage V.

This quotation shows

that in 1478 procession

and plays were distinct.

Stage V.

In 1493, the plays and

the procession w«re sep¬

arate.

Pierson — The Corpus Christi Procession,

DOCUMENTS— Continued.

York.

Date Document

1501. Antiq., XXIII, 29.

1505.. Eng. Hist. Rev.

IX, 301. '

1535. Davies' p. 258.

1547. Davies, p. 260.

1548. Davies, p. 262.

1550. Davies, p. 262.

1552. Davies, p. 262.

1554. Davies, p. 263.

1559. Davies, p. 266

1560. Davies, p. 266.

1562. Davies, p. 266.

1563. Davies, p. 266

1564. Davies, p. 266.

1565. Davies, p. 266.

Content

Thomas Drawswerd

was admitted into Holy

Trinity Guild on the

condition “ ‘that the said

Thomas shall mak the

Pagiant of the Dome

belonging to the Mrcha-

unts of newe substan-

ciale in eury thing pr

vnto belonging.’ ”

Drapers ask for help

in supporting their pa¬

geant.

Performance of Cor¬

pus Christi play suspen¬

ded.

Fines on gildmen for

not furnishing torches

for “ ‘procession the Fri¬

day after Corporscristy

day.’ ”

Plays given except

pageant of “ ‘the denyng

of our lady, the assump¬

tion of our lady, and

the coronacion of our

lady.’ ”

No play.

“ ‘Corpus Xpi playe

shall (God willyng) be

played this yere, and

billets to be made forth

as hath been accus¬

tomed, and that thoes

pagiauntes tharof that

were last forth shall be

played ageyne as before

tyme they were; and

also that the xij and

XXIIIjor, and all other

occupacions accustomed

to have torches shall

have warnyng to pre¬

pare every man for their

torches ageynst the

sayd Corpus Xpi day.’ ”

No Corpus Christi

play.

No Corpus Chlristi

play.

Corpus Christi play

given.

Corpus Christi play

given.

Observance of Corpus

Christi play suspended.

Observance of Corpus

Christi play suspended.

153

MM

Interpretation

Stage V.

In 1547, the plays and

the procession were dis¬

tinct.

Stage V.

Does the reference to

torches in this entry,

mean that the proces¬

sion and the plays were

on the same day?

Stage Y.

154 Wisconsin Academy of Sciences, Arts, and Letters.

DOCUMENTS— Continued.

York.

Pierson — The Corpus Christi Procession .

155

DOCUMENTS— Continued.

York.

Conclusion: The material for York presents some difficul¬

ties. After 1426, procession and pageants were on different

days. Before 1426, the plays may, if the entry for 1426 has

been correctly interpreted, have been acted during the proces¬

sion. The course of the two through the city was the same.

Both started at Holy Trinity (entries 1399, 1426) and stopped

at St. Peter’s and at St. Leonard’s. Obviously more material,

covering the period from 1325 to 1425, must be found to settle

the matter.

The following material is too fragmentary to be conclusive.

The first part of it concerns towns where only the procession

is mentioned; the second concerns towns where only the playa

are mentioned.

DOCUMENTS.

156 Wisconsin Academy of Sciences , Arts, and Letters.

Pierson — The Corpus Christi Procession ,

157

Date Document

Durham .

Longstaffe, in Aeliana.

N. S., II, 59.

Great Yarmouth.

1388-9. Gross, I, 119,

note.

Leicester.

1349-1350. Davidson, p.

93.

DOCUMENTS— Continued.

Content

Also there was a

goodly shrine in Sacte

Nicholas church. or-

deyned to be caryed ye

sayd daie in Procession

— wherin was enclosed

the holy sacramt of

thaulter and was caryed

ye said daie with iiij

preistes vp to ye place

grene & all ye hole pros-

sessio of all ye churches

in ye said towne goyng

before ytt and when it

was a litle space wth in

Wyndshole yett yt dyd

stand still, then was

Sacte Cuthb:. B a n n’

browghte fourth wth two

goodly faire crosses to

meete yt and ye por

& covent wth all ye

whole companye of ye

Quere all in there best

copes dyd meet ye said

shrine sytting on there

kneys and praynge. The

prior did sence yt (fetch

it, Cos.) and then cary-

inge yt forward into the

abbey church ye por and

covent wth all the quere

following yt. — all ye

Bann’ of ye occupac’ons

dyd followe ye saia

shrine into ye church

goyng Rownde about

Saincte Cuthb: fereture

lyghtinge there Torches

& burning all ye s’vice

tyme. Then yt was car¬

yed frome thence wth ye

said p’ssessio of ye

towne back againe to ye

place from whence it

came & all the Ban’p

of ye occupac’ons fol¬

lowing it, & setting yt

againe in ye church.”

This procession was

continued until about

1770.

Society of Corpus

Christi maintained a

light “ ‘circa Corpus

Christi annuatim in die

Corporis Christi.’ ”

“The Gild of Corpus

Christi of Leicester,

which contributed to

the most splendid pro¬

cession in the city ex¬

cept that of St. George.”

Interpretation

From a roll of 1600.

Although the crafts

carried banners in the

procession, pageants ap¬

parently never devel¬

oped.

Stage II.

Cos. refers to Cosin’s

MS.

The crafts apparently

were not concerned with

the procession.

The crafts apparently

did not have charge of

the procession.

158 Wisconsin Academy of Sciences, Arts, and Letters.

DOCUMENTS— Continued.

Pierson — The Corpus Christi Procession.

159

DOCUMENTS — Continued.

CONCLUSION.

My conclusions as to the relation between the Corpus Christi

procession and the Corpus Christi play may be enumerated as

follows :

I. In no case are all the stages to be found.

II. In Beverley, in Chester, in King’s Lynn, and in Lincoln,

stages I and V are represented. In Aberdeen stages I, II, Y,

VI ; in Newcastle I, Y, VI ( ?) ; in Norwich I, II, Y ; in York I,

II, IY to 1426 ( ?), Y ; in Coventry I, Y, VI, (?) are found.

III. Stage III is found apparently at Bury St. Edmunds,

Bungay, and Hereford, but in each case the material is too frag¬

mentary to be conclusive. The material for Dublin, I believe,

refers to complete plays, processional in nature, but not given

in the Corpus Christi procession.

IY. In many places there seem to have been plays and no .

procession, or a procession and no plays.

1610 Wisconsin Academy of Sciences, Arts, and Letters.

V. The length of time required to give the plays precludes

any long connection with the procession.

YI. The date of composition of the plays in each town must

be discovered before one can say definitely that the plays grew

out of the procession.

VII. If one wishes to find the real relationship between the

Corpus Christi procession and the Corpus Christi plays, he must

find it between 1311 and MOO. After 1400, we have spoken

drama in almost all the towns considered. Therefore all con¬

nection between plays and processions after 1400 must be

external, merely hinting at the earlier internal relationship.

We have as yet no material covering the period fully. The

source material for the history of the plays in each town must be

examined separately, and the conclusions obtained must be ap¬

plied only to the individual places from which they were drawn.

Pierson — The Corpus CJiristi Procession.

161

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166 Wisconsin Academy of Sciences , Arts, and Letters.

THE HEAT BUDGETS OF AMERICAN AND EUROPEAN

LAKES.

Edward A. Birge.

Notes from the Laboratory of the Wisconsin Geological and

Natural History Survey. VII.

This paper was suggested during the preparation of a report

on the Finger lakes of New York, which was recently completed

by Mr. C. Juday and myself, under the direction of the United

States Bureau of Fisheries. (Birge and Juday, ’14.) That

part of the report which deals with the temperatures of these

lakes was written by myself and its preparation led me to com¬

pare the results of our observations with those made on similar

lakes in Europe.

In this paper an inland lake of the first class is defined as one

whose size and depth are such as to permit the lake to acquire

the maximum amount of heat possible under the weather condi¬

tions of the season. The lower limits for such lakes in the east¬

ern and central United States seem to be about 10 km. of length

with, at least, 2 km. of breadth ; and 30 m. of mean depth, which

means 50 m. or more of maximum depth. Such lakes must also

lie under ordinary conditions of topography and altitude. Lakes

whose conditions of climate or location are exceptional, such as

those of alpine lakes at considerable elevations, cannot be com¬

pared directly with those in lower and more normal situations.

(See Birge and Juday, ’14, p. 561.)

The European lakes selected for comparison are, with one or

two exceptions, all of the first class as regards size and depth.

They lie at elevations rarely exceeding 500 m., so that the eleva¬

tion can not play any considerable part in determining their

heat cycle. These lakes are chiefly taken from lists used by

Birge—Heat Budgets of American and European Lakes. 167

Forel and Halbfass in their discussion as to the effect of latitude

on the heat budget. The mean temperatures employed are

chiefly taken from two papers by Halbfass (Halbfass ’05, TO).

Without these compilations of numerous temperature observa¬

tions from many authorities, gathered and computed with great

care and labor, this paper could not have been written. In some

cases I have checked Halbfass’ results and usually have found

that my computations closely agree with his. In some other

cases, I have computed for myself the mean temperature ot

European lakes and such results are marked by a star in

Table A.

No doubt more complete study will show that the heat bud¬

gets of the several lakes are influenced by elevation, surround¬

ings, size of affluents and effluent, climate, cloudiness, latitude,

and other conditions as well, but at present a direct and unmis¬

takable effect of any one of these conditions can be pointed out

in very few cases and it is best to consider the budgets in gross

and without too much attention to particular circumstances.

Certain preliminary questions must be discussed before pro¬

ceeding to the comparison of American and European lakes.

These are: (A.) The definition of the heat budget. (B.) The

unit which should be employed in stating it. (C.) The value of

temperature observations, such as those from which the budgets

have been computed.

A. Three things may be understood by the heat budget of a

lake :

1. The amount of heat necessary to raise its water from 0° C.

to the maximum temperature found in summer. This may be

called the gross or crude heat budget.

2. The amount of heat necessary to raise its water from the

minimum temperature of winter to the maximum summer tem¬

perature. This may be called the annual heat budget.

3. The amount of heat necessary to raise its water from 4°C.

to the maximum summer temperature. This may be called, for

reasons explained in the report on the Finger lakes, the wind-

distributed heat or the summer heat-income. (See Birge and

Juday, T4, p. 562.)

Of these three conceptions, the first is of least value, since

it does not correspond to any facts in nature. No lake falls to

0° in winter, so that this temperature is not a starting point

for any actual gains or a terminal point for losses. Still further ;

168 Wisconsin Academy of Sciences, Arts, and Letters.

the same conditions that limit the rise of temperature of a lake

in summer limit also its fall in winter. Thus the gross heat

budget may differ widely in lakes whose actual gain of heat per

unit of surface are closely similar.

A single example of this may suffice. If the mean summer

temperature, as given in Table A, of Owasco, Cayuga, and Sen¬

eca lakes is multiplied by the respective mean depths of the

lakes, the products will be 39,800 cal., 50,500 cal., 68,300 cal., re¬

spectively — sums which represent the number of gram calories

per sq. cm. of surface of the lake necessary to raise its water

from 0° to the summer temperature. These sums differ very

greatly; but if the mean temperature of the water in winter is

multiplied by the mean depth the result shows how much heat

was left in each lake at the winter minimum. These amounts

are for Owasco lake 2,400 cal., for Cayuga, 12,200 cal., and

for Seneca, 30,000 cal. When these sums, which are no part

of any actual heat budget, are subtracted from the gross

sums, the remainders (which represent the annual heat bud¬

get) are closely alike; 37,400 cal., 38,300 cal., and 38,300 cal.,

respectively. These numbers show that the indication of the

gross heat budget is incorrect, and that each unit of surface of

these lakes, in spite of wide differences in area and depth (see

Table A.), is capable of taking up the same amount of heat from

sun and sky ; and this is a fact of no little interest.

The second conception, that of the annual heat budget, is a

statement of facts of the first order of importance in the heat

cycle of a lake. This method, therefore, which was used by

Halbfass, in his comparisons of lakes, is by far the most funda¬

mental of these three conceptions, and should always be used

where the data are at hand.

The third conception, that of the wind-distributed heat, or

summer heat-income, is one which serves much the same purpose

as the second in cases to which it is applicable. It may be used

in many cases where winter data are lacking, as is still true for

many lakes. It applies only to the temperate lakes of ForePs

classification. It has no significance for polar lakes, which do

not rise above 4° ; and in reference to tropical taxes it has the

same difficulties that apply to the gross heat budget, as noted

above. But for temperate lakes, the temperature of 4° con¬

stitutes, not a terminal point, but an important turning point

in their annual temperature cycle, and most such lakes may

Birge—Heat Budgets of American and European Lakes. 169

fairly be compared with each other on the basis of their wind-

distributed heat as well as on that of the annual heat budget.

As shown by Table A, the conclusions to be drawn from the two

types of budget are closely similar.

This statement holds for lakes which lie so far to the south

that the temperature of 4° is reached early in the open season.

If a lake lies far to the north it may not reach that temperature

until a date so late as seriously to affect the remainder of its

budget. This is clearly the case with Ladoga (See Table A, Fig.

1, and p. 45), in which 15,000 cal., or nearly one-half of its bud¬

get, were required to raise the water from the winter tempera¬

ture to 4°.

This is in itself a large amount, although it is not so large as

that demanded by Mjosen in 1901, and it is only slightly larger

than that called for by Skaneateles lake in 1911. But the water

of Mjosen reaches 4° early in June and the New York lakes

reach that temperature early in May or even in April. Homen

states (’02, p. 3) that Ladoga does not warm to 4° until the end

of July. If this is the case, it is surprising that Ladoga accumu¬

lates so much heat above 4° as 18,000 cal. per sq. cm., since Ho¬

men also states that the temperature of the surface begins to fall

by the end of August. In any case, it is clear that the summer

heat-income of Wettern, Mjosen, and Ladoga is reduced by the

late date at which the temperature of 4° is reached. It is also

clear that the summer heat-income of these lakes cannot fairly

be compared with that of lakes further south.

But the summer heat-income of lakes which reach 4° in April

or early May in but little, if at all, affected by the amount of heat

necessary to raise the water to 4°. The gains of heat may go on

well into August, or even later, and so much heat is delivered to

the lake which cannot be absorbed in any case, that in these lakes

the summer heat-income is practically independent of the date

when the temperature of 4° is reached, or of the amount of heat

needed to warm the water to that point.

Thus, the size of each of the two members of the annual heat

budget— -that below and that above 4° — is independent of that

of the other in many lakes. The variations in the lower part

of the budget are due to winter conditions and are especially

dependent on irregular accidents of weather rather than on the

average conditions of the season. This part of the budget,

therefore, introduces a considerable element of variation into

170 Wisconsin Academy of Sciences, Arts , and Letters .

the annual heat budget, which has little, or nothing, to do with

the capacity of the lake for absorbing heat. This capacity may

often be more exactly shown by the wind-distributed heat than

by the annual heat budget.

The fact that winter losses and summer gains are measurably

(although not wholly) independent of each other may also be

seen in the budgets of tropical lakes, like Como and Geneva.

Both lakes show greater variations in the winter than in the

summer temperatures, and the annual heat budgets of both lakes

are more variable than is common in lakes which regularly

freeze during the winter.

B. The heat budget in any of its forms may be expressed in

various ways, as follows:

1. The number of calories necessary to warm a column of

water of unit base in the deepest part of the lake from the se¬

lected minimum (0°, 4°, winter minimum) to the summer tem¬

perature. This is the method followed by Forel.

2. The total sum of the calories necessary to warm in a simi¬

lar way the whole mass of the water of the lake from the selected

minimum to the summer temperature. This is the method em¬

ployed by Halbfass in his paper of 1910.

3. The total number of calories necessary to warm in a sim¬

ilar way a column of water of unit base and a height equal to

the mean depth of the lake. This method is used in this paper.

The first method was that employed by Forel ( ’95, p. 400, and

’01, p. 41). It is well suited to very deep and very large

bodies of water in which the annual temperature changes are

confined to the surface strata, and whose area is so great that

we may, without appreciable error, neglect the difference be¬

tween the volume of the upper part and the lower part of the

stratum in which these temperature changes take place. Such

bodies of water are oceans and seas, and possibly also the very

largest fresh water lakes. In the ordinary inland lake, how¬

ever, neither of these conditions is met, and the method of stat¬

ing the heat budget is correspondingly defective. It is also

true, as stated by Forel, that it is not possible by this method

to compare the heat budget of different lakes.

Forel at first took as his data the temperature found at the

surface and at the successive 10 m. levels of' the lake and added

them. Since a column of water 1 cm. square and 10 m. high

Birge — Heat Budgets of American and European Lakes. 171

contains 1 liter, this sum gives the number of large calories

necessary to raise a column of water, whose base is 1 sq. cm.,

and whose height is the maximum depth of the lake, from 0°

to the temperature at the time of observation. This method was

later modified by employing the mean temperature of each 10 in.

section of the column of water instead of the temperature of the

ends of the section. The unit of 'area was altered to the sq.

dm., thus multiplying the number of calories in the result by

100 and escaping the necessity of using fractions of a calorie.

This method permits a study of the gain and loss of heat in

any one lake, but it does not permit a comparison of different

lakes.

If the method is applied to the New York lakes, the following

results will be reached; 'which should be compared with those

of Table A.

TABLE 1

ANNUAL HEAT BUDGETS OF NEW YORK LAKES, COMPUTED BY FOREL’S

METHOD

The method can also be applied to measure the gains of heat

above 4°, and the result for 1910 will be as follows:

TABLE 2

SUMMER HEAT-INCOME OF NEW YORK LAKES, COMPUTED BY FOREL’S

METHOD

* This Wisconsin lake is smaller than the New York lakes, but of the same type. It

is constantly used in comparison with them in Birge and Juday ’14.

172 ! Wisconsin Academy of Sciences, Arts, and Letters.

In these tables I have stated the results in small calories pel*

sq. cm. of surface, in order to make them comparable with those

given elsewhere in this paper. To agree with Forel’s notation

they should be divided by 10.

The figures are much higher for the deeper lakes than are

those given by the other methods. They are correct so far

as they go, but they may easily deceive. It is doubtless true- —

as the defenders of Forel’s method replied to Halbfass — that

these figures state correctly the amount of heat necessary to

warm the given column of water in accordance with the observa¬

tion. They do not show, however, how much of this heat came

direct from sun and sky, and how much came in laterally from

columns of water in the lake which are shorter, and therefore

contain less heat. In other words, if the result for the column

is carried over to the lake, the successive levels of the lake are

treated as if each contained the same volume, and as a result

the figures given in Table 1 show a much greater gain of heat

than could possibly come directly from sun and sky, or from any

other source outside of the lake. It is not probable that more

than 70,000 gr. cal. per sq. cm. could be thus received by the

surface of a lake during the five months, April to August, in¬

clusive. It is quite beyond possibility that so many as 62,000 of

these could be stored up in a lake. Thus, while a record of this

sort may have a certain value for comparing results within a

single lake at different times or in different years, it has little

value for such a purpose as that for which it was used by Forel,

viz., for comparing the relative gains of heat in lakes which lie

in different latitudes.

In Table 2 the gains shown for the shallower lakes, Canadice

and Otisco, are much lower, relatively to the deeper lakes, than

would be shown by a comparison based on mean depth rather

than maximum (see Birge and Juday ’14). This is merely an¬

other illustration of the fact that different lakes can not be com¬

pared by this method. It deals with columns of water, not

with lakes, and results reached by it can not be applied to lakes.

Not only may this method make the results larger than can

be true for the lake, but the amounts of heat shown for the

same lake in different years may differ more widely than they

should do. In Seneca lake, for instance, the budget for 1910,

as shown in Table 1, is nearly 24% greater than in 1911. This

Birge — Heat Budgets of American and European Lakes . 173

is true for the columns of water whose gains are shown, and in

which the rise of temperature of a 10 m. section of the deep

water involves as much heat as that of the water near the sur¬

face. Since, however, the volume of the lower strata of the lake

is smaller than that of the upper, the amount of heat necessary

to warm a given stratum of the deeper water of the lake is less

than that needed to warm a corresponding stratum of the up¬

per water, and when the volumes of the several strata are con¬

sidered, the budget of 1910 was less than 11% greater than that

of 1911, as is shown in Table A.

Forel’s method, therefore, states the losses and gains of heat

in a way which, while true for the column in which the temper¬

atures were measured, may be obviously impossible when applied

to the whole lake. Thus he gives for Ladoga (’01, p. 46) be¬

tween certain dates a mean daily gain of 101 kg. cal. per sq. dm.

and in a similar way for lake Enare a gain of 163 kg. cal. pei^

day. These sums are equal to 1010 and 1630 gr. cal. per sq. cm.,

respectively. This amount of heat is far greater than can pos¬

sibly be furnished to a lake by sun and sky. They are, as Bruck¬

ner states “vollkommen unverstandlich. ” The sun would very

rarely deliver so much as 600 gr. cal. per sq. cm. per day for a

month on a horizontal surface, and 1000 cal. are impossible.

Thus it is plain that much of the apparent gain of heat in the

column of water observed has come from other sources than sun

and sky. As a matter of fact, it has been contributed by other

parts of the lake whose column of water is shorter and whose

gains are correspondingly smaller. If the gains of Ladoga are

computed on the basis of the mean temperature and mean depth

of the lake, they amount during the period named to about 160

gr. cal. per sq. cm. per day, instead of more than 1000 cal. The

smaller figure is very moderate and wholly intelligible. The

gains of lake Enare can not be thus computed since the mean

depth of the lake is not known ; but they are probably no greater

than those of Ladoga and can not equal one-tenth of the amount

computed by Forel. Obviously, the results of computations

based on the maximum depth of lakes afford no basis for estab¬

lishing laws concerning the relation of heat budget and latitude.

Any conclusion on such a subject must be based on a knowl¬

edge of the heat gains of the average of the lake and not on those

of a selected column of water.

174 Wisconsin Academy of Sciences , Arts, and Letters.

Bruckner (’09, p. 305) thinks that in spite of the objections

of Wojeikoff and Halbfass, Forel’s units are the best for the

study of the physics of lakes, since they give exactly the facts

of temperature for that part of the lake which covers the cen¬

tral plain (“der sogenannte Sehweb”). We do not know ex¬

actly the temperature of the shallow parts of the lake, nor do we

know exactly its volume, so that we cannot apply the needed cor¬

rections to this temperature. These considerations have a cer¬

tain value, but, as already indicated, they are not of very much

weight. The statement that Forel’s units give a better idea of

the physics of the lake may hold perhaps against Halbfass ’ meth¬

od, but not against that of Wojeikoff, or that used in this paper.

Indeed, as has already been indicated, if the student desires to

correlate the heat cycle of the lake with that of the season, units

of surface, depth, and temperature must be employed which will

state the facts for the entire lake and not merely for a part of

it.

The second method was introduced by Halbfass and was first

applied by him in the papers to which reference has already

been made. It has the fundamental advantage over Forel’s

method of basing itself on the mean temperature of the entire

water of the lake and not upon that of part of it. But the use

of the entire volume of the lake as a factor for computing total

gains and losses of heat destroys for comparative purposes a

large part of the advantage gained from the use of the mean

depth in ascertaining the mean temperature of the water. This

method employs a factor which varies with the individual lake,

and thus restores in the result a difficulty similar to that which

Forel’s method suffers from his use of the maximum depth in¬

stead of the mean depth. A method by which the daily gains of

loch Garry amount to 7 cal. ; of loch Ness to 140 ; lake Geneva,

1430; and Wettern, 3400 cal. can give no really comparable re¬

sults. Halbfass finds that in comparing lakes, he must select

those of the same volume, and these he can roughly compare.

This is true also under Forel ’s method if confined to lakes whose

maximum depth is similar and in which the ratio JIm. is nearly

Dmx

the same. But in both these methods the opportunity for com¬

parison is greatly restricted.

Halbfass states gains and losses of heat in calories per day

during the interval between the observations. This method re-

Birge — Heat Budgets of American and European Lakes. 175

duces the number of calories to a comprehensible sum, but the

expression is little more than a mathematical result under the

circumstances to which it is applied. If temperatures had been

tkken daily, or even weekly, during the warming period of these

lakes, it might be worth while to compute daily gains. But the

case is far different. We must depend on a few — perhaps two —

isolated observations in each year, one for winter and a second

one for summer. Under such conditions a statement of “mean

daily gains” has little meaning. There is really no significance

in a ‘'mean value” for the daily gain between (say) Feb. 5 and

Sept. 7. The first month of this period may represent a daily

loss ; then may come daily gains increasing in amount and reach¬

ing a maximum in April or early May ; then a slowing of gain

leading to an almost stationary condition by late July and Au¬

gust, and passing probably into small losses in the later part of

the period. Under such conditions, a statement of the mean

daily gain or loss means little or nothing. A good illustration

may be taken from lago di Como in 1904 (Halbfass, ’10, p. 62).

The winter temperatures were as follows: Jan. 26, 7.11° ; Feb.

26, 7 :00° ; Mar. 28, 7.15°. The maximum recorded temperature

of 8.96° was noted on Aug. 27. If the winter temperature in

January only had been recorded, the mean daily gain would have

been stated as about 160 cal. per day; if February had been

taken, about 200 cal. ; if March, about 220 cal. As a matter of

fact, the temperature was substantially stationary during two

months of the winter, and the mean daily gains stated for the

season will vary nearly 40%, according to the winter date se¬

lected as the point of departure. The annual heat budget, how¬

ever, is much the same whichever winter date is chosen.

The third method was first proposed, so far as I know, by

Wojeikoff (’02, pp. 193-199) in a discussion of Forel’s results.

The same method was suggested by Wedderburn* (’10, p. 134).

This method could not be applied widely until such a compila¬

tion of lake temperatures had been made as Halbfass’ papers

have furnished, and neither of the authors named has made any

serious attempt to apply it.

The Wisconsin Geological and Natural History Survey has

used this method in all of its work on Wisconsin lakes, which

* It may be noted that the figures given for lake Geneva by Wedder-

burn (76,000 gr. cal. per sq. cm.) are certainly much too high. Loch

Ness is assigned 34,000 cal., which is lower than my computations.

176 Wisconsin Academy of Sciences , Arts, and Letters.

has been going on since 1898. It is the only method applicable

under our circumstances, which involve a comparison between

the heat budgets of lakes differing widely in form and area, but

similarly situated as regards topographic and climatic condi¬

tions. It allows us to determine the influence of such factors

as the area and depth of lakes on the amount of heat taken in

by them. This unit, therefore, is employed throughout this

paper.

If the heat budgets of lakes are to be widely studied, units

must be selected, both for temperature and area, such that lakes

may be compared in spite of great differences in area and depth.

We have, therefore, computed the mean temperature of a lake

on the basis of its mean depth. We derive the number of gram

calories per square centimeter above zero, represented in this

temperature, by multiplying it by the mean depth expressed in

centimeters. We employ the gram calorie as the unit of heat

and the square centimeter as the unit of area, because these units

are employed by the meteorologist in stating the amount of heat

received by the earth’s surface from the sun.

C. The third preliminary question concerning the character of

the observations from which our conclusions regarding lake tem¬

peratures are derived. The results shown in Table A are based

on series of temperatures taken at the deepest part of the lake

concerned. There is no difficulty in ascertaining with sufficient

accuracy the mean temperature of the column of water in which

such observations were taken. Nor is it difficult to compute

from such observations the mean temperature oi the water of

the lake, provided a hydrographic survey of the lake has been

made. But does such an expression give a correct idea of the

mean temperature of the water? On this matter Wedderburn

states (’07, p. 408) : “It is in a far greater degree impossible

to deduce the average temperature of a large loch from observa¬

tions made in any one place.” Halbfass also speaks (’13, p. 471)

of “die Bedeutingslosigkeit einer Beobachtungsserie in verti-

kaler Richtung in einem vereinzeltem Punkt eines Sees”.

If these statements are correct, then plainly such a paper as

this, or those of Halbfass, are of no value, since they are based

on just such observations as Wedderburn calls “futile” in the

paper quoted, and which Halbfass thinks are “bedeutungslos”

These are practically the only kind of observations now published

for any lake. So far^as I am aware, there is no lake whose tern-

Birge — Heat Budgets of American and European Lakes. 177

perature at all depths has been followed from day to day for a

year, still less for a series of years. The Wisconsin Survey has

done this for lake Mendota, but the results are still unpublished.

The same is also to be said of the observations made by the sta¬

tion at Lunz. There is no lake the mean temperature of whose

water is known in the same way as the mean temperature of the

air is known for thousands of places all over the world.

In the paper on the Finger lakes, reasons are presented for

believing that a single series of observations taken near the

middle of a lake gives a fair account of the mean temperature of

its water at the time of observation. (Birge and Juday T4,

p. 556.) I shall not repeat these arguments at this place, but

will add somewhat to them, mainly from the facts contained in

Table A.

The mean temperatures of Green lake in nine seasons on dates

varying from August 14 to September 8 show a total range of

less than 1°, and a maximum range of about 12% in the wind-

distributed heat. In those years in which observations were

made on different dates, that one was selected which shows the

highest result. I can not believe that so close an agreement

would be present if the figures did not fairly show the mean

temperature of the water. Certainly if the temperature of the

water in different parts of the lake varies greatly, such a dif¬

ference ought to cause a more considerable variation in results.

If such observations are “ futile”, the futility ought to show

itself by large variations in the temperatures indicated.

In the paper on the New York lakes, attention was directed

to the close agreement between the mean temperature of lakes

as deduced from observations made at the center and that de¬

rived from the mean temperature of several series of observa¬

tions made along the axis of the lake. A maximum variation in

the New York and Wisconsin lakes was found not exceeding 5%.

The same result is reached if similar observations on European

lakes are compared. Wedderbum (’07) gives several instances

where temperatures in loch Ness were taken at Inverfarigaig at

the center of the lake and at the same time near Ft. Augustus

and Dores at its ends. The mean temperature of the lake, de¬

rived from the middle series and that derived from the three

series, do not often differ by more than 0.1° C. This difference

of temperature would correspond to a difference of about 1300

cal. in a budget of about 40,000 cal. Such a difference of about

12— S. A.

178 'Wisconsin Academy of Sciences, Arts, and Letters.

3% is very far from the exactness expected in a laboratory ex¬

periment; but in the present state of our knowledge of lakes,

a possible variation of 3% or 5%, or even a larger error, does not

render an observation “ futile’’ for purposes of general discus¬

sion.

The same general results as to the relation of observations at

the center of the lake and those made simultaneously at the

center and ends can be derived from the observations made by

Wedderburn and Halbfass (Halbfass 10a) on the Madii-See in

Pomerania.

In short, so far as I can learn from my own observations and

from those of others, the statement made in the paper on the

Finger lakes is correct: “It is fair to conclude that the mean

temperature of the water of a lake of simple form may be de¬

rived from a single series of observations taken at or near the

center of oscillation of the water.” (Birge and Juday T4, p.

558.)

I am confident that this statement is true for American lakes

in our latitudes up to 10 km. — 15 km. of length. My experience

of larger lakes is limited, but I believe that it holds true for

them also. How far it is true for European lakes must be

shown by observations made there. I am prepared to accept

any conclusion which such studies may warrant, but I shall be

surprised if observations made at the center of a lake are shown

to be in general either “futile” or “ bedeutungslos ” as evidence

of the mean temperature of the water of the lake.

I believe, therefore, that the numbers stated in Table A rep¬

resent with general truthfulness the heat budgets of their re¬

spective lakes. The results obviously include several variable

features, and they are, therefore, approximate and provisional.

Yet they come from so many places and so many different years

that they constitute a “random selection” of the facts. They

so far agree with each other that certain general conclusions

may be drawn which further study will doubtless modify, but

which are not likely to be overthrown.

Birge — Heat Budgets of American and European Lakes . 179

The results of Table A may be summarized as follows :

TABLE 3

ANNUAL HEAT BUDGETS OF EUROPEAN AND AMERICAN LAKES

This table discloses the following facts:

1. The European heat budgets are on the whole smaller than

the American. More than one-third of the whole number are

below any American budget. Green lake (see Table A) has a

larger average budget than any European lake except three of

the giants — Geneva, Ness, and Mjosen. 0 wasco lake exceeds

Green and is only slightly behind the other three. This is espe¬

cially significant when we note that these two American lakes

are shallower, both as regards maximum and mean depth, than

any European lake recorded in Table A. and that their area is

much below the average. Only five of the twenty-live European

lakes are decidedly smaller than Owasco. Yet only three lakes

besides those already named — lago di Como, Traun-See, Zuger

See — have even single budgets that exceed Owasco ’s largest, or

indeed, its mean budget.

It must also be noted that the small average size of the Euro¬

pean budgets is due in great measure to the numerous mid-

European lakes, whose budgets are in general very small. If

only those lakes were considered which lie south of the Alps or

north of latitude 55°, the difference in favor of America would

be much less.

The larger amount of heat in the American lakes is due chiefly

180 'Wisconsin Academy of Sciences , Arts, and Letters.

to their warmer epilimnion, as is shown elsewhere by a com¬

parison of Cayuga lake and Wiirm-See.

There are also some very large budgets in the European se¬

ries. Seven of them from five lakes exceed 40,000 cal., while

only one American budget exceeds this sum, and that but slight¬

ly. In the paper on the Finger lakes (Birge and Juday, ?14

p. 565) reasons were given for believing that the budget of

Skaneateles lake exceeded 45,000 cal. in 1912, and no doubt this

figure is reached and exceeded by other lakes, and a series cov¬

ering a greater number of years would contain such budgets.

It is plain that budgets close to 50,000 cal. may be expected

occasionally in both continents, but this seems to be near the

possible maximum. In case of budgets that reach or exceed this

figure, the data on which they are based should be very carefully

scrutinized.

2. The range of the European budgets is much greater than .

that of the American. This might be expected, since the

European lakes are more numerous than the American, they ard

distributed over a much wider area, and they lie in very differ¬

ent topographic and climatic situations. On the other hand, it

is to be noted (a) that Green lake lies about 1000 km. west of

the New York lakes and yet closely agrees with them; (b) that

the variations in European lakes seem measurably independent

of situation — so far as concerns single lakes — and (c) that

single European lakes may show variations that cover nearly the

whole range of the series. Lago di Como, with eleven budgets,

has a range from 17,000 cal. to nearly 42,000 cal. and appears in

five of the six classes of Table 3. Lake Geneva, with six bud¬

gets, appears in five classes and ranges from 22,000 cal. to nearly

46,000 cal. Zuger See, with four budgets, appears in four

classes and ranges from 18,000 cal. to 44,000 cal. Certain lakes,

like Traun-See, Mjosen, and perhaps lac du Bourget, have one

budget much exceeding the others. Reasons are given elsewhere

for believing that the great apparent size of Mjosen ’s budget of

1901 is at least partially deceptive ; and very possibly a longer

series of budgets would show a considerable range of variation

in the other lakes, and fill in the gap between the small budgets

and the large one. This is true of lac du Bourget, but not of

Birge — Heat Budgets of American and European Lakes. 181

Traun-See, where the large budget of 1895 is in striking con¬

trast with the ten others , and must be regarded as exceptional.

On the other hand, certain lakes with several budgets seem quite

as regular as the American. Such are loch Ness, lago di Bol-

sena, Zurieher See, Boden-See, and probably Wiirm-See. This

regularity, where it exists, is the more noteworthy since the years

from which budgets are reported cover a much wider range than

is the case with American lakes. i

The range of European budgets is so large that there is no use

in calculating their mean. That of the American lakes is 36,000

cal., and the mean departure of each observation is less than

1,800 cal., or 5%. The maximum departure is +5,900 and

— 3,100, or a range of 25%. So far as the evidence goes, there¬

fore, the heat budget of American lakes of the first class lying in

the region of lat. 43° is in general much higher and more uni¬

form than that of European lakes of similar character.

The amount of the summer heat-income in the temperate

lakes is shown in the following table:

TABLE 4

SUMMER HEAT-INCOME OF EUROPEAN AND AMERICAN LAKES

In this table the same facts are evident as in Table 3. More

than 20% of the European budgets lie below any American ; over

40% more lie in the class which contains less than 20% of the

182 | Wisconsin Academy of Sciences, Arts , and Letters.

American budgets; while 80% of the American budgets are in

the upper region that contains only 40% of the European. It

must be noted further that eleven budgets, or more than half

of those found in Europe which exceed 25,000 cal., are from one

lake — Traun-See, which is obviously exceptional in this partic¬

ular. Some of the other exceptionally high budgets are dis¬

cussed in the notes to Table A.

The uniformity of the American budgets is as noteworthy as

their size. 80% of them lie between 25,000 and 30,000 cal. I do

not suppose that a larger collection would show the same uni¬

formity, but I believe that, on the whole, the American lakes will

be closer to each other than are the European.

ForeFs theory ( ’01) that the heat budget of northern lakes is

greater than that of lakes farther south cannot be proved or

disproved by the records of the European lakes, or by these with

the addition of the American lakes. The budgets that exceed

40,000 cal. are as follows:

According to Halbfass’ figures, loch Ness would have a bud¬

get of 43,000 cal. Of the American lakes, Skaneateles has one

budget of 41,900 cal. and reason is given in our paper for be¬

lieving that in this lake budgets as great as 45,000 cal. may be

found. The same is doubtless true of other deep lakes of the

Finger lake series. It seems certain that the budget of Mjosen

is larger than the facts warrant. At any rate, these budgets

show no decided preference for northern lakes and do not confirm

Forel ’s theory. On the other hand, the. average budgets of loch

Ness and Mjosen (omitting the exceptional one) are somewhat,

though but little, greater than those of Geneva, and possibly

the average of many years is as great as 40,000 cal. Very prob¬

ably a longer series would confirm this relation of superiority on

the part of the northern lakes. But there is no good reason to

attribute this difference to the northern latitude. There is more

Birge — Heat Budgets of American and European Lakes. 183

difference between Como and Geneva, separated by only 0° 21'

of latitude, than between Geneva and Mjosen, which lies over 14°

further north. In view of the fact that Wettern and Ladoga

show rather small annual heat budgets and that lochs Katrine,

Lochy, and Morar, near neighbors of loch Ness, have budgets of

no extraordinary size, it may well be questioned whether the

large average budgets of Ness and Mjosen are not due to other

causes than their northern situation.

So far as the data now before us can warrant a conclusion,

it would be that the variations in the annual heat budgets of

these lakes lie within the range due to the variations of the

weather of the season. Thus lakes, whose latitude ranges from

that of Mjosen (lat. 60°) to Geneva (lat. 46°) or Seneca (lat.

42°) may have identical budgets. We can not say that the mean

budgets of the several lakes will not differ; indeed we may be

sure that they will not be identical. But there is no reason to

think that those of the northern lakes will have any great su¬

periority or that any difference found will be due to the latitude.

In a word, there is no evidence that the annual heat budget in¬

creases with latitude, within the limits of the zone between 40°

and 60° north. Still further, the data from these lakes do not

show that a temperate lake has a larger heat budget than a trop¬

ical lake of comparable area and depth.

The lakes of central Europe show on the whole smaller heat

budgets than would be expected from American experience. The

high winter temperature of these lakes which regularly freeze

is noticeable, and this reduces the amount of that part of the

annual heat budget which lies below 4°. Something of the same

sort appears in the upper part of the budget also. While in the

American lakes of the first class the wind-distributed heat will

ordinarily exceed 25,000 cal., a decidedly lower figure would nat¬

urally be selected for those of central Europe.

An instructive comparison may be made between lake Cajmga

and Wurm-See which have closely similar mean and maximum

depth. The comparison of their heat volumes shows that the

main difference between these lakes lies in the upper 10 m. or

20 m. (Figs. 2-3). This region in Cayuga lake, down to 15 m.,

has a temperature substantially uniform and high, while in

184 Wisconsin Academy of Sciences, Arts, and Letters.

Warm-See the temperature begins to decline almost from the

surface. The amount of wind-distributed heat in the lower

strata of Wiirm-See is as great as in those of Cayuga. But the

European lake seems to lack the continuous sunshine, heat, and

light winds of mid-summer which give the American lakes the

peculiar character of their August temperature curves in the

upper strata. This difference is apparently the characteristic

one found when the temperatures of mid-European lakes are

compared with those of lakes in our latitudes of America, and

this is the usual cause of the lower heat budget, where such ex¬

ists.

The August temperature conditions in the lakes of central

Europe, in general, resemble those of American lakes in June.

In the European lakes a relatively thick epilimnion is not formed

in early and mid-summer as in ours, and a well-marked

epilimnion is hardly developed in these lakes until the surface

begins to cool. Prom this fact comes the statement, not uncom¬

mon in European writers on lakes, that the thermocline is a phe¬

nomenon which develops during the cooling period of a lake.

No student of American lakes would make such a statement,

since the epilimnion is fully formed in July, even in a large

lake, while in smaller lakes it may be well developed in June or

even in May.

In Fig. 1 the number of gram calories in the annual heat bud¬

get of each lake is indicated by the length of the line assigned

to it. The position of the line on the figure indicates the char¬

acter of the lake as tropical or temperate, and its position in the

class to which it belongs. The line marked “0” indicates the

temperature of 4°. In the case of the temperate lakes the lake

lines cross the zero line and the length of that part of the line

to the left of zero indicates the number of calories per square

centimeter which its water loses below 4° ; or in other words the

number of calories per square centimeter of its surface required

to raise its water from the winter temperature to 4°. That part

Of the line to the right of the zero line indicates the number of

calories per square centimeter necessary to raise its water from

4° to the summer 'temperature, or the summer heat-income. In

the case of the tropical lakes the left end of the line is placed

Fig. 1. This diagram shou

of the lake. Each division of til

The place of the line on the dias

get of a tropical lake is shown ;

same line; that of a temperate ]

The broken line extending 1

ing winter. No winter observat

eluding in the budgets of that b

In general, where there ar<

recorded, the mean is taken for

n to 5

Cayuga, |

Owascol

SHaneatjjles

Green

Bolsenaj

Orte. J

Como J

Geneva j

Thun |

Zua !

Zurich |

Mil I shaft

Bourftefj

Annecy

Traun

Tefiei

Ness

Morar

Lochy

Katrine J

Wetterrj

Ladoga

40 45 TO 55 60 65 70 75 80 65 90

-H—

I i i I • !

of th^* 1' -J11 * **8 dia?r.am shows the annual heat budgets of American and European lakes, expressed in gram calories per square centimeter of the surface

Thpnin % Each division of the diagram indicates 5,000 cal. per sq. cm. The length of the line for any lake indicates the amount of its annual heat budget.

Btifi .t the line on the diagram indicates the position of the lake as tropical or temperate. The zero line is drawn at the temperature of 4°. The bud-

s tr°Plcai lake is shown by a line wholly to the right of the zero line. That of a polar lake, if one were present, would lie wholly to the left of the

-‘Me lme; that of a temperate lake crosses the line.

ine win*! htoken line extending the budget lines of Millstatter See and Ammer-See to the left indicates the supposed loss of heat below 4° in these lakes dur-

eintnncTi rVv ™nter observations have been made on them. The broken line extending the budget line of Miosen to the right shows the effect of in-

dueling in the budgets of that Jake the budget of 1901. See p. 43.

remraTaStuera ’ whflre tllere are more than two budgets for a lake, the mean is taken. In those tropical lakes for which several winter temperatures are

eoraed, the mean is taken for the starting point of the lake line.

Birge — Heat Budgets of American and European Lakes. 185

at the point indicating the number of calories necessary to raise

its water from 4° to the winter temperature. It represents in

some sense the permanent stock of heat in the lake above 4°.

The diagram shows, for instance, that Cayuga lake is a tem¬

perate lake; that its annual heat budget is about 37,000 gram

calories per square centimeter of the surface of the lake; that

of these calories more than 9,000 lie below 4° ; that about 28,000

cal. constitute the summer heat-income. It shows that lago di

Bolsena is a tropical lake; that at its winter temperature the

water still contains wind-distributed heat to the amount of about

29,000 gram calories per square centimeter of its surface; and

that its annual heat budget is about 32,000 calories.

The temperature of 4° is taken as the starting point for four

reasons :

1. The lakes all approach within a measurable distance of 4°

in the autumn or winter of each year.

2. If 0° C. were taken as the starting point the distances

from zero line to beginning of lake line would vary greatly. In

Owasco lake in 1911 only about 2,400 cal. would remain in the

lake ; in Mjosen some 60,000 cal. ; in Seneca nearly 64,000 cal. ;

in Como 126,000 cal. Yet Mjosen had lost below 4° some 15,000

cal. per sq. cm. and Owasco only about 6,200, and it is well worth

while to bring out this fact.

3. Losses of heat below 4° and gains of heat up that point are

subject to very different laws from those at a higher temperature

and should be treated separately. '

4. The principal reason, however, for placing the zero line

at the temperature of 4° is the fact that in this way the place of

the lake in Forel ’s classification can be shown in the diagram, as

well as its position in the class to which it belongs.

The diagram shows the main facts of Table A in a way which

conspicuously strikes the eye. Among th.em I should name:

1. The small heat budget of a series of mid-European lakes

from Thuner See to Traun-See, and their general' uniformity in

spite of differences of size, depth, situation, etc. The differences

(with two possible exceptions — the largest of Zuger See and

Traun-See) all seem to be within the possible range of annual

variation of any one of the lakes.

186 Wisconsin Academy of Sciences, Arts, and Letters.

2. The small loss of heat below 4° in these lakes as compared

with north European or American lakes. In the case of Mill-

statter See and Ammer See, where no winter temperatures are

recorded, this item is conjectural and is so indicated by the use

of dotted lines.

8. The uniformity oF the American lakes, both as to losses be¬

low 4° and gains above it. The diagram also brings out a pos¬

sible error in the observations on Seneca lake. We can give no

reason why that lake, which does not freeze, should not have

lost as many calories per sq. cm. in 1911 as the other lakes.

Cayuga lake, which also does not usually freeze, shows substan¬

tially the same losses as Owasco and Skaneateles, whose temper¬

atures were taken through the ice. Yery likely, therefore, the

water at the point of observation in Seneca lake showed a tem¬

perature somewhat above the mean of the lake. It is quite pos¬

sible, however, that the observation correctly shows the facts.

4. The three northern lakes of the continent of Europe show

a budget of similar character in 'spite of great differences in area

and depth. In each case there is a very large loss below 4°, so

large as to affect the possible gains above 4°. The gains of

Ladoga and Wettern seem to be closely similar, although more

evidence is needed on this point, either to confirm or refute it.

5. The very large and deep lakes, Mjosen, Ness, Geneva, and

Como, have on the whole decidedly larger budgets than the lakes

which are their neighbors. This inference is weakened by the

small budget found for Thuner See in 1908-09, but this 'may be

due to local reasons, or a larger series of observations may show

that its great mean depth — 50% greater than any of its neigh¬

bors — gives it a certain advantage over them, when the average

of a considerable number of series is taken.

6. The large budgets of lago di Bolsena show (a the favor¬

ing effect of area on budget; (b) a high winter temperature

is not incompatible with large gains of heat.

7. The fact that the Scottish lakes are tropical in spite of

their high latitude is a testimony to the general mildness of the

winter. It is surprising that summer gains of heat are so large.

Continuous sunshine, however, is by no means especially favor¬

able for large gains of heat, as it warms the surface too rapidly.

Birge — Seat Budgets of American and European Lakes . 187

8. No clear relation can be discovered between latitude and

size of heat budget.

9. More important than any of these specific results is the

conclusion that, by the method employed, it is possible to ex¬

press in similar units and to compare the heat budgets of lakes

which differ widely in area, depth, etc. ; that, even from the im¬

perfect temperature records now available, important generali¬

zations can be made as to the heat cycle of these lakes ; and that

it is plainthatmoreabundant data will enable us to correct and

to extend these generalizations.

Figure 2 and figure 3 are diagrams showing tne quantity of

heat in the several 10 m. strata of Cayuga lake and Wiirm-See.

Each of the larger divisions of the network represents 2° hori¬

zontally and 5 m. vertically. The horizontal lines marked 10,

20, etc. are so placed as to show the ‘‘reduced thickness’ ’ of the

several 10-m. strata ; that is, the thickness of each 10-m. stratum

if its area were extended to that of the surface of the lake and

if its sides were vertical. This thickness is computed by divid¬

ing the volume of the stratum by the area of the surface of the

lake.

The area in the diagram, bounded by the lines representing

winter and summer temperature and by any two of the 10-m.

boundary lines is proportional to the amount of heat delivered

to that stratum from the surface of the lake. Each of the

larger squares included in such a quadrilateral represents 1000

gr. cal. per sq. cm. of the surface of the lake delivered

to the stratum and each small square represents 40 such

calories. The area bounded by any two of the 10-m. lines, the

temperature line of 4°, and the temperature curve gives the sum¬

mer heat-income of the stratum in question.

On the dates selected for summer temperatures the tempera¬

ture of the two lakes at the surface was almost identical. In

Cayuga lake the temperature of the epilimnion falls very slowly

and has declined less than a degree at 15 m. In Wiirm-See the

temperature declines rapidly below 5 m. and at 15 m. is more

than 10° below that of Cayuga. This lake was also warmer at

depths immediately below 15 m., the difference falling to 0.4°

at 40 m. At 50 m., 60 m., 70 m., and 80 m. Wiirm-See was

188 -Wisconsin Academy of Sciences , Arts , and Letters.

Fig. 2. — Diagram to show the heat income of Cayuga lake. The date of the

winter temperature was Feb. 13, 1911 ; those of the summer temperatures were

Aug. 11-12, 1910 (full line) and Sept. 2, 1911 (dotted line).

warmer, and at 100 m. and below the lakes were of the same

temperature. The heat budget of the lakes differs widely, being

about 44% greater (11,400 cal.) in Cayuga lake than in Wiirm-

See. But little of this difference comes in the upper 10 m.

where the greater reduced thickness of Wurm-See compensates

for its lower temperature. Nearly 70% of the difference comes

in the strata between 10 m. and 40 m., where the temperature

of Wurm-See is much below that of Cayuga; and a great part

of the remainder comes in the bottom strata whose volume is

much smaller in Wurm-See, corresponding to its smaller max¬

imum depth.

Birge — Heat Budgets of American and European Lakes. 189

A large part of the difference between the budgets is due to

the higher winter temperature of Wurm-See, 3.30° as compared

with 2.23° in Cayuga. This difference on a mean depth of 54 m.

would make the annual heat budget of Cayuga about 5,800 cal.

larger (more than one-half the difference in the budgets) than

that of Wurm-See. In the wind-distributed heat therefore,

the two lakes differ much less than in the annual heat budget.

The summer heat-income of Cayuga lake is about 5,600 cal.

larger than that of Wiirm-See. Over 80% of this difference lies

190 Wisconsin Academy of Sciences, Arts, and Letters.

in the stratum between 10 m. and 20 m. and therefore in the

thermocline region. The following zones down to 40 m. more

than make up the rest, while the wind-distributed heat in W iirm-

See between 40 m. and 80 m. is greater than in Cayuga lake

at the same depths. (See Table 5.)

These facts show (1). That the agents for distributing heat

are substantially as effective in /Wurm-See as in Cayuga lake ;

and that the smaller heat budget of the European lake is not

to be attributed to its smaller size or to less efficient agents for

the distribution of heat. (2). That the gains of heat in early

spring are not essentially different in the two lakes. (3). That

the possibilities of gaining heat become less for the European

lake as the season advances. (4). That the main difference in

the temperature and in gain of heat lies in the epilimnion and

the thermocline.

COMPARISON OF CAYUGA LAKE AND WURM-SEE

Birge — Meat Budgets of American and European Lahes. 191

192 , Wisconsin Academy of Sciences , Arts , and Letters,

TABLE A— HEAT BUDGETS OF AMERICAN

Birge — Heat Budgets of American and European Lakes,

193

AND EUROPEAN LAKES.

13— S. A.

194 Wisconsin Academy of Sciences, Arts, and Letters ,

TABLE A— HEAT BUDGETS OF AMERICAN

Birgc — Heat Budgets of American and European Lakes. 195

AND EUROPEAN LAKES— Continued.

196 Wisconsin Academy of Sciences , Arts, and Letters.

TABLE A— HEAT BUDGETS OF AMERICAN

Birge—Heat Budgets of American and European Lakes. 197

AND EUROPEAN LAKES— Continued.

198 Wisconsin Academy of Sciences , Arts 9 and Letters.

TABLE A— HEAT BUDGETS OF AMERICAN

Birge — Heat Budgets of American and European Lakes. 199

AND EUROPEAN LAKES— Continued.

+ Including 01—01,

200 f Wisconsin Academy of Sciences, Arts, and Letters .

Notes on Table A.

All temperatures for American lakes have been computed by

myself ; the same is true for those European temperatures which

are marked with a star. The data for Ammer See come from

Geistbeck ’85 and Ule ’06; those for Tegern-See come from

Breu, ’06. All other data are from Halbfass’ papers of ’05 and

’10.

In the table annual gains of heat are shown by subtracting

the winter temperature from that of the summer following, and

are indicated thus in the table, ’95- ’95. Losses of heat are

shown by subtracting the winter temperature from that of the

preceding summer, and are shown thus, ’95- ’96. Both changes

in temperature are taken as the basis for computing the annual

heat budget. Thus observations in two successive summers and

the intervening winter afford a basis for stating two budgets.

This is the method followed by Halbfass in the papers to which

reference has been made. The formula is Dm (Tms-Tmw), in

which Dm represents the mean depth in centimeters; Tms, sum¬

mer mean temperature ; Tmw, winter mean temperature.

The summer heat-income is computed by subtracting 4° from

the summer mean temperature and multiplying the remainder

by the mean depth expressed in centimeters. The formula is

Dm (Tms-4).

The variations in the number of calories reported for a single

lake m the table are caused by a variety of conditions. Chief

among them is the variation' in the temperature of the water

in the different years. Exceptional seasons cause exceptional

budgets. This may be well illustrated by the temperature of

Traun-See in 1895. The winter temperature that year was only

2.66°, or nearly 1.5° below the other winter temperatures re¬

ported. The summer temperature was high, though not so far

above the average as to be obviously exceptional, but the com¬

bination caused a budget of nearly 50,000 cal., undoubtedly very

close to the possible maximum. Probably 6,000 cal. — 9,000 cal.

of this total are due to the exceptionally low winter tempera¬

ture, which is considerably below that reported for any other

European lake of approximately equal depth. A second case

may be that of Mjosen in the summer of 1901, but this will be

discussed further on.

Secondly, the temperature of the particular period in which

Birge — Heat Budgets of American and European Lakes. 201

the observations were made may cause minor variations in the

record. This period may have been cold, warm, or average, and

the temperature of the water will vary accordingly. In Wet-

tern, the mean temperature for July 11, 1900, was 7.39° (Halb-

fass, ’10, p. 62) ; Aug. 12, 6.48°; Sept. 2, 8.09°. There is thus

indicated a loss of 3,000 cal. between the first and the second

observation and a gam of 6,300 cal. between the second and third.

The middle of August normally gives a reading close to the

maximum, but on this occasion the series taken on August 12

would have given an annual budget nearly* 20% smaller than

it really was.

Thirdly, the mean temperature derived from the observations

may not fairly represent the mean temperature of the lake. This

general question has been discussed elsewhere in this paper.

There are but few cases which seem to be of this character. The

most conspicuous is that of Mjosen in 1901, which is discussed

later in these notes.

Nos. 1-7. — The mean depths of the American lakes are given

to tenths of a meter, not to indicate greater accuracy of the sur¬

vey, but that the number of calories may agree with those stated

in the paper of Birge and Juday ( T4) to which reference has

been made. This paper should be consulted for a more com¬

plete account of the heat budgets of these lakes.

In the case of Green lake, the summer temperatures are given

to the first decimal place only. This has been my usual custom

with the temperatures of Wisconsin lakes, and it did not seem

worth while to recompute the temperatures of Green lake, sincer

in the case of any lake the value of the figure in the second

decimal place is wholly uncertain. In all summers when the

lake was visited more than once, the highest reading has been

taken.

The uniformity of the heat budget is noteworthy, especially

in comparison with the irregularities of most European lakes.

The maximum departures of the annual heat budget from the

mean are about 7% in either direction; those of the summer

heat-income are less. This last fact means that there is more

difference between the loss of heat below 4° than in the gain

of heat above this point. That part of the heat budget which

lies below 4° ranged from 6200 cal. in 1911 to 10,500 cal. in

1901. It should be noted, however, that the higher temperature-

was taken in March and would be above the minimum temper-

202 Wisconsin Academy of Sciences, Arts, and Letters.

ature of the lake, which would come at the time of freezing.

Green lake seems to gain about 25 cal. per sq. cm. of surface

per day during the ice-period. The temperature at the time

of freezing might have been as much as 0.5° (1600 cal.) colder

than the record in March. But there is no doubt that the lake

might freeze at any of the temperatures indicated.

No. 8. Lago di Bolsena. — The large I heat-budgets of this south¬

ern lake are noteworthy, especially in comparison with those of

the lakes north I of the Alps. It will be observed that the low

temperature record of 1900 can not be used as basis for a bud¬

get. It no doubt indicates a very small heat budget for that

year. A longer series from Bolsena should show variations like

those of Como and Geneva.

No. 9. Lago cl ’Orta. — No definite conclusion can be based on

the single i budget from this lake, although it is probable that

Como, like Geneva, has a larger budget than its smaller and

shallower neighbors.

No. 10. Lac du Bourget. — Very probably the temperature of

March 4, 1894 is above the minimum for that year, and probably

the temperature in Feb. 1895 was below the minimum of the

preceding year. This is clearly the case in the neighboring lac

d ’Annecy. This lake and lac d ’Annecy lie near the southwest¬

ern end of lake Geneva. They evidently are lakes (which are

near the dividing line between Forel’s classes of tropical land

temperate lakes.

No. 11. Lac d ’Annecy. — This lake was measured from Dele-

becque’s large map land its volume was computed from my

measurements. The result for total volume was substantially

the same as that given by Delebecque. The maximum I depth

should be stated as 64.7 m. since this is the maximum depth in

the central plain of the lake. The depth 81.6 m. is found in the

Boubioz, a large sub-aqueous spring, but so small that it has

no appreciable effect on the total volume of the Hake. From

these measurements of volume, I have computed the tempera¬

ture. The result are higher for the years 1890 and 1891 than

are those of Halbfass. They are substantially the same in 1893.

No. 12. Lago di Como. — The mean temperatures given by

Halbfass in his two papers differ considerably. I have used the

later figures, since they have probably been revised and corrected.

These changes throw some doubt Ion the temperature for the

summer of 1898, which is given only in the paper of 11905, and

Birge — Heat Budgets of American and European Lakes. 203

which is lower than in l any other year except one. The changes

made in the figures for 1894 and 1895 alter these from i the small¬

est budgets to the largest. The excess is due to the exception¬

ally low winter temperature for 1895, which is nearly 1.0°

(18,500 cal.) i below the mean of the other five years recorded.

I have not included in the record the figures given in TIalbfass,

paper of 1905 I for the years 1887 and 1889, since these should

probably be increased like the numbers for later years. If the

budget ’98-’99 is omitted, the mean of the total number record¬

ed will be increased 1500 cal. It will probably not be far wrong

if (the mean for Como is reckoned at 32,000 cal. — 33,000 cal., and,

therefore, very close to that recorded for lago di Bolsena.

The exceptionally small budget ’98- ’99 (depends on the com¬

bination of an unusually low summer temperature, and an un¬

usually high (winter temperature. The difference as measured

in degrees is not very great, but since the mean depth of the

lake is nearly 200 meters, the difference in the heat budget is

very large.

No. 13. Lake Geneva.— -For (this lake, north of the Alps, we

have a considerable number of budgets as we have for lake Como

south of 'the Alps. The maximum budget is above that of Como ;

the minimum is higher than Como’s minimum, and the average

is also considerably greater. These lakes lie in substantially the

same latitude. Lake Geneva is the larger which would tend

to increase its (budget, but it is also the shallower, which, if this

fact has any influence in lakes so deep, would be in the opposite

direction. The main cause for the greater heat budget of lake

Geneva seems to lie in the colder winter, wdiich reduces the tem¬

perature to a point where the heat can be more rapidly absorbed

than is possible in Como.

It may be noted that the minimum budget for lake Geneva is

less than half the maximum, a ratio not very different from that

found in Como.

During the summer the Rhone carries an enormous volume of

cool water into lake Geneva and an equally large volume of

warm water is drawn off from its surface. It does not appear,

however, that this fact operates to reduce the annual heat bud¬

get of the lake. The annual heat budget of lake Geneva is

higher than that of any other lake in the table except loch Ness

and Mjosen. It is substantially the same as that for Mjosen

if the single exceptionally high budget of Mjosen is omitted.

204 Wisconsin Academy of Sciences, Arts, and Letters.

It will be noted that the winter temperature for 1891, 1892,

and 1894 are lower than any others recorded. Since the tem¬

peratures of 1892 and 1894 are in January, they are probably

above the minimum for those years. If we had summer tem¬

peratures corresponding to these winter observations the bud¬

gets wTould almost certainly have been unusually high, probably

over 40,000 cal., and would have raised the average for the lake.

Nos. 14-25. — There are twelve lakes in the table which be¬

long to the region of the Alps north of Italy and east of lake

Geneva. To these may be added lac d ’Annecy and lac du

Bourget. These lakes furnish thirty-five budgets of which seven¬

teen lie below 25,000 cal., fourteen are between 25,000 and 30,000

cal., and four are exceptionally high. Omitting the four high

budgets the mean of the remaining thirty-one would be below

25,000 cal. If all were included it would be somewhat below

27,000 cal. The Vierwaldstiitter See has been omitted from the

table on account of its complex form, and there are no annual

heat budgets recorded for Ammer See or Millstatter See, since

winter temperatures are lacking. But it is evident that were

the budgets from these three lakes added to the list the mean

would not be essentially altered.

These lakes range in mean depth from 38 m. to 135 m., al¬

though only one exceeds 90 m. The area varies more than one

hundredfold, from 822 ha. to 83,850 ha. The elevation above

the sea ranges from slightly below 400 m. to 725 m. The lakes

lie within the Alps in Switzerland, Austria, and France and on

the north side of the Alps in Germany and Austria. The gen¬

eral uniformity and small amount of the annual heat budgets

in lakes so numerous, so various, and so widely distributed is

remarkable. Equally noteworthy are the exceptionally high

budgets which appear in two of the lakes, and the large average

budgets which appear in Traun-See.

The conclusion seems warranted that lakes in the region of

the Alps, and north of Italy, have annual budgets in the region

of 25,000-27,000 calories, and that lake Geneva with its mean of

36,500 cal. is far above the average for such lakes, though not

beyond the amount which other lakes may reach in exceptional

years. <

Nos. 14-17. — These Swiss lakes lie to the east of lake Geneva.

Their -annual heat budgets are all low and are all fairly uniform

with the exception of Zuger See. This lake shows an excep-

Birge — Heat Budgets of American and European Lakes. 205

tional range in its budgets due to a considerable variation both

in the winter and in the summer temperatures. I cannot assign

any reason for this wide variation, nor can I state why its bud¬

gets should rise so much higher than those recorded for the

other lakes.

If the temperatures of Thuner See are derived from Halb¬

fass’ table of 1910, they will be found somewhat lower than • those

given, which are from his paper of 1905. The annual heat bud¬

gets, however, are but little altered by the change.

The summer temperatures for Ziiricher See were all taken in

July and may be lower than the maximum.

Nos. 18-20. — These lakes continue in Austria, at a consider¬

able distance, the series of Swiss lakes which ends with Con¬

stance and they lie substantially in the same latitude. Their

heat budgets are also approximately the same.

It will be remembered that there is a doubt both as to the

area and depth of Hallstiitter See. I have taken the smaller

figures. If the larger are taken, the [budgets will be increased

by about 15%.

No. 19. MilSatter See. — Halbfass (13a, p. 312) compares the

temperatures of Gmundener (Traun-) See and Millstatter See as

taken on Aug. 20, 1913. Tie finds the latter lake “erheblich

warmer” than Traun-^ee, and thinks that the difference may

perhaps be due to the east-west direction of the long axis of

Millstatter See, while that of Traun-See lies north and south.

The case is, however, not so simple as this suggestion would

imply, and Halbfass has not called attention to the peculiarity

which seems to be the most important one. The upper strata of

Millstatter See are indeed warmer than those of Traun-See, as

are those of Atter-See (see below). But from a depth of 15 m.

down Traun-See is the warmer and this is the more significant

difference between the temperatures of these lakes.

The observations on Aug. 20, 1913, did not extend to such

depths that the mean temperatures of the lakes can be computed ;

but there can be no doubt that on this occasion, as on all others

in late summer, Traun-See contained much more heat than

Millstatter See. The latter lake agrees in general with the

other lakes of its class in the Alps, both as to the amount and

the distribution of its heat. Traun-See is the exceptional case

among these lakes. The interesting question is not so much

206 | Wisconsin Academy of Sciences , Arts, and Letters.

how the upper 10 m. — 15 m. of Millstatter See come to be warmer

than those of Traun-See; for in this respect Millstatter See

agrees with Atter-See, whose long axis, it may he remarked, ex¬

tends from north to south. We should like rather to know how*

the deeper strata of Gmundener See are able to gain such an ex¬

ceptionally large amount of heat and how its water accumulates

a total quantity of heat so much in excess of that gained by

other and neighboring lakes of comparable area and depth.

Nos. 21-25.— These are illustrations of lakes from the north

slope of the Alps. Wiirm-See and Ammer See lie close together

in the neighborhood of Munich and (are in the relatively low foot

hills. Atter-See and Traun- or Gmundener See lie further east

in [Austria and are in mountain valleys, though their surface

is not so high above the sea as that of the first two named. Their

mean depth) is also much greater. These lakes have heat bud¬

gets entirely comparable both in amount and variation with those

of the group of Swiss Takes which lies on the other side of the

Alps.

I have included Ammer > See whose dimensions are somewhat

less than those of Wiirm-See in order to show how nearly iden¬

tical in these lakes was the summer heat-income in 1881, the

only year when we can compare them. Geistbeck ( ’85 p. 30)

places Wiirm-See among the cold lakes and Ammer See among

the warm, but the mean temperatures taken on successive days

on [the two lakes agree within 5% and that of Ammer See is

the lower. I have no doubt that further observations will show

the same general result since Ammer See has a smaller mean

depth than Wiirm-See.

Nos. 23, 24. Atter-See, Traun-See. — The large budgets of

Traun-See, both annual and summer, are especially noticeable.

Two of the three winter temperatures reported are above 4.0°

and one is much below that temperature, making a great differ¬

ence in the annual heat budgets. I have, however, computed

the wind-distributed heat in each year in which there is a late

summer or early autumn record, relying. on Milliner’s (’99 p. 3)

statement that the water of all these lakes goes annually below

4.0°. I have made a similar computation for the neighboring

Atter-See. The eleven records for Traun-See are both high and

uniform. Traun-See indeed has a much greater amount of wind-

distributed heat than any other central European temperate

lake which is recorded. A comparison of the records of)Atter-

Birge — Heat Budgets of American and European Lakes. 207

See and Traun-See show that in every case where temperatures

were ) observed on closely adjacent days, Atter-See had a higher

temperature at the surface and in the upper strata; but at a

depth of 50 schuh (15.8 m.) or more, Traun-See was the warmer.

It is plain that the larger amount of heat in Traun-See depends

not Jon the reception of a greater supply at the surface, but on

the greater efficiency of the means of distributing it to the deeper

water, or on the larger supply brought to the middle and lower

strata of the lake by the river Traun. The temperature records

for both lakes come from Milliner ’99.

No. 25. Tegern-See. — The budget of this lake is very probably

affected by its altitude which is greater that that of any other

lake in the table. The temperatures for this lake come from

Breu ’06.

No. 26. Arend-See. — This small lake is placed in the table to

show how close the heat budget of a small European lake comes

to that of the larger lakes. An American lake of the same size

would show a smaller heat budget. This lake is not used in the

general discussion of results.

Nos. 27, 28. Loch Ness, Loch Morar. — I have included bud¬

gets from four of the Scotch lakes. All of them have been com¬

puted by myself from data derived from Murray and Wedder-

burn. I have taken as the winter temperature of loch Ness

Wedderburn’s estimate of 41.2° F., 5.10° C. The temperature

given in the table for Sept. 15, 1904 is the mean of five series

taken near the center of the lake on such dates in mid-September

as to make the mean date fall about Sept. 15.

The winter temperature for loch Morar in 1887 is taken from

the bottom temperature found on April 29th of that year. The

lowest temperature recorded is 41.8° F. Halbfass’ figures (’05

p. 227) for Sept. 9, 1887 are undoubtedly] too high.

It is difficult to understand why the budgets of loch J Morar

should be so much smaller than those of loch Ness. The maxi¬

mum depth of loch Morar is greater, and both its mean depth

and area are ample to insure a maximum heat budget. No such

difference appears in the budgets of the mid-European or the

American lakes. It is possible, that, since loch Morar is situated

nearer the sea, the greater cloudiness prevents the temperature

of the water from falling as low in winter as does that of loch

Ness. If, however, the budgets were to be as great as those of

loch Ness, the summer temperatures must also be decidedly

208 | Wisconsin Academy of Sciences, Arts, and Letters.

higher than those recorded. It is possible that this is also due to

lack of sunshine. In any case, it seems plain that the large

budgets of loch Ness are exceptional among the Scottish lakes

in much the same way that those of lake Geneva are among those

of the Swiss lakes.

The figures given by Halbfass (’10, p. 61) would assign a

budget to loch Ness for 1904 amounting to 43,000 cal. This is

somewhat higher than my computation shows, but is a wholly

possible budget and one that loch Ness has doubtless reached.

No. 30. Loch Katrine. — Halbfass (’05, p. 226-227) gives the

temperature of loch Katrine on Mar. 10, 1900 as 5.52° ; Sept. 6,

1900, 11.16°. These figures he repeats (TO, p. 62). If, how¬

ever, the mean temperature of the water is derived from table

1 of his pvper of 1910, by dividing the total heat by the volume

of the lake, the temperature for Mar. 10 would be 4.42° ; Sept.

6, 8.91°. I have computed mean temperatures for loch Katrine,

basing my results for volume on the areas given by Murray ; and

my result for Sept. 9, 1812 is 8.00°, the same as that of Halbfass

( ’05, p. 227) ; so that my hydrographic data seem to be the same

as his. The temperature records which I have used for comput¬

ing the temperatures of 1900 come from Forel ( ’01, p. 37) and I

cannot see that my results are in error. In the temperature rec¬

ords, no reading is given at 25 m., and the temperature depends

somewhat on the reading inserted there, but the mean tempera¬

ture could not go above 8.8° in any case. The mean depth of the

lake as given by Murray (Yol. II, p. 3) is 199 ft. which equals

61 m., not 62 m. as stated by Halbfass. If the temperature de¬

rived from table 1 in Halbfass’ paper of 1910 are employed, the

heat budget would be 27,400 cal., substantially the same as mine.

No. 31. Wettern. — The annual heat budgets of this lake are

by no means extraordinarily large. The wind-distributed heat

is small, owing evidently to the low temperature of the water in

winter and the relatively late date at which the lake reaches the

temperature of 4°. This lake lends no support to Forel ’s doc¬

trine of increasing budgets with increasing latitude. The com¬

ment on Fig. 1 (p. 21) should be consulted for a discussion of

the low summer heat-income of this lake and the next two.

No. 32. Mjosen. — The average heat budget of this lake is large,

though if the budget for 1901 be omitted it is smaller than that

of loch Ness. There is no question but that the temperatures

of the lake for 1901 are correctly recorded and computed but it

Birge — Heat Budgets of American and European Lakes. 209

does not seem possible that they give correctly the mean tem¬

perature of the lake.

Professor Halbfass has kindly informed me of certain correc¬

tions to be made in his computations for Mjosen. The volume

of the lake should be 67.1 cu. km. not 69.1 eu. km. as given in

Halbfass’ paper of 1910. This change reduces the mean depth

to 186.7 m. instead of 192.9 m. I have computed again

the temperatures of the lake on this basis and find them some¬

what different from those assigned by Halbfass. A source

of difference besides the change in volume is in the method of

computation. If in the 0-10 m. level, for instance, readings are

given at 0 m., 5 m., 10 m. Halbfass takes that at 5 m. as repre¬

senting the mean of the stratum; while I use the mean of the

three readings.

The changes in computation make no essential difference ex¬

cept in the summer of 1901. There can be no doubt that the

water south of the island of Helgoen was unusually warm dur¬

ing that season. The mean temperatures are as follows : Apl. 17,

3.17° ; May 26, 3.62° ; July 1, 5.27° ; Aug. 7, 6.41°. The follow¬

ing table shows the gains computed for a mean depth of 187 m.

TABLE 5

GAIN OP HEAT, MJOSEN

A gain of nearly 61,000 cal. in 122 days is quite impossible. So

great a gain as nearly 600 cal. per day during July seems be¬

yond the credible and a gain of over 850 cal. during June is very

far above anything that can be accepted. The gains during

April and May are normal.

The observations were made south of the island Helgoen, which

is about 4.0 km. long and lies some 50 km. from the north end

14— S. A.

210 Wisconsin Academy of Sciences, Arts, and Letters,

of the lake and 34 km. from the south end.* * The 400 m. contour

extends within about 16 km. (less than one-fifth of the total

length of the lake) of the south end of the lake. The observa¬

tions of 1901 did not extend below 400 m. If they were made

at the south end of the deep water the apparently high mean

temperature may be due in part to a temperature seiche and in

part to transport of warm water from the very large north part

of the lake and its accumulation in the south part. If the read¬

ings were taken near Helgoen the latter cause must have pre¬

dominated. This was probably the chief factor in any ease,

since the lake has a form somewhat resembling a funnel with the

broad part toward the north and with a narrow extension toward

the south. This form would make it easy for a north wind to

force large masses of the warm water into the south end of the

lake. In any case, the high temperature of the surface water

shows that the budget for 1901 was unusually large.

In 1901 the temperature taken through the ice on Apl. 17

(3.17°) was lower than that taken on Mch. 9 (3.27°). In the

latter case the readings were taken only to a depth of 320 m. and

probably in shallower water than the April series, which ex¬

tends to 420 m. Very likely the mean of 3.22° would fairly

represent the winter temperature. The water in April was

colder at all depths than in March and of course there was no op¬

portunity for the water to cool under the ice if the lake was com¬

pletely frozen.

The readings of the other years for Mjosen are wholly

reasonable : but it should be noted that the observations taken

on Sept. 14, 1899 extend only to 200 m. and must be conjectur-

ally supplied for the deeper water.

There are no other series from any lake included in Table A

which give clearly incorrect indications regarding the heat bud¬

get. Mjosen is a lake in which such errors would be more likely

to occur than in any other lake included in the list. The shape

of the lake in general, the situation of the island, the eccentric

position of the deep water, the great length of the relatively

shallow north arm, all combine to make it anything but a lake

of simple form in which a single series of temperature observa¬

tions taken at the center may be fairly expected to indicate the

mean temperature of the lake. (See map, Huitfeld-Kaas ?05.)

* The measurements are taken from the small map in Huitfeld-Kaas

*05.

Birge — Heat Budgets of American and European Lakes. 211

No. 133. Ladoga. — The summer temperature given for this

lake is very probably too high. It comes from a series taken on

October 17. The mean temperature derived from a series on

Sept. 7 of the same year was 0.85° lower, indicating a gain of

nearly 4800 cal. per sq. cm. between these two dates. Such a

gain at this time of the year is quite impossible, since it must

be as much as half the total radiation from sun and sky. Either

the earlier series is too low or the later one is too high and no

important conclusions should be based on this single budget.

It may very likely be true, as stated by Halbfass (’10, p. 63),

that ‘‘die Warmezunahme des Ladogasees iibertrifft die des

Wurmsees ganz ausserordentlich ’ ’ ; but the maximum budget for

Wurm-See is 25,600 cal., while if the September temperature is

taken for Ladoga, its budget would be 28,500. This difference

or that shown in the table hardly warrants so strong an asser¬

tion. Very probably, however, the mean of a series of years

would show that the budget of Ladoga is the greater, since its

temperature is very low both in winter and summer. A large

part of its gain of heat, therefore, lies below 4° and just above

that point, and at these temperatures, a larger percentage of in¬

cident heat will be stored than at higher temperatures, like those

of Wiirm-See in summer.

In concluding these notes on Table A, I will add that I have

given care to secure accuracy in the numerical data which it

presents. I can not even hope that complete accuracy has been

attained; when I follow the work of others in this field I find

what seem to me to be errors and I can not doubt that similar

mistakes occur in my data. But I believe that such errors as

may be found to exist in the table — as in the other cases to which

I referred — are not numerous or serious enough to invalidate or

weaken the general results announced in the paper.

I ought to conclude this paper, as I began it, with acknowl¬

edgment of indebtedness to the work of Professor Halbfass.

( ’05, ’10) from which so much of my data has been derived.

212 Wisconsin Academy of Sciences, Arts, and Letters.

Literature Cited.

Birge and Juday ’14. A Limnological Study of the Finger

Lakes, New York. E. A. Birge and C. Juday. Bulletin

Bureau of Fisheries, Yol. XXXII, 1912. Washington, 1914.

Breu ’06. Der Tegern-See. Limnologische Studie. Georg

Breu. Mitt. Geog. Gesell. in Miinchen. Bd. II, H. 1, 1906.

Bruckner ’09. Zur Thermik der Alpenseen und einige Seen

Nord-Europas. E. Bruckner. Geog. Zeitschrift. Jg. XY.

H. 6, Leipsic, 1909.

Delebecque *98. Les Lacs Frangais. A. Delebeeque. Paris,

1898.

Forel ’95. Le Leman. Yol. II. F. A. Forel. Lausanne, 1895.

Forel ’01. Etude thermique des Lacs du Nord de l’Europe.

F. A. Forel. Arch, Sci. Phys. et Nat., T. XII. No. 7. Gen¬

eva, 1901.

Geistbeck ’85. Die Seen der deutschen Alpen. A. Geistbeck.

Leipsic, 1885.

Halbfass ’96. Der Arendsee in der Altmark. W. Ilalbfass.

Petermanns Mitteilungen. Yol. 42, p. 173. Gotha, 1896.

Halbfass ’05. Die Thermik der Binnen-Seen und das Klima.

W. Halbfass. Petermanns Mitt. Yol. 51, p. 219. Gotha,

1905.

Halbfass ’10. Ergebnisse neurerer simultaner Temperaturmes-

sungen in einigen tiefen Seen Europas. W. Halbfass. Peter¬

manns Mitt. Yol. 56, II, p. 59. Gotha, 1910.

Halbfass ’10a. Gibt es im Madusee Temperaturseiches ?

W. Halbfass. Inter. Rev. d. Gesamt. Hydrobiol. u. Hydro-

graphie. Bd. III. Leipsic, 1910.

Halbfass ’13. Review of A. Hamberg: Dichteunterschiede u.

Temperaturverteilung u. s. w. ; W. Halbfass. Int. Revue d.

gesamt. Hydrobiol. u. Hydrographie. Bd. Y. p. 474-475.

Leipsic, 1913.

Birge — Heat Budgets of American and European Lakes . 213

Halbfass 13a. Einfluss der geographischen Lage auf die Warme-

verhaltnisse von Seen. W. Halbfass. Petermanns Mitteil-

ungen. Vol. 59, p. 312. Gotha, 1913.

Homen, ’02. Die Temperaturverhaltnisse in den Seen Finlands.

Th. Homen. C. R. Congres des Nat. et M&decins dn Nord.

Sec. Y, 1. Helsingfors, 1902.

Huitfeld-Kaas ’05. Temperaturmessungen in dem See Mjosen

und in drei anderen tiefen Norwegischen Seen. H. Huit-

feld-Kaas. Arch, for Math. og. Naturvid. Bd. XXII.

Kristiania, 1905.

Miillner ’95. Die Temperaturverhaltnisse der Seen des Salz-

kammergutes. J. Miillner. Jahresber. der K. K. Staats-

Oberrealschule in Graz. 1895.

Murray and Pullar ’10. Bathymetrical Survey of the Fresh

Water Lochs of Scotland. J. Murray and L. Pullar.

Yol. 1. Edinburgh, 1910.

Ule ’01. Der Wiirmsee in Oberbayern. W. Ule. Yer. fur Erd-

kunde zu Leipsic. Bd. Y. 1901.

Ule ’06. Studien am Ammersee in Oberbayern. W. Ule. Landes-

kundliche Forschungen, Geog. Gesellsch. in Munchen, Heft.

I. 1906.

Wedderburn ’09. The Temperature of the Fresh-water Lochs

of Scotland, with special reference to Loch Ness. E. M. Wed¬

derburn. Trans. Roy. Soc. Edinburgh, Yol. XLY, pt. II.

1907.

Wedderburn ’10. The Temperature of Scottish Lakes. E. M.

Wedderburn. Bath. Survey of the Fresh Water Lochs of

Scotland. Yol. I. pp. 91-144. Edinburgh, 1910.

Wojeikoff ’02. Der jahrliche Warmeaustausch in den nordeuro-

paischen Seen. J. A. Wojeikoff. Zeitsch. fiir Gewasser-

kunde. Bd. Y. p. 193-199. Leipsic, 1902.

214 Wisconsin Academy of Sciences, Arts, and Letters.

LIMNOLOGICAL STUDIES ON SOME LAKES IN

CENTRAL AMERICA.

BY C. JUDAY.

During the month of February, 1910, four lakes situated in

Central America were visited for the purpose of making some

limnological observations on them. The purpose of these in¬

vestigations was to make such studies on these tropical lakes as

had been made on a considerable number of lakes in the tem¬

perate zone so that comparisons could be made with respect to

dissolved gases and net plankton content. It was the purpose,

also, to obtain data concerning the vertical circulation of the

waters of deep lakes possessing such markedly tropical charac¬

teristics. A permanent stratification, resulting in a permanent

stagnation of the lower strata of water, doubtless, would have

presented some interesting biological problems, but such condi¬

tions do not obtain in any of these lakes. Lastly, since they are

situated in volcanic regions information was desired with re¬

spect to the effect of such phenomena on the dissolved gases as

well as on the other substances held in solution by their waters.

These four lakes lie on the Pacific slope of Central America.

Two of them are situated in the republic of Guatemala and are

known as lakes Amatitlan and Atitlan. The other two, lakes

Ilopango and Coatepeque, are situated in the republic of Sal¬

vador.

Cljmate.

These four lakes lie far within the tropics, being located near

the middle of the north tropical zone. Lake Atitlan is the most

northerly and it is less than 15° from the equator. The general

region in which they are situated has two seasons during the year,

Juday — Lakes in Central America. 215

a dry or winter season which lasts from November until April,

and a wet season from May to October. December and January

are the coldest months of the year. ' During the period covered

by these studies the prevailing winds blew from the southwest

and at such times there was always very little or no wind at all

during the earlier part of the day, but between 10 a. m. and noon

a southwesterly wind would spring up and blow until about sun¬

set, reaching a maximum strength about the middle of the after¬

noon. Cool northerly winds were noted twice. At lake Atitlan

a strong northerly wind arose about noon on February 12 and

blew continuously for a little more than three days, making the

lake, so rough that work had to be discontinued during this in¬

terval of time. A few days later another norther, called

‘ ‘ norte 7 7 by the natives, was noted at lake Coatepeque.

Some idea of the daily range of the temperature of the air at

two of the lakes during the visits may be obtained from the fol¬

lowing table :

Table I. Air Temperatures at Lakes Atitlan and Ilopango.

At lake Amatitlan in the latter part of January, 1906, Meek1

found by means of maximum and minimum self -registering ther¬

mometers that the lowest temperatures of the day were reached

between 3 a. m. and 5 a. m. and the highest between 2 p. m. and

4 p. m. The total range varied from a minimum of 11.7° C.

(53° Fahr.) to a maximum of 26.1° C. (79° Fahr.)

1 Field Columbian Museum, ZooL, Vol. VII, p. 178. 1908. «,

)

261 Wisconsin Academy of Sciences , Arts, and Letters.

Physical Features of Lakes.

LAKE AMATITLAN.

Lake Amatitlan is situated in 90° 30' west longitude and

14° 25' north latitude. By rail it is 29.6 km. (18.5 mi.) from

Guatemala City and 89.6 km. (56 mi.) from San Jose, the most

important seaport of the republic of Guatemala on the Pacific

coast. Its surface is about 1180 m. (3870 ft.) above sea level and

some of the neighboring mountains rise to an elevation of 250 m.

to 400 m. or more above the lake, thus giving the body of water

the general characteristics of a typical mountain lake. A short

distance southwest of the lake is the volcanic peak Pacaya, which

reaches an altitude of about 2550 m. while to the south lies an¬

other volcano known as Aqua, whose altitude is given by some

authorities as 3750 m. and by others as 4100 m. To the north¬

west of the lake the mountains consist of hard granitic rock but

all of the others in this vicinity are composed largely of loose

material, such as volcanic ash and pumice which are easily

eroded.

The depression occupied by the lake owes its existence to the

forces which produced the surrounding mountains, but the dam

which impounds the waters was produced in part, perhaps, by

volcanic agencies.

Along the greater portion of the lake the shore is rugged and

possesses a steep slope, but about half of the north shore con¬

sists of a fairly broad, low plain which has been built by the

Lobos river and the temporary streams which enter the lake

from the north. (See fig. 1, p. 217.) The higher land back of

this plain is composed of material which is easily eroded so that

the waters coming from this region carry a large amount of

debris. Even during the height of the dry season the Lobos

river brings down a. fairly large amount of this material which

is deposited at the mouth as a typical delta formation. This

plain is an excellent example of the encroachment of land on a

lake. Apparently more than a third of the original area of the

lake is now occupied by this plain, which is about 6 km. long by

about 2 km. wide.

The lake is long and narrow, with its main axis extending al¬

most east and west, being slightly inclined to the northwest and

Juday — Lakes in Central America .

217

southeast. It is irregular in outline. The maximum length is

about 13 km., and the maximum width, about 4 km. The mini¬

mum width is less than half a kilometer. The encroachment of

the land on the north side has narrowed the lake very much a

little east of its middle thus separating it into two basins. At

the narrowest part the lake is crossed by the Ferrocarril Central

de Guatemala and the resort station called Laguna is situated

on the south shore in this locality.

FIG. 1— SKETCH MAP OF LAKE AMATITLAN (AFTER MEEK).

The depths are shown in meters.

The somewhat smaller, eastern portion has a maximum depth

of about 29 m. and the western portion, about 34 m.

The bottom of the lake is uniform and presents a rather nar¬

row marginal shelf ; beyond this the slope is rather steep.

The chief affluent, and in fact the only permanent stream en¬

tering the lake, is the Lobos river. Along the south side and

at the east end of the lake there are several warm or hot springs,

the largest and hottest one being found at Laguna. The water

of the latter is hot enough to boil eggs, about six minutes time

being required for soft boiled eggs.1 The waters contributed by

these hot springs affect the temperature of the lake water only a

few meters from the shore and then only at the surface. The

surplus waters leave the lake at the western end through the

Michatoya river which flows into the Pacific ocean.

The transparency of the water was rather low, a Secchi’s disc

10 cm. in diameter disappearing from view at a depth of 2.75 m.

1 Meek, loc. cit., p. 167.

218 Wisconsin Academy of Sciences, Arts, and Letters.

With a 30 cm. disc, Meek found a transparency of about 3.5 m.

The water is slightly brackish, the total solids amounting to 421

parts per million. Of this amount, sodium and potassium chlor¬

ide constitute 210 parts per million. The other important con¬

stituents are as follows: Silica, 40 parts per million; iron and

alumina, 6; calcium, 56; and magnesium, 7.8.

The larger aquatic plants, such as Typha, Scirpm, Potamoge-

tpn, Ceratophyllum, and Chara, are fairly abundant in the shal¬

low water along the shores, but they grow most luxuriantly along

the lowland on the north side of the lake. A floating plant, Sal -

vinia natans, is widely distributed also, but it is most abundant

in the vicinity of hot springs. Another floating plant, the

water-lettuce or Pistia obcordata is found in the lake, but it is

confined mainly to the low shore on the north side.

LAKE ATITLAN.

Atitlan is a mountain lake situated on the Pacific slope of

Guatemala, about due west of Guatemala city. It is about 40

km. from the nearest railway station, viz., that of the village of

Patulul. The lake lies at an altitude of about 1500 m. above sea

level and is said to have an excellent climate during the entire

year, being free from excessive heat in summer as well as un¬

usually cool weather in winter. Coffee and tropical fruits, such

as lemons, oranges, and bananas, are grown in abundance in

this vicinity. With the exception of a small portion of the

southern shore, the lake is bordered by mountains which rise

to a general altitude of 750 m. or more above its surface. In

many places the shores rise perpendicularly from the water’s

odge to a height of 50 m. or more.

Several craters of extinct volcanoes are found on the south

side of the lake and give mute evidence of the volcanic activities

that once took place in this region. The most prominent cones

are Atitlan and San Pedro. The former rises about 2100 m.

above the surface of the lake, and the latter about 1500 m.

The lake is fairly regular in outline with a prominent bay ex¬

tending southward. (See sketch map, fig. 2, p. 219.) The

maximum length is about 38 km. and the longest axis lies in a

northeast-southwest direction. The maximum width is about

half the length, but the mean width is not more than a third of

the length. It seems probable that the basin occupied by the

Juday — Lakes in Central America .

219

lake resulted from the damming of an ancient valley by the vol¬

canoes which are situated on the south side. This is the only

portion of the shore that is low and beyond it is San Lucas pass

through which the lake probably discharged its surplus waters if

it ever possessed an outlet.

FIG. 2. — SKETCH MAP OF LAKE ATITLAN (AFTER MEEK).

The depths are shown in meters.

There are a few small affluents chiefly on the north side of the

lake. The largest stream enters the lake near Jairal and the

next in size enters near Panajachel. These streams drain small

valleys that are about 1.5 km. wide and 3 km. to 5 km. long. A

small hot spring is situated at the village of Atitlan. There is

no visible outlet.

The marginal shelf along the edge of the lake is very narrow

in most places and the bottom has a steep slope, a considerable

depth of water being found close to the shore. This is shown

also by the very small delta formations found at the mouths of

the streams. While these streams are small, the height of the

adjacent uplands indicates that they have removed a relatively

large amount of material.

220 Wisconsin Academy of Sciences , Arts , and Letters.

The water has a bluish-green color and is much more trans¬

parent than that of lake Amatitlan. In 1906 Meek1 found that

a white disc 30 cm. in diameter could be seen at a depth of

13.7 m. On February 15, 1910, a disc 10 cm. in diameter dis¬

appeared from view at a depth of 10 m. Meek found a maxi¬

mum depth of 322 m. and a rather large portion of the lake, ap¬

parently, has a depth of 300 m. or more.

Owing to the narrowness of the marginal shelf and the steep

slope of the bottom, the larger aquatic plants are rather scarce

along the shore, not being able to gain a foothold in most places.

They are found more frequently along the deltas of the small

affluents and consist of such forms as Typha, Scirpus , Potamo-

geton, and Ckara.

LAKE ILOPANGO.

Lake Ilopango lies in a picturesque valley about 10 km. south¬

east of the city of San Salvador, the capital of the republic of

Salvador. It lies jn 89° west longitude and 13° 42' north lati¬

tude. The surface of the lake is about 490 m. above sea level.

The longest axis of the lake lies in an east-west direction and is

about 9.2 km. in extent; the maximum width is about 7.3 km.;

and the area is 54.3 sq. km. (See fig. 3, p. 221.) The basin oc¬

cupied by the lake appears to have had a volcanic origin, prob¬

ably the coalescence of several points of eruption.2 The region

is still subject to seismic disturbances which have affected the

level of the lake at various times.

Brigham states that most of the small towns in the neighbor¬

hood of Lago de Ilopango were destroyed by earthquakes on De¬

cember 27 and 30, 1879. On January 11, 1880, the water of the

lake had risen more than a meter and it is estimated that on the

following day nearly 14,000,000 cu. m. of water were discharged

through the outlet of the lake, making a stream of greater volume

than the Seine at Paris. On January 20, 1880, about 11 p. m., a

great disturbance was noted near the middle of the lake and the

next morning a pile of rocks was seen from whose midst arose a

column of vapor. These rocks now constitute two small islands

near the middle of the lake. The larger island is 90 m. to 100 m.

long and 20 m. to 30 m. wide and rises from 8 m. to 10 rn. above

1 Log. cit, p. 180.

2 Brigham. Guatemala, the Land of the Quetzal, p. 402. N. Y., 1887.

Juday — Lakes in Central America .

221

the surface of the water. The smaller island is conical, about

20 m. in diameter and 10 m. high. The smaller lies about 200 m.

north of the larger and the water between them has a depth of

45 m.

The shores are fairly steep, the east and southeast shores rising

most abruptly. The surrounding country soon rises to a height

of 300 m. or more above the surface of the water, the only break

in the elevated rim being at the outlet. A short distance east of

The depths are shown in meters.

the lake is the lofty volcanic cone known as San Ramon. At the

western end it receives a small affluent, Qa. del Arenal. This

stream has built a rather large delta at the point where it en¬

ters the lake. The outlet, El Desagiie, is located at the south¬

western eorner of the lake, and this stream is a tributary of El

Chorro, which flows into the Pacific ocean.

Several kilometers of the shore line were examined and com¬

paratively little shallow water was found, a depth of several

meters being found very near the shore. Along the northwest¬

ern shore, for example, the water descends to a depth of 52 m.

within 12 m. to 15 m. of the shore cliff. This cliff has the ap¬

pearance of a fault line. Soundings were not made in all parts

of the lake, but those that are indicated on the accompanying

map (fig. 3) show that the slope of the bottom is steep. Several

years ago a maximum depth of 209 m. was found ; but in the ob-

222 Wisconsin Academy of Sciences, Arts, and Letters.

servations made in 1910, one sounding showed a depth of 215 m.

The slopes of the volcanic islands below the water seem to be

very steep. A sounding about 200 m. south of the larger island

showed a depth of 170 m. and a depth, of nearly 200 m. was

found a short distance east of the islands.

Like lake Atitlan, the larger aquatic plants are not very

abundant in lake Ilopango owing to the steep slopes of the bot¬

tom near shore. Chara seemed to be the commonest form in the

shoaler water.

The water was just a little more transparent than that of

Atitlan, the disc disappearing from view at a depth of 10.5 m.

The fishes inhabiting lake Ilopango are chiefly small and not

very abundant.

LAKE COATEPEQUE.

The Laguna de Coatepeque is situated in the province of Santa

Ana, republic of Salvador, some distance northwest of the city of

San Salvador. Its geographical position is 89° 34' 25" west lon¬

gitude and 13° 49' 33" north latitude. It lies about 8 km. south¬

west of the city of Coatepeque, which is one of the stations on

the Santa Ana branch of the Salvador railroad.

The accompanying sketch map (fig. 4, p. 223) shows that the

lake is almost quadrangular in outline. The east-west axis of

the lake has a length of about 6.5 km. and the north-south one,

about 5.5 km. The basin appears to be of volcanic origin and

possesses crater-like characters. The lake is surrounded by a

narrow margin of valley which has a gentle slope, but beyond

this the shores rise abruptly to a height of 200 m. or more above

the surface of the water, forming a continuous, unbroken rim.

A short distance west of the lake are two volcanoes, Santa Ana

and San Marcelino. The former rises to a height of 1830 m.

above sea level. The surface of the lake is about 760 m. above

sea level.

The marginal valley affords an excellent location for chalets,

a number of which are situated along the southern and eastern

shores. A small island is located at the southwestern corner of

the lake. The lake possesses neither stream inlet nor outlet.

At the time of this visit, February 25, 1910, the water seemed

to be somewhat higher than usual, as it was undercutting banks

which had previously been above the horizon of the waves. The

Juday— Lakes in Central America . 223

lake is said to have a maximum depth of 120 m. but only two

soundings were made during this investigation. One near the

middle of the lake showed a depth Of 110 m. while another

nearer the southern shore also gave a depth of 110 m.

The larger aquatic plants were unusually scarce along that

portion of the shore which was examined.

The depths are shown in meters.

The water of this lake was the most transparent of the four,

a Secchi’s disc 10 cm. in diameter just disappearing from view

at a depth of 12.5 m.

The water contains considerable mineral matter in solution as

shown by the following analysis by Renson.1 The results are

stated in parts per million :

Residue at 180° . 1063.0

Si02 . 6.5

Mg . 86.8

Ca . 36.8

1Barherena, Monografias Departmental VI, Departmento de Santa Ana.

1910, p. 13.

224 Wisconsin Academy of Sciences , Arts , and Letters.

Na

K

197.7

34.5

253.2

301.5

Cl

It will be noted that this water contains about two and a half

times as much matter in solution as that of lake Amatitlan.

Especially significant is the large amount of sodium and chlor¬

ine.

Temperatures.

All of these lakes belong to the tropical class whose waters are

disturbed throughout their entire depths but once each year.

That is, the so-called process of overturning and circulation is

confined to the late autumn and winter months. With the ad¬

vent of cooler weather in the autumn, the temperature of the up¬

per water begins to fall. The cooling takes place at the surface

and it results in the formation of convection currents which aid

in the mixture of the upper strata of water. These currents af¬

fect the water to greater and greater depths as the cooling pro¬

gresses. The wind, however, is a more active as well as a more

important agent in causing the mixture and circulation of the

upper water. As the cooling progresses and the temperature of

the upper water approaches that of the lower, the thickness of

the stratum that is disturbed by the wind gradually increases

Finally the thermal resistance to the mixture of these two strata

becomes so small that the wind is able to produce a general mix¬

ture of the water from surface to bottom. This is known as the

autumnal overturning and it probably takes place between thk

middle of November and the middle of December. Owing to

differences in altitude as well as in latitude the time is different

for the different lakes. There are, doubtless, variations in the

same lake from year to year owing to annual differences in the

weather.

This overturning is followed by a period during which the

water is subject to disturbance by the wind throughout its en¬

tire depth. This is the socalled period of circulation. During

its existence the water has a uniform temperature from surface

to bottom ; that is, the lake is homothermous. The exact time at

which this circulation ceases is not known, but in 1910 it took

Juday — Lakes in Central America .

225

place in all four lakes previous to the time of these observations.

The minimum temperatures which were reached by the waters

of these lakes in the winter of 1909-10 are shown by the bottom

temperatures obtained in these observations. The minimum

temperature depends upon the severity of the winter and doubt¬

less it varies somewhat from year to year. In late January and

early February, 1906, Meek found that the temperature of the

bottom water of lake Amatitlan was 20.5° C. while on Febru¬

ary 5, 1910, it was 19.6°. In lake Atitlan the bottom tempera¬

ture was 20.0° in February, 1906, and 19.2° in February, ’1910.

In this latitude day and night are about equal in length so that

the diurnal period of radiation is substantially equal to that of

insolation. Such a condition is not favorable for the storing

of a large amount of heat in the upper stratum, but during the

latter part of January this water begins to gain heat and this

process continues until the temperature of the upper water rises

a few degrees above that of the lower. As the temperature of

the surface water rises it becomes lighter than the water below

and offers a resistance to mixture with it. Soon this thermal

resistance reaches a point where the wind is no longer able to

mix the upper water with the somewhat cooler water below and

the water of these tropical lakes becomes directly stratified just

as it does in temperate lakes in early summer. Such a stratifi¬

cation was found in these southern lakes in February, 1910.

The temperature differences were not great enough to show the

exact limits of the three different strata, that is, the layers which

correspond to the epilimnion, the thermocline or mesolimnion,

and the hypolimnion in temperate lakes after their stratification ;

but taken in connection with the results obtained for dissolved

gases, they show that similar strata existed at the time of these

observations. Owing to differences in area, depth, and climatic

conditions, the thickness of these layers differed in the different

lakes.

In lake Amatitlan, the middle stratum or mesolimnion lay be¬

tween 10 m. and 20 m. and the temperature at the latter depth

was only 0.5° less than at the former. In lake Ilopango this

zone had the same extremes or limits with a difference in tem¬

perature of 0.6°. In lake Coatepeque, however, the middle

stratum lay between 20 m. and 30 m. and there was a decrease

of 0.5° in this layer. In lake Atitlan the difference in tempera¬

ture between surface and bottom was only about 0.4° in the early

15— S. A.

226 | Wisconsin Academy of Sciences , Arts , and Letters.

part of the day and the greater part of this difference was

found in the upper 10 in. The middle or insulating stratum

had not yet become definitely defined thermally, but the results

obtained from dissolved gases showed that the lower strata no

longer took part in the general circulation of the water. The

decrease in the alkalinity between 75m. and 100 in. indicates

that this stratum was at the lower limit of the circulation. As

the season advanced and the upper water became warmer, this

transition zone undoubtedly came much nearer the surface, very

probably lying at a depth not exceeding 20 m. to 30 m.

The effectiveness of the slight differences in temperature in

preventing the mixture of the water at all depths doubtless finds

its explanation in the fact that the waters had such a high

temperature. The thermal resistance to mixture due to a rise

in temperature of water from 19° to 20° is 25 times as great as

that due to a rise from 4° to 50.1 That is, it would require

twenty-five times as much work to mix two given volumes of

water at temperatures of 19° and 20° as it would take to mix

the same volumes of water if their temperatures were 4° and 5°

respectively. Thus, since the lowest temperature in these trop¬

ical lakes was about 19°, a slight rise in the temperature of the

upper water would produce a marked increase in its thermal

resistance to mixture with the somewhat cooler water below.

The maximum rise in the temperature of the upper water dur¬

ing the summer most probably does not exceed a few degrees

because day and night are about equal in length in this latitude,

thus giving as much time for the radiation as for the accumula¬

tion of heat. But this relatively small increase in the tempera¬

ture of the upper water undoubtedly causes the three strata to

become sharply defined. In fact, it seems probable that they

become so well marked that the resistance to mixture offered

by the transition zone, or middle stratum, is comparable to that

offered by the thermocline or mesolimnion of the deeper lakes

of the temperate zone in summer.

The waters of lake Ilopango were warmest and those of lake

Atitlan were coolest. (See table II, p. 244.) The differences

in temperature were due chiefly to differences in the altitude of

the various lakes, the former lying at the lowest altitude, while

the latter lies at the highest. The maximum difference between

surface and bottom temperature was found in lake Coatepeque.

Birge, Trans. Wis. Acad. Sci., Arts, & Letts., Vol. XVI, Part 2, 1910, p. 992.

Judmj — Lakes in Central America .

227

The results obtained on these lakes are not in agreement with

those obtained by Downes1 on water reservoirs situated in the

Panama Canal Zone. He found that these artificial bodies of

water were permanently stratified and that the transition zone

lay at a depth of only 2 m. to 3 m.

Dissolved Gases.

Oxygen. Samples for the estimation of the dissolved gases

were obtained by means of a closing water bottle which had a

capacity of about one liter. The sample bottles had a capacity

of about 250 cc. so that three samples of water were obtained

from each depth at a single haul. Two of these samples were

used for the determination of the dissolved oxygen and the third

for carbon dioxide. The water was transferred from the water

bottle to the sample bottles by means of a rubber tube and pre¬

cautions were taken to prevent contact with the air. The sur¬

plus water was used to flush out the bottles containing the oxy¬

gen samples. The Winkler method was used for the determina¬

tion of the quantity of dissolved oxygen and in general the re¬

sults given in the table are the mean of duplicate samples.

In this method of determining the dissolved oxygen, 1 cc.

of a solution of maganous chloride is added to the sample and

then 1 cc. of a solution containing sodium hydroxide and potas¬

sium iodide. The sample is thoroughly shaken and the precipi¬

tate is allowed to settle. It is now treated with 2 cc. of con¬

centrated, chemically pure hydrochloric acid, which dissolves

the percipitate. In the chemical reactions which take place,

iodine is liberated in proportion to the amount of dissolved

oxygen present. The amount of the free iodine is then deter¬

mined by titration with a standard solution of sodium thiosul¬

phate. One cubic centimeter of a N/10 sodium thiosulphate

solution is equivalent to 0.0008 gm., or 0.5598 cc. of oxygen at

0° and 760 mm. since 1 1. of oxygen weighs 1.429 gms. at nor¬

mal temperature and pressure.

During the winter circulation the general mixing of the water

at all depths is sufficient to produce a fairly uniform distribu¬

tion of the dissolved gases and other substances held in solution.

The water at various depths is exposed to the air from time to

1A Study of the Water Supplies of the Isthmus of Panama. Proc. Med.

Asso. of Isthmus of Panama, vol. Ill, 1911, p. 133-150, 7 pis.

228 ! Wisconsin Academy of Sciences, Arts, and Letters.

time where it may obtain more oxygen in ease its supply of this

gas is deficient. Through the influence of the wind this aerated

water is forced down into the lower strata and mixed with the

deeper water, thus carrying down a supply of dissolved oxygen.

This process continues during the period of complete circula¬

tion and results in a pretty thorough aeration of the water at

all depths. The upper water also receives some oxygen through

the process of photosynthesis, since in this process the chloro¬

phyll-bearing organisms, under the influence of light, take up

carbon dioxide and liberate oxygen. The decrease in the tem¬

perature of the water in autumn and winter increases its

capacity for oxygen, since this gas is more soluble in cold than in

warm water. As a result of the circulation and the increase in

the capacity of the water for this gas, the largest quantity of

oxygen, held in solution by the waters of these lakes is found

just at the close of the circulation period.

As soon as the lower water ceases to take part in the circula¬

tion it is cut off from further addition to its supply of dissolved

oxygen because this gas diffuses so slowly through water that

the amount transferred in this way is negligible. Hence the

supply is limited to the amount which this water contains at the

time that it ceases to take part in the circulation. Soon an

appreciable decrease in the dissolved oxygen of the lower water

is noted. This decrease is the resultant of two processes, namely,

respiration and decomposition. The lower strata are inhabited

by various organisms which consume dissolved oxygen in their

process of respiration and liberate carbon dioxide. But prob¬

ably a much larger portion is consumed in the decomposition of

organic material. Especially is this true when the upper water

supports a large population of plankton organisms, more particu¬

larly phytoplankton. The decrease is most rapid at the bottom

because decomposable material is most abundant there.

But decomposition takes place at all depths. The floating

devices possessed by the various phytoplankton forms reduce

their specific gravity so that it is but little greater than that

of the water. Thus such organisms sink very slowly when they

are dead and they may pass through the early stages of decay,

at least, at almost any depth. When this decomposition takes

place in the upper water the dissolved oxygen which is con¬

sumed in this process may be replenished because this stratum

is kept in circulation by the wind. Also, the photosynthesis

Juday— Lakes in Central America. < 229

which takes place in this layer aids in maintaining a supply of

oxygen. But the water which lies below the zone , of circulation

is not able to replenish its losses of oxygen until the next period

of overturning and circulation.

Table II, (p. 244) shows that the bottom waters of lakes Am¬

atitlan and Ilopango contained a distinctly smaller amount ;of

dissolved oxygen than their surface waters. The difference be¬

tween surface and bottom in the former lake was 2.5 cc. per

liter of water and in the latter almost 2 cc. In both lakes the

most marked decrease came in the 10-15 meter stratum. There

was an appreciable difference between surface and bottom in

lakes Atitlan and Coatepeque, but it was not nearly so marked

as in the other two lakes. In the former it amounted to only

about 0.5 cc. and in the latter about 0.6 cc.

As the season progresses the difference between surface and

bottom undoubtedly becomes more marked in all of the lakes.

In fact, it seems very probable that the dissolved oxygen is

entirely exhausted in some of the lower water of both Amatitlan

and Ilopango lakes before the autumnal overturning takes place.

In lake Atitlan the volume of the lower water is relatively very

great so that it holds a correspondingly large amount of oxy¬

gen in solution ; the amount is so large, in fact, that the demands

for oxygen in the lower strata may not be great enough to en¬

tirely exhaust the supply in any of this water.

Downes1 found that the water in some of the reservoirs of the

Panama Canal Zone was permanently stratified and that, below

a depth of 3 m., there was practically no dissolved oxygen

throughout the year.

The percentages of oxygen saturation shown in the last col¬

umn of table II (p. 244) are based upon the quantity of oxygen

required for the saturation of perfectly fresh or distilled water

as shown by Fox’s2 determinations. It will be noted that these

percentages are low, even at the surface where the water is=

freely exposed to the air. The highest percentage of saturation

was found in lake Amatitlan and the lowest in lake Atitlan.

The quantity of oxygen absorbed by the waters of these lakes'

is affected by two factors, namely, the salinity of the waters and

the elevation of the lakes above sea level. The presence of

sodium chloride reduces the capacity for oxygen. Chemical

1 Proc. Med. Asso. of Isthmus of Panama, vol. Ill, p. 133-150, 7 pis. 1911.

2 Publications de circonstance, No1. 41, Part I, 23 p.,-1 pi. 1907.

230 Wisconsin Academy of Sciences , Arts y and Letters.

analyses of the water of lakes Amatitlan and Coatepeqne show

that rather large amounts of this substance are found in them

and this would reduce materially their capacity for the absorp¬

tion of oxygen. With increase in altitude there is a decrease in

atmospheric pressure; this means a corresponding decrease in

the tension of the oxygen and the amount absorbed by water will

be correspondingly smaller. The decrease in the amount of

oxygen absorbed is about one per cent, for every 82 m. (270 ft.)

above sea level.

On this basis the saturation point at the altitude of lake

Amatitlan is 14.3 per cent, below that at sea level, or 85.7 per

cent. Since the quantity of oxygen in the upper water amounted

to about 90 per cent, of saturation at sea level, it was actually

somewhat above the saturation point for the altitude of the

lake. The excess was due to the activities of the chlophyll-bear-

ing organisms which were abundant in the upper strata.

At the altitude of lake Atitlan the saturation point is only

about 82 per cent, of that at sea level. But the quantity of

dissolved oxygen in the upper strata amounted to only 66 to

69 per cent, which still left a deficiency of 13 to 16 per cent.

The altitude of lake Ilopango accounts for only 6 per cent, of

the 16 per cent, deficiency, thus leaving 10 per cent, to be at¬

tributed to other factors. In the upper 20 m. of lake Coate-

peque the quantity of dissolved oxygen was from 15 to 19 per

cent, below the saturation point at sea level. About 9 per cent,

of this deficiency can be attributed to the altitude of the lake

which leaves 6 to 10 per cent, still unaccounted for.

In the latter part of table II (p. 244) are shown the results

for dissolved gases which were obtained on two of the Finger

Lakes in the state of New York. These two bodies of water,

namely Cayuga and Seneca lakes, belong to the deeper class

of temperate lakes and they compare very favorably in depth

with three of these tropical lakes, lake Amatitlan being the ex¬

ception. It will be noted that there was a marked difference

between the temperate and tropical lakes with respect to the

amount of dissolved oxygen found in their waters. The quan¬

tity was distinctly larger in the former than in the latter at all

depths. The large amount of this gas found in the lower waters

of the temperate lakes was due to the low temperatures in these

strata which gave the water a much greater capacity for oxygen.

At the time of the observations the surface layers of Amatitlan

Juday — Lakes in Central America .

231

and Cayuga lakes had the same temperature, yet the former

contained one cubic centimeter of oxygen per liter of water less

than the latter. But if the altitude of the two lakes is taken

into account, the amount of oxygen in the surface stratum of

lake Amatitlan was about 4 per cent, above the saturation point

while in Cayuga lake it was somewhat less than 2 per cent, in

excess.

The surface temperatures of Atitlan and Seneca lakes were the

same but the dissolved oxygen of the former amounted to 2.1 cc.

less than that of the latter. If corrections be made for the al¬

titudes of the two lakes, the quantity of oxygen in the surface

water of Seneca lake was about 6 per cent, above saturation

while that in the surface water of lake Atitlan was about 13 per

cent, below the saturation point. With respect to the volume of

oxygen found at the various depths it may be said that the ob¬

servations were made on the temperate lakes in August when the

quantity of this gas was near the minimum for the year, while on

the tropical lakes they were made soon after the close of the win¬

ter period of circulation, and at that time their waters should

contain about the maximum amount for the year.

Carbon dioxide. The Seyler method1 was used for the deter¬

mination of the carbon dioxide. In this method the degree of

alkalinity or acidity of the water is obtained by titration with a

standard solution of an acid or an alkali, as may be necessary,

using phenolphthalein as an indicator. The fixed carbon diox¬

ide is determined by titrating with a standard solution of an

acid, using methyl orange as an indicator. For these determi¬

nations N/44 Na2C03 and N/44 HC1 were used.

The greater portion of the carbon dioxide of lake waters is

generally found in chemical union with other substances, chief

among which are calcium and magnesium. This combined gas

exists in two different states. One portion is in a close chemical

union with these substances and forms their normal carbonates.

This is known as the fixed or combined carbon dioxide. The

other part is in a loose union and comprises that portion of this

gas which converts the normal carbonates into the bicarbonates.

It is called the half -bound or bicarbonate carbon dioxide. The

latter is in such a loose chemical union that the chlorophyllous

organisms are able to use the greater part of it in their photo-

1 Chem. News, vol. 70, 1894, p. 82, 104, 112, 140 and 151.

232 Wisconsin Academy of Sciences , Arts , and Letters .

synthetic activities. This makes the half bound carbon diox¬

ide a very important factor from the biological standpoint.

Whenever this gas is present in excess of the amount required

for the conversion of the normal carbonates into bicarbonates,

it appears in the form of free carbon dioxide and such a water

will give an acid reaction with phenolphthalein. On the other

hand, when no free carbon dioxide is present and some of the

half bound carbon dioxide is lost, leaving an excess of the nor¬

mal carbonate, the water gives an alkaline reaction with this in¬

dicator. In table II (p. 244) the degree of alkalinity or acid¬

ity of the water is shown in terms of carbon dioxide. The minus

sign shows that the water gave an alkaline reaction with phe¬

nolphthalein and the numbers indicate the amount of free carbon

dioxide necessary to produce a neutral reaction. The plus sign

indicates that the water gave an acid reaction and the degree of

acidity is stated in terms of free carbon dioxide.

In lake Amatitlan the water was distinctly alkaline down to

a depth of 15 m. so that it would have required 3.34 cc. of free

carbon dioxide per liter to make it neutral. It was neutral at

20 m. and 25 m. and acid thence to the bottom where the acidity

was equivalent to 2.78 cc. of free carbon dioxide per liter of

water. Similar results for free carbon dioxide are found in tem¬

perate lakes soon after the summer stratification has been well

established.

In lake Atitlan the water gave an alkaline reaction at all

depths. In the upper 70 m. the alkalinity was equivalent to 2.23

cc. of free carbon dioxide per liter of water and it amounted to

only 1.11 cc. at and below 200m. The water of a thermal spring

located at the village of Atitlan had an acidity equivalent to

23.26 cc. of free carbon dioxide per liter, while the fixed carbon

dioxide amounted to 124.65 cc.

The water of lake Coatepeque had the highest degree of alkal¬

inity. In the upper 20 m. it amounted to 5.57 cc. while below

75 m. it was equivalent to 3.23 cc. (See table II, p. 244.)

During the period of complete circulation the water in each of

these three lakes doubtless had a substantially uniform degree

of alkalinity, but when the lower water ceased to take part in the

circulation its alkalinity began to decrease. This result was due

to the liberation of carbon dioxide in the respiration of the or¬

ganisms which occupied this stratum and in the decomposition

of organic material which also took place in this region.

Juday — Lakes in Central America .

23a

Unlike the other three lakes the water of lake Ilopango gave

an acid reaction at all depths. In the upper 25 m. the acidity

was equivalent to 2.78 cc. of free carbon dioxide per liter of

water, while in the bottom stratum it amounted to 7.23 cc. Such

a marked acidity following so soon after the winter period of

circulation would seem to indicate that the water of this lake at

all depths may have an acid reaction throughout the year. Sim¬

ilar results have been obtained on soft water lakes in Wisconsin,

except that such a high degree of acidity has not been found in

the upper water of the latter lakes, their acidity rarely exceeding

one cubic centimeter per liter of water in the epilimnion. Also

in temperate lakes which possess as large an amount of fixed car¬

bon dioxide as lake Ilopango the water becomes alkaline soon

after the vernal overturning has taken place and the water of

the epilimnion, at least, remains more or less alkaline until the

time of the autumnal overturn. But the bottom water of such

a lakq may possess a very high degree of acidity toward the

close of the summer period of stratification. At this time it

may be equivalent to as much as 30 cc. to 50 cc. of free carbon

dioxide per liter of water. Doubtless the acidity of the lower

water in lake Ilopango becomes much greater as the season ad¬

vances, and it may equal or exceed that of these temperate lakes

just before the winter period of circulation is inaugurated.

In general there are four sources of carbon dioxide for lake

waters. They are (1) the air, (2) the decomposition of organic

substances, (3) the respiration of the organisms inhabiting the

water, and (4) ground waters. The atmosphere contains from

three to four parts of carbon dioxide per 10,000 so that water

which is freely exposed to the air absorbs a small amount of this

gas. But only a relatively small amount is obtained from this

source because it is absorbed only in proportion to its partial

pressure which is slight. Lake waters are generally well popu¬

lated with organisms which furnish a supply of decomposable

material at their death. The decomposition takes place at all

depths but is most vigorous at the bottom where the organic ma¬

terial is most abundant. In their respiration these organisms

consume dissolved oxygen and liberate carbon dioxide. Many of

the lower organisms are able to live in water which contains no

free oxygen, but in the intra-molecular respiration which they

carry on a certain amount of carbon dioxide is doubtless im¬

parted to the water.

234 Wisconsin Academy of Sciences , Arts y and Letters.

Rainwater possesses a small amount of carbon dioxide when it

reaches the earth and in passing through the ground it obtains

still more. The. normal carbonates of calcium and magnesium

are only slightly soluble in pure water, but this carbonated water

readily changes the comparatively insoluble normal carbonates

to the much more soluble acid carbonates or bicarbonates. Thus,

when this ground water reaches a lake through springs, it gen¬

erally contains a liberal amount of both bicarbonates and free

carbon dioxide. But, unless the volume of spring water enter¬

ing a lake is relatively large in proportion to the volume of the

lake, the free carbon dioxide content of the whole body of water

is not greatly affected from this source.

The unusual acidity of the water in lake Ilopango was not due

to the absorption of carbon dioxide from the air because the

quantity was larger than would naturally be obtained from this

source owing to the low partial pressure of this gas in the atmos¬

phere. Neither was there any evidence that it was due to car¬

bon dioxide derived from decomposition and respiration. These

processes would have to be taking place very vigorously in order

to maintain such a high degree of acidity in the upper water

which |was kept in circulation by the wind and freely exposed

to the air where the tendency would be to dispose of any excess

of this gas. But there was no indication that these processes

were proceeding with such vigor. The fact that there were

about 3 cc. of dissolved oxygen per liter of water at the bottom

shows also that the 7.23 cc. of carbon dioxide in this water did

not come entirely from respiration and decomposition because, if

it had been produced by these processes, the free oxygen would

all have been consumed in its production.

Of the sources mentioned above, there remains, then, only

the ground water to be considered. Ordinary spring waters

generally contain an abundance of free carbon dioxide and

when the volume of water derived from springs is relatively

large ini proportion to the volume of the lake, the acidity of the

lake water is affected appreciably. But in a lake having as

large a volume of water as lake Ilopango, it would require an

unusually large inflow of spring water to produce and main¬

tain such a high degree of acidity as was shown by the waters

of this lake. No definite data were obtained relative to the

volume of spring water entering lake Ilopango, but about a

quarter of the shore was examined and no springs were noted

Judmj — Lakes in Central America.

235

along this portion. Also the volume of the water discharged

through the outlet stream does not seem to indicate that the lake

receives an unusually large amount of spring water. As noted

above, the water of the thermal spring at lake Atitlan possessed

a high degree of acidity and, since lake Ilopango is situated in

a region that is affected by volcanic disturbances, it seems more

probable that the acidity of its water is mainly of volcanic origin

rather than derived from the usual sources noted above.

Lake Amatitlan had the smallest amount of fixed carbon

dioxide with 32.28 cc. per liter of water and Atitlan came next

in order with 37.84 cc. (See table II, p. 244.) The other two

lakes had distinctly larger amounts, Ilopango having an aver¬

age of about 48 cc. and Coatepeque about 56 cc. It will be

noted that all of these tropical lakes possessed a distinctly

larger amount of fixed carbon dioxide than Cayuga and Seneca

lakes in the state of New York. In fact Ilopango and Coate¬

peque contained more than twice as much as these temperate

lakes.

With respect to the amount of this gas these four Central

American lakes belong to the same class as the majority of the

lakes in southeastern Wisconsin. In this quarter of the state

the waters of most of the lakes that have been tested so far,

possess between 30 cc. and 45 cc. of fixed carbon dioxide per

liter. Only one of these lakes, however, reaches an average of

48 cc. to 50 cc. Toward the end of the summer period of stag¬

nation the bottom water in some of these temperate lakes con¬

tains a larger amount of fixed carbon dioxide, but this is found

only in lakes where the bottom water becomes highly charged

with free carbon dioxide. This highly carbonated water acts

upon the comparatively insoluble normal carbonates in the

bottom mud and converts them into the readily soluble bicar¬

bonates. The maximum amount of fixed carbon dioxide that

has been found in the bottom water of any of the Wisconsin lakes

was found in Garvin lake on two different dates in 1909. It

amounted to 63.2 cc. but the surface water on these dates con¬

tained respectively 35.9 cc. and 34.7 cc.

The Net Plankton.

Plankton catches were made in each of these four tropical

lakes at the same time that samples of water were obtained for

the chemical tests. They were taken with a vertical closing

236 Wisconsin Academy of Sciences, Arts, and Letters.

net whose straining surface was made of No. 20 silk bolting

cloth. No attempt was made to study those plankton organisms

which are so small that they readily pass through the meshes

of the bolting cloth, to which the term nannoplankton has re¬

cently been applied. The hauls were quantitative in nature

and they give a fair idea of the vertical distribution of the vari¬

ous forms constituting the net plankton. The number of or¬

ganisms in each catch was determined by the counting method.

This number was multiplied by the factor representing the co¬

efficient of the net. This factor was determined by means of a

tube 3 m. long and 10 cm. in diameter. At the lower end this

tube was fitted with a sliding door which carried a bolting cloth

strainer and a removable plankton bucket. The tube was low¬

ered into the water while open, then the door was drawn over

the lower end by means of a line and the tube raised, so that

the water within it was strained and the plankton concentrated

in the bucket. Then the net was hauled through the same

stratum and the number of plankton organisms obtained in the

two catches was ascertained by counting.

The results given in table III (p. 246) show the number of

individuals per cubic meter of water in the various strata.

Lake Amatitlan. — The phytoplankton of this lake was charac¬

terized by the great preponderance of diatoms, of which Melosira

was by far the most abundant form. It reached a maximum of

nearly eleven million filaments per cubic meter of water in

the 0-5 m. stratum. Synedra came next in point of abundance.

Clathrocystis was the most abundant blue-green alga, but the

maximum number of this form was found in the bottom stratum.

This was probably an indication of senility; the form having

passed its period of maximum development, the senile indi¬

viduals had sunk into the lower water. The material from lake

Amatitlan contained a greater variety of phytoplankton forms

than that from any of the ‘other three lakes.

Ceratium was the most abundant protozoan. The maximum

number, more than a million per cubic meter, was found in the

0-5 m. stratum.

The rotifers were represented by Triarthra longiseta, Anuraea

stipitata, and Pedalion fennicum. Triarthra was found only

in the 15-20 m. stratum, and Anuraea was rather evenly dis¬

tributed through all strata of the lake. Pedalion was not found

in the upper 10 m. but was present at all depths below this

stratum. It was most abundant in the 20-25 m. stratum.

Juday — Lakes in Central America.

237

Copepods and their nanplii were found at all depths, but

“they were most abundant in the 0-5 m. stratum. Diaptomus

was more abundant than Cyclops in the upper 10 m., but below

this depth the latter was more abundant.

The cladocera were represented by Daphnia longispina var.

Lyalina, Ceriodaphnia pulchella, Ceriodaphnia lacustris, and

Bosmina longirostris. The first three forms were present at all

depths, but Bosmina was confined to the upper 20 m. Daphnia

was more aboundant than the other three cladocera and they

all reached their maximum numbers in the 0-5 m. stratum.

Lake Atitlan. — The phytoplankton was very abundant in this

lake but consisted almost entirely of Melosira gramdata. This

form was most numerous in the upper 15 m. where the average

number was more than 46 million per cubic meter. A few

specimens of Gloeocapsa and Zygnema were found in the upper

15 m.

Four rotifers were found in the catches, Polyarthra platyp-

iera, Anuraea stipitata, Brachionus pala, and a Bdelloid form.

Polyarthra was noted in only one catch, that made in the 5-15

m. layer. Anuraea was found only between 5 m. and 30 m.

Brachionus was by far the most abundant rotifer and was pres¬

ent in the catches from all depths. The maximum number was

in the catch from the 15-30 m. stratum. This form was also

found in considerable numbers in the littoral as well as in the

limnetic region. A very few specimens of a Bdelloid rotifer

were noted.

Cyclops was the only copepod noted in the catches from

lake Atitlan. It did not occupy the 0-5 m. stratum but was

present at all other depths. It was most abundant between 5 m.

and 30 m. The nauplii were distributed from surface to bottom

but they, too, reached their maximum number between 5 m.

and 30 m.

Five cladocerans were found in the catches, namely Daphnia

longispina var. hyalina , Daphnia pulex, Diaphanospma brach-

yurum, Ceriodaphnia pulchella, and Bosmina obtusirostris.

All were absent from the 0-5 m. stratum and D. pidex was not

noted in the upper 100 m. Both Diaphanosoma and Ceriodaph¬

nia were confined to the upper strata as in temperate lakes in

summer; they were found only between 5 m. and 15 m. Both

Daphnia hyalina and Bosmina were present in all catches be¬

low 5 m., but they were most numerous between 5 m. and 15 m.

238 ^Wisconsin Academy of Sciences , Arts , and Letters.

In one of the bottom hauls two specimens of Hyalella dentata

were obtained.

Lake Ilopango. — The net plankton of lake Ilopango was char¬

acterized by the scarcity of the phytoplankton. Relatively

small numbers of Cyclotella and Synedra were obtained in the

upper 25 m. Zygnema was the only other alga noted in the

catches. A few specimens of it were found in the upper 10 m.

The protozoa were represented by a few specimens of Dino-

bryon in the upper 10 m. and by a considerable number of

Tintinnus. The latter was found throughout the depth of the

lake, but was most numerous in the 10-25 m. stratum.

Pedalion fennicum was the only rotifer obtained throughout

the depth of the lake. It was very abundant in the upper 10

m. Anuraea stipitata and Notlnolca longispina were found only

in the 25-50 m. catch.

The only copepod noted was Diaptomus siciloides which was

present at all depths, but was most abundant between 10 m. and

25 m. Nauplii were present in small numbers at all depths.

Only one cladoceran was noted. A few specimens of Ceri-

odaphnia pidchella were found between 10 m. and 50 m.

Coatepeque lake.— -The net plankton of this lake was also poor

in phytoplankton, but it was not as poor as that of lake Ilopango.

Here, too, a diatom, Cyclotella , was the most abundant form.

It was scarce in the upper 25 m., but was distinctly more plenti¬

ful below this depth, the maximum number being found in the

bottom stratum. This distribution seems to indicate that this

form was on the decline.

The blue-green algae were represented by a small number of

Clathrocystis in the upper 25 m. and by a still smaller num¬

ber of Aphanizomenon in the upper 10 m.

Ceratium and Tintinnus were the protozoan representatives.

The former was present in relatively small numbers throughout

the depth of the lake. The latter was much more abundant,

with the maximum number in the upper 25 m.

The rotifers were represented by a contracted Bdelloid form,

probably belonging to the genus Rotifer , and by Pedalion fenni¬

cum. The former was abundant in the upper 25 m. and a very

few of the latter were found in this stratum.

The copepods were represented only by Diaptomus sicilis

which was found at all depths, but which was most abundant in

the upper 10 m. A small number of nauplii was noted from

surface to bottom.

Juday — Lakes in Central America.

239

The cladoeera consisted of Daphnia tongispina var. Tiyalina

and Ceriodaphnia pulchella. A very few specimens of the

former were found below 50 m. The latter was fairly abundant

in the upper 25 m.

The general results that have been obtained by marine plank-

tologists show that plankton organisms are present in much

smaller numbers in the tropical parts of the sea than in the tem¬

perate latitudes, or even within the polar circles. To cite but a

single instance, Lohmann1 found a much larger number of organ¬

isms north of 30° north latitude and south of 25° south latitude

in the Atlantic ocean than between these two parallels. In the

main the greater number in the higher latitudes was due to a

greater variety of forms, the most conspicuous increase being

due to diatoms. But the Coccolithophoridae and the Peridinidae,

which were the common forms from the English Channel to

Buenos Ayres, also reached their maximum numbers beyond the

parallels mentioned above.

Table III (p. 246) shows that there was no marked paucity of

net plankton in these four tropical lakes at the time of these ob¬

servations. In fact the material compares very favorably in

quantity with that of lakes situated in temperate latitudes. The

shallowest member of these tropical lakes, Amatitlan, possessed

the greatest variety of forms. With the exception of its rotifer

population the abundance and variety of forms were quite as

great as might be expected from temperate lakes of similar size

and depth.

A few months after the observations were made on these tropi¬

cal lakes i. e., in August, 1910, similar observations were made

on several of the Finger Lakes in the state of New York. In

both instances, the catches were made with the same net and the

material was preserved and counted in the same manner. The

results obtained with the vertical closing net on the shallower

of the Finger Lakes have already been published, but those ob¬

tained in the same manner in Cayuga and Seneca lakes, the

deeper of the Finger Lakes, have not been published hitherto and

are included here in the latter part of table III (p. 246) for the

purpose of comparing these catches with those obtained on the

deeper tropical lakes.

A comparison of the cladoceran population of lakes Atitlan

1 Veroeffentlieh. d. Instituts f. Meeresk., Univ. Berlin, N. F., Geogr. Natur-

wiss. Reihe, Heft 1, 1912. 92 p., 2 pi.

240 Wisconsin Academy of Sciences, Arts, and Letters.

and ‘Cayuga shows that the number was somewhat greater in

the upper water of the latter, but the former possessed a greater

variety of forms. The copepods were represented only by the

genus Cyclops in lake Atitlan, but by both Cyclops and Diap-

tomus in Cayuga. The number of copepods was very much

greater in Atitlan than in Cayuga.

There was a very marked difference in the rotifer population

of these two lakes, Cayuga having a much larger number as well

as a greater variety. The protozoa were not represented in the

catches from Atitlan but these forms constituted a very import¬

ant element bf the plankton of Cayuga.

The total number of algae was much greater in Atitlan but this

superiority in numbers was due to the presence of only one

species, namely, Melosira granulata.

When lakes Atitlan and Seneca are compared the results are

still more favorable to the former. It possessed a larger number

and variety of cladocera and a distinctly larger number of cope¬

pods. The rotifer population of the two lakes did not differ so

greatly in number but there was a distinctly greater variety in

Seneca lake. The protozoan population was comparatively small

in Seneca and absent in Atitlan. The total number of algae was

very much smaller in Seneca lake but it possessed a greater var¬

iety of forms.

Lake Ilopango possessed a very much smaller number of clad¬

ocera than either Cayuga or Seneca lakes and the copepoda were

represented only by Diaptomus siciloides and its nauplii. In

lake Ilopango the rotifers belonged to but three genera while

in Cayuga and Seneca at least nine genera were present. In

point of number of individuals Ilopango far outranked the tem¬

perate lakes owing to the presence of such large numbers of

Pedalion fennicum, a brackish water rotifer.

The protozoan population of Ilopango greatly exceeded in

number that of Seneca lake, but was smaller than that of

Cayuga. In Ilopango the algae were represented only by a few

diatoms, while the net catches from Seneca lake contained a few

blue-greens and a fair number of diatoms ; those of Cayuga lake

also contained a relatively small number of blue-green algae,

but a fairly large number of diatoms.

The cladoceran population of lake Coatepeque consisted of two

forms, Daphnia hyalina and Ceriodaphnia pulchella. The cope¬

pods were represented only by Diaptomus sicilis whose numbers

Juday — Lakes in Central America. 241

greatly exceeded the total copepod population of either of the

two temperate lakes.

Here, as in the other tropical lakes, the rotifer population was

limited to very few forms, but two in this instance. Protozoa

were nearly as numerous as in Cayuga lake and greatly exceeded

the number in Seneca.

Excluding the diatoms, the number of algae was not very dif¬

ferent in the three lakes. The diatoms were more abundant in

Coatepeque than in Seneca lake, but the number in the former

was much smaller than in Cayuga lake.

List of Plankton Crustacea.

In addition to the collections made at the four lakes which

were visited, material was also obtained at the following places :

Puerto Barrios, Los Amates, and from the mangrove swamp at

San Jose, Guatemala; from the Sunken Gardens and Lakeside

Tivoli at Mexico City, and in the vicinity of San Cristobal near

Mexico City. In the following list the various forms which were

found in the material are indicated, together with the localities

in which they were obtained.

CALANIDAE.

Osphranticum labronectum Forbes. Puerto Barrios, Los

Amates.

Diaptomus siciloides Lilljeborg. Lake Ilopango.

Diaptomus albuquerquensis Herrick. Lake Amatitlan, Mex¬

ico City.

Diaptomus marshi Juday. Puerto Barrios, Los Amates.

Diaptomus sicilis Forbes. Lake Coatepeque.

CYCLOPIDAE.

Cyclops ater Herrick. Mexico City.

Cyclops viridis Jurine. Mexico City, San Cristobal.

Cyclops albidus Jurine. Puerto Barrios, Los Amates, Mexico

City.

Cyclops fuscus Jurine. Lake Atitlan, Mexico City.

Cyclops serrulatus Fischer. Lakes Amatitlan and Atitlan,

Los Amates, Mexico City, San Cristobal.

\

16— S. A.

242 Wisconsin Academy of Sciences, Arts, and Letters.

Cyclops prasinus Fischer. Lakes Amatitlan and Atitlan, San

Cristobal.

Cyclops phaleratus Koch. Los Amates.

Cyclops bicolor Sars. Mangrove swamp at San Jose, Mexico

City.

Cyclops fimbriatus Fischer. Puerto Barrios.

OLADOCERA.

Diaphanosoma brachyurum Lievan. Puerto Barrios, lake

Atitlan, Mexico City.

Pseudosida bidentata Herrick. Puerto Barrios.

Pseudosida tridentata Herrick. Los Amates.

Parasida ramosa Daday. Puerto Barrios.

Daphnia longispina var. hyalina Leydig. Lakes Amatitlan,

Atitlan, and Coatepeque.

Daphnia pulex He Geer. Lake Atitlan.

Simocephalus vetulus 0. F. Mueller. Mexico City.

Simocephalus serrulatus Koch. Los Amates.

Ceriodaphnia pulchella Sars. Lakes Amatitlan, Atitlan, Ilo-

pango, and Coatepeque, and Mexico City.

Ceriodaphnia lacustris Birge. Lake Amatitlan.

Bosmina longirostris 0. F. Mueller. Lakes Amatitlan and

Ilopango, Mexico City.

Bosmina obtusirostris Sars. Lake Atitlan.

llyocryptus spinifer Herrick. Lake Atitlan, Mexico City.

Macrothrix laticornis Jurine. Lake Atitlan.

Macrothrix rosea Jurine. Puerto Barrios, Los Amates, Mexico

City.

Eurycercus lamellatus 0. F. Mueller. Mexico City.

Camptocercus rectirostris Schoedler. Lake Atitlan.

Kurzia latissima Kurz. Mexico City.

Alona costata Sars. Lake Atitlan, Mexico City.

Graptoleberis testudinaria Fischer. Lake Atitlan, Mexico

City.

Dunhevidia setigera Birge. Mexico City.

Pleuroxus denticulatus Birge. Mexico City.

Chydorus globosus Baird. Mexico City.

Chydorus sphaericus 0. F. Mueller. Lakes Amatitlan and

Atitlan, Mexico City, Puerto Barrios.

Juday — Lakes in Central America .

243

In his report on the fresh-water copepoda from Panama

Marsh1 states that the general character of the copepod fauna

of the Canal Zone is much more closely related to the South

American fauna than to that of North America. This, how¬

ever, does not appear to be true of the copepod fauna of Guate¬

mala and Salvador since it consists chiefly of North American

forms. Among the Calanidae only one form, Diaptomus marshi ,

is so far known to be common to the fauna of Panama and that

of Guatemala and Salvador.

I am greatly indebted to Mr. H. K. ITarring for the identifi¬

cation of the various rotifers.

1 Smithso. Miscel. Col., vol, 61, no. 3, 1913, 30 p., 5 pi.

244 Wisconsin Academy of Sciences , Arts, and Letters.

Table II. Observations on Dissolved Gases.

The depth is given in meters, the temperature in degrees centigrade, and the gases

in cubic centimeters per liter of water. The last column shows the per cent of satura¬

tion of the oxygen. In the free carbon dioxide a minus sign indicates that the water

was alkaline, a plus sign that it was acid, and neut. that it was neutral to phenolph-

thalein. The degree of acidity or alkalinity is indicated by the number of cubic

centimeters of carbon dioxide that would have to be removed or added to make the

water neutral.

Lake Amatitlan, February 5, 1910.

Lake Atitlan, February 12, 1910.

Lake Ilopango, February 23, 1910.

)

Juday — Lakes in Central America,

245

Table II. — Continued

Lake Coatepeque, February 25, 1910.

Cayuga Lake, New York, August 11, 1910.

Seneca Lake, New York, August 3, 1910.

246 Wisconsin Academy of Sciences , Arts, and Letters ,

Table III. The Distribution of the Plankton Organisms.

The verticle distribution of the various organisms is shown by giving the number

of individuals per cubic meter of water in the different strata. The members grouped

in the different columns are indicated as follows:

1. Cladocera, D. h .=Daphnia hyalina, D. p ,=Dapbtnia pulex, C.—Ceriodaphnia,

B.=Bosmina, Bi.— Diaphanosoma, P.=Polypiiemus , Ch =Chydorus. 2. Copepods,

I>.=Diaptomus, Li.=Limnocalanus, G.=Cy clops, N .=Nauplii. 3. Rotifera, A. a.=

Anuraea aculeata, A. c .—Anuraea cochlearis, A. s.=Anuraea stipitata, N .=Notholca,

S.=Synchaeta, P .=Polyarthra, Vl.—Ploesoma, A . =A sp lane hna , C .=ConocUilus, Tr.=

Triartlira, P e.=Pedalion fennicum, R .=Rattulus, Bd =Bdelloid rotifer , B. p.=

Brachionus pala. 4. Protozoa, C .—Ceratium, D ,—Dinobryon, P =Peridinium, M.=

Mallomonas, T.=Tintinnus, V.=Vorticella. 5. Algae exclusive of diatoms, C.=Clath-

rocystis, Aph.= Aphanizomenon, A=Anabaena, G.=Gloeocystis, G1 .=Gloeocapsat

Cl .=Closterium, S.~Staurastrum, Z.=Zygnema. 6. Diatoms, A.=Asterionella, C.=

Cyclotella, F .=Fragilaria, M.=Melosira, S.=Synedra, T =Tabellaria.

Lake Amatitlan, February 5, 1910.

Juday — Lakes in Central America,

247

Lake Atitlan, February 11, 1910.

Lake Ilopango, February 23, 1910.

248 Wisconsin Academy of Sciences, Arts, and Letters.

Lake Ilopango, February 23, 1910. — Continued.

Lake Coatepeque, February 25, 1910.

Cayuga Lake. New York, August 12, 1910.

Juday — Lakes in Central America ,

249

Cayuga Lake, New York, August 12, 1910.

Seneca Lake, New York, August 2, 1910.

250 Wisconsin Academy of Sciences, Arts, and Letters.

Seneca Lake, New York, August 2, 1910. — Continued.

Davis — Parasitic Fungi in Wisconsin — III.

251

NOTES ON PARASITIC FUNGI IN WISCONSIN— III.

Supplementary to a provisional list of parasitic fungi in Wis¬

consin. Trans. Wis. Acad. Sciences, Arts & Letters 17:2:846-

984.

J. J. Davis.

The fungus recorded in the provisional list under the name

Synchytrium decipiens Farl. is referred to the chytridiaceous

genus Woroninella by H. Sydow using the combination W. aeci-

dioides (Pk.) Syd. (Ann. Mycol. 15:5:484). Peck’s original bi¬

nomial was TJredo aecidioides which had been proposed previ¬

ously for another fungus which fact has been held by some my¬

cologists to invalidate the publication ; hence the use of another

specific name.

Oospores occur in Wisconsin collections of Plasmopara ribi-

cola Schroet. They are globose, brown, smooth, 33-36/x in di¬

ameter; endospore 3 — 4y thick; oogonia 37 — 40/x filled by the

oospores.

Pcronospora parasitica (Pers.) Tul. Guy West Wilson pro¬

poses the division of this into two species and a like treatment of

P. effusa (Grev.) Ces. (Mycologia 6 :197 et seq.).

Pcronospora trifoliorum D By., does not occur on clover in

Wisconsin as far as observed even in fields where both Trifolium

and Medicago are abundant and the latter infected. The conidia

252 Wisconsin Academy of Sciences, Arts, and Letters.

exceed the dimensions given for this species. I append meas¬

urements made from the conidia of two collections :

These measurements indicate that the conidia were larger on

the earlier date. The meteorological records show that May

13th was a day of low temperature and low relative humidity

(44°-62°. 39-25) while on May 25 the temperature ranged

62°-84° and the humidity 99-66. I take it that to the low

temperature may be credited the larger conidia on May 13th.

This reminds one of Melhus’ finding that a comparatively low

temperature favors germination of conidia of P eronpsporales.

During one season, somewhere in the ’nineties, there appeared

at one station in the suburbs of Racine a destructive outbreak of

Erysiphe on Galium aparine. On examination from time to

time no spores were found in the asci and no specimens were

preserved for that reason as I did not know at that time that

they were not formed during the season. The mildew was

looked for during subsequent years but was not again seen.

From an examination of specimens of Lophodermium pinastri

(Schrad.) Chev. on Pinus Banksiana collected at Millston June

5, 1914, the following measurements were made : asci 115-185 x

22-30 fx : ascospores 55-100 x 314-4/a. It has been distinguished

on the label in the herbarium as var. amplum. The affected

leaves were still in situ.

Phyllosticta paviae Desm. is connected by Y. B. Stewart with

the ascigerous fungus Laestadia aesculi Pk. (Phytopath.

4:399.)

Davis — Parasitic Fungi in Wisconsin — III. 253

Phomopsis vexans (Sacc. & Syd.) Harter is the name given

by Harter to the fungus recorded in the provisional list under

the name Phyllosticta hortorum Speg. (Journ. Ag’l Research

2:338).

The Septoria which occurs on Agrimonia in Wisconsin bears

smaller sporules than the Septoria agrimoniae-eupatorii Bomm.

& Rouss. of Europe as described. They are usually 25-40 x 1/*.

*

There is considerable variation in the appearance of Septoria

on Echinpcystis in Wisconsin. The spots are commonly small,

round and arid such as are attributed to Septoria sicyi Pk. and

S. brencklei Sacc. Sometimes, however, they are angular, inter-

venular, green becoming brown. This is more nearly the kind

of spot described under Septoria echinocystis E. & E. Both

types of spot are sometimes found on the same leaf. Dr. R. A.

Harper kindly compared a Wisconsin specimen with green to

brown angular spots with the type of Septoria echinocystis E.

& E. in the Ellis herbarium and wrote as follows: “The spores

agree as to size etc. The spots in the type are larger, more

brownish in color and rounded with a well defined center. It

seems however that it ought to be the same thing.” (In lit.

Apr. 30, 1914) As these two kinds of spots intergrade I cannot

consider them as due to specific distinctness of the infecting

agents but rather as shade and moisture forms on one hand and

the results of sunshine and dry air on the other, the latter con¬

ditions favoring a process of delimitation. As to the size of the

sporules I find them to range from 20-60 x 1— 2/x. In the form

with round arid spots they are usually shorter than in the one

with angular green-brown spots. For instance in a collection

that could be referred to S. brencklei Sacc. most of the sporules

are about 36/* long with an extreme length noted of 48/*. The

collection of the S. echinocystis type from which a specimen was

sent to Dr. Harper for the comparison has sporules 35-55 x

1-1%/*. There seems to be no reason as yet for changing the

record of these forms from Septoria sicyi Pk.

The entry Septoria stachydis Rob. & Desm. in the Wisconsin

lists seems to have been founded upon immature specimens of

another fungus.

In returning a portion of the type specimen of Septoria inter¬

media E. & E. Mr. Ellis wrote as follows on the packet: “There

254 Wisconsin Academy of Sciences, Arts, and Letters.

was only one leaf; this is part of it. It seems to differ from

S. solidaginicola in its shorter spores but it may turn out after

all to be only a var. of the species”, and then by way of post¬

script, “Try and look into it.” I think that it is now safe to

say that the name should be eliminated by reason of being ap¬

plied to a short spored specimen of a Septoria that occurs in

Wisconsin on both Solidago and Aster and known as S. solidagi¬

nicola Pk. According to the description the sporules of that

species are 4/x in diameter while in our specimens they are

iy2-2y2p. Through the kindness of Dr. H. D. House I have

had an opportunity to examine type material and find the spor¬

ules about 1%^ thick.

Examination of Wisconsin specimens that were referred to

Phlepspora oxyacanthae (Kze. & Schm.) Wallr. shows a fine

branched mycelium, inter- and intra-cellular, ramifying through

the affected portions of the leave's. The aerial branches of this

mycelium constitute the conidia which are assurgent, more or less

strongly curved sometimes even horse shoe shaped, pluriseptate,

60-100 x 4-5/x. These form a loose white felt in patches on the

lower surface of the leaves which suggest a powdery mildew.

No spots are caused but the affected tissues finally become dead

and brown.

Leptothyrium dryinum Sacc. Specimens on Quercus rubra

collected at Minocqua have sporules 15 x 10ft like those of Lepto-

fkyrium maculicolum Wint. but the small fruit bodies borne on

large pale leaf areas are characters of L. dryinum Sacc. A

specimen collected at Racine is probably on Quercus ellipsoidalis.

Quercus alba should be stricken from the list of hosts of this

fungus in the provisional list as I find that the specimen in my

herbarium on that host is Phyllosticta phomiformis Sacc.

Gloeosporium septoriodes Sacc. Saccardo in his description

states that the sporules are always continuous. Winter in his

description of Marsonia quercina Wint. which Saccardo gives as

a synonym, describes the sporules as uniseptate. Ellis & Ever¬

hart in their description of Gloepsporium septorioides Sacc. var.

major E &. E. state that the endochrome is often indistinctly

divided in the center. Wisconsin specimens on Quercus rubra

show occasional sporules with a median septum and the two

halves of the sporule separate at this point resulting in two in-

Davis — Parasitic Fungi in Wisconsin — III.

255

dependent sporules. Some of the spornles attain a length of

30/x.

Gloeosporium thalictri Davis. In specimens collected at

Phlox the spots are larger (10-15 mm.) and sometimes less defi¬

nite than in the type. They become sordid-arid above and the

central portion falls away. The aeervnli are light brown and

amphigenons.

The fungus recorded in the provisional list under the name

Cylindrosporium leptospermum Pk. was originally described as

a Cercospora and the change, for which I was perhaps in some

degree responsible, seems to me to have been ill advised. As I

see it the fungus belongs in Hyphales, Mucedinaceae, microne-

meae, scolecpsporae and I know of no genus into which it fits.

As a result of inoculation experiment by B. B. Higgins the

Cylindrosporium padi Karst, of the provisional list has been

divided into three species and connected each with an ascigerous

stage upon the fallen leaves the following spring. According to

this classification our Wisconsin species would stand as follows:

Cylindrosporium hiemale Higgins

On Primus pennsylvanica

cuneata

Cerasus (cult.)

Ascogenous state Coccomyces hiemalis Higgins.

Cylindrosporium prunophorae Higgins

On Prunus domestica (cult.)

Ascogenous state Coccomyces prunophorae Higgins.

Cylindrosporium lutescens Higgins.

On Prunus serotima.

virginiana

Ascogenous state Coccomyces lutescens Higgins. I assume

that the fungus on Prunus cuneata is identical with that affect¬

ing other members of the host group. Inasmuch as the host of

the typical Cylindrosporium padi Karst, is a member of the

same group as are the hosts of C. lutescens Higgins the distinct¬

ness of the latter species is not established.

Ramularia dioscoreae Ell. & Evht. (Proc. Acad. Nat. Sci.,

Phila., 1891, p. 85) was founded upon leaves of Smilax bearing

Ramularia siibrufa Ell. & Hoi. It is therefore to be elided.

256 Wisconsin Academy of Sciences , Arts , and Letters.

Solidago ulmifolia where given as a host of Eamularia vir-

gaureae Thnem. in the provisional list should be placed under

Eamularia serotina E. & E. instead. Eamularia virgaureae

Thuem. seems to vary from an Ovular ia to a Cercosporella type.

Piricularia parasitica Ell. & Evht. When well developed the

conidia are produced into long slender tips as in Cercospora and

may attain a length of 5Qy.

Fusicladium radiosum (Lib.) Lind. In the provisional list

this combination was erroneously attributed to Lindr. I have

assumed that this is a widespread and variable species which

includes the fungus occurring in Wisconsin the conidia of which

vary from 15-30 x 6-11//,. Venturia tremulae Aderh. in that

case is the ascogenous state. Peck’s description of his Clado-

sporium letiferum (40th Report, p. 64) applies very well to the

fungus occurring in Wisconsin and I therefore take it to be a

synonym. Quite different material was collected on Populus

tremuloides at Pepin in August and referred to var. microscop-

icum (Sacc.) Allesch. The following notes were made from this

material.

Spots 1-4 mm. in diameter, orbicular, sordid-arid above with

a narrow, dark, raised margin, alutaceous below also with a dark

margin and a central paler portion on which the conidiophores

are borne ; conidia continuous, 15-18 x 4/x. This differs from the

variety, as described, in the narrower, continuous, hypophyllous

conidia as well as in the character of the spots.

In the 4th supplementary list, p. 78, I gave notes of a speci¬

men that I had referred to Cercospora meg alopot arnica Speg. To

indicate something of the variation I add notes regarding two

further collections: Spots suborbicular, definite, immarginate,

blackish brown becoming paler with age and finally white in the

center, 2-10 mm. in diameter; conidiophores tufted, septate,

smoky brown, 55-80 x 4-5 y ; conidia slender, straight, attenuate,

pluriseptate, 60-125 x On Bidens connata, Price County,

Sept. 9th, 1911.

Spots suborbicular, light brown with a purple margin and a

white center, paler and devoid of purple below, 4-8 mm. in di¬

ameter; conidiophores amphigenous, fasciculate, varying from

subhyaline to brown, straight, flexuose or bent, continuous, en¬

tire or toothed, 15-45 x Sy ; conidia hyaline, straight, attenuate,

Davis — Parasitic Fungi in Wisconsin — III.

257

pluriseptate, 60-100 x 3-4/a. On Bidens connata, Fountain City,

August 12, 1914.

Telia of Uromyces albus (Clint.) Diet. & Hoi. on leaves of

Vida americana were collected at Sharon in 1889. I am unable

to perceive an arrangement of the verrucosities of the spore walls

in rows.

The rust on Hystrix patula referred to Puccinia apocrypta

Ell. & Tracy in the provisional list is probably Puccinia impa-

tientis (Schw.) Arth. as I am informed by Dr. Arthur.

Gymnoc&nia peckiana (Howe) Trotter is made up of the

aecial Caeoma nitens Schw. and the telial Puccinia peckiana

Howe. Kunkel doubts the relationship (Am. Journ. Bot.

1:37-45).

Telia of Melampsoropsis cassandrae (Pk. & cl.) Arth. were

collected at Solon Springs, June 15th, 1914.

Additional Hosts.

Plasmopara australis (Speg.) Swingle. On Echinpcysiis lob-

ata. Galesville.

Peronospora parasitica (Pers.) Tul. On Cardamine bulbosa.

Madison.

Peronospora trifoliorum D By. On Astragalus canadensis.

St. Croix Falls.

Peronospora chamaecysis Wils. is the species that has been

formed to include American forms such as were recorded in the

provisional list as P. euphorbiae Fckl. (Mycplogia 6 ; 204). On

Euphorbia glyptosperma. Trempealeau.

Sclerospora graminicola (Sacc.) Schroet. On Setaria glauca.

Bridgeport.

Bphaerotheca humuli (D C.) Burr. On Viola canadensis.

Clintonville. Abundant at this station but confined to the sin¬

gle species of violet. Viola scabriuscula was abundant but en¬

tirely free from the mildew.

Erysiphe polygoni DC. On Delphinium (cult.) Poynette.

(H. L. Russell?)

258 Wisconsin Academy of Sciences , Arts, and Letters.

Asterma rubicola Ell. & Evht. On Rubus occidentalism

Grant County, opposite Bridgeport.

Plowrightia mprbosa (Schw.) Sacc. On Prunus pumila.

Shore of Lake Superior in Ashland or Iron County. (L. S.

Cheney.) A specimen in the herbarium of the University of

Wisconsin, collected in 1896, appears to be on this host.

Phyllosticta minima (B. & C.) Ell. & Evht. On Acer sac-

ckarinum. Galesville.

Phyllosticta phomifprmis Sacc. On Quercus bicolor. Alma.

Apparently no one has referred this species to Macrophoma.

Phyllosticta* decidua Ell. & Kell. What I take to be this fun¬

gus has been collected on Hieracium aurantiacum at Phlox.

Sept or ia alnifolia Ell. & Evht. On Alnus crispa. Vilas

County.

Septoria dentariae Pk. On Cardamine bulbosa. Madison.

Sept or ia astericpla Ell. & Evht. On Aster Shortii. Potosi.

Sporules 22-33 x 1/x.

Septoria ribis Desm. On Rib’es nigrum (cult.) Madison.

This was collected in November and the triseptate sporules, as

is often the case with conidia late in the season, show a tendency

to divide at the septa.

Septoria sambucina Pk. On Sambucus racemosa. Neopit.

Septoria atrppurpurea Pk. On Aster sagittifolius. Grant

County opposite Bridgeport. On Aster laevis. Wyalusing.

This has the dark purple spots with small central white areas

but the sporules are 90-1 10ft long as in S. punicei Pk.

Phleospora ulmi (Fr.) Wallr. On TJlmus fulva. Richland

Center (Harper & Reed). Maiden Rock. If Phleospora ulmi

(Fr.) Wallr. is a conidial state of Euryachora ulmi (Fr.) Sehroet.

and the ascigerous condition in Wisconsin is distinct from this

and is what is known as Dothidella ulmea (Schw.) E. & E. then

the conidial state must be distinct also and should have another

name. Rehm has replaced the generic name Dothidella Speg.

by the older Euryachora Fckl. (Ann. My col. £:516). It would

be best perhaps to designate the states as Euryachora ulmea

Davis— -Parasitic Fungi in Wisconsin - — III.

259

(Schw.) and Septogloeum ulmeum. Both the American and

European specimens that I have seen have shorter sporules

(often 30—40/0 than the description indicates and are variable.

Gloeosporium septorioides Sacc. On Quercus rubra. St.

Croix Falls. In this collection the sporules are mostly 18-22/a

long, continuous.

Gloeosporium caryae Ell. & Dearn. On Cary a cordiformis .

Trempealeau and St. Croix Falls. In these collections the acer-

vuli are epiphyllous. This fungus received a second description

under the same name by Ellis & Everhart, hence the citation of

these authors in the provisional list.

Glpeosporium ribis (Lib.) Mont. & Desm. On Ribes gracile .

Trempealeau.

Marssonina potentillae var. tormentillae Trail. On Rubus

triflorus. Phlox. Subcuticular. Sporules 15-18 x 3^/t. Also

collected at Solon Springs.

Microstroma juglandis (Bereng.) Sacc. On Juglans nigra .

Galesville.

Monilia fructigena Pers. On fruit of Prunus virginiana ,

Millston and P. pennsylvanica , Solon Springs.

Ramularia rosea (Fckl.) Sacc. On Salix rostrata. Alma.

Salix pedicellaris. St. Croix Falls.

Ramularia pratensis Sacc. On Rumex altissimus. Maiden

Rock. COnidia about 30 x 3 y. continuous. Conidiophores mostly

shorter.

Ramularia rufomaculans Pk. On Polygonum scandens. St.

Croix Falls. Not abundant on this host.

Ramularia aequivoca (Ces.) Sacc. On Ranunculus septen -

trionalis. St. Croix Falls.

Piricularia grisea (Cke.) Sacc. On Setaria viridis. Bridge1-

port.

Passalora fasciculata (C. & E.) Earle. On Euphprbia ser-

pyllifolia. St. Croix Falls.

Cercpsport circumscissa Sacc. On Prunus pennsylvanica ,

Neopit.

260 Wisconsin Academy of Sciences , Arts, and Letters.

Cercospora althaeina Sacc. On Callirhoe triangulata. Prairie

du Chien and Grant County. The relationship to the form on

Althaea is questionable. The conspicuous black-purple raised

border of the spots in this collection give it a quite different ap¬

pearance. I have labeled it var. praecincta.

Cercospora pentstemonis Ell. & Kell. On Pentstemon grandi-

florus. Pepin.

Cercospora varia Pk. On Viburnum acerifolium. Devils

Lake.

Uromyces acuminatus Arth. Aecia on Steironema ciliatum.

Madison. But a scanty development on this host.

TJromyces proeminens (DC.) Lev. (U. euphorbiae of the pro¬

visional list) . On Euphorbia Geyeri I, III, Pepin.

humistrata , III. Pepin.

heterophylla, III. Pepin.

The rust on the latter is U. poinsettiae Tranz. which is here con¬

sidered a race.

Uromyces hyperici-frondpsi (Schw.) Arth. Aecia, uredinia

and telia on Hypericum majus. Devils Lake.

Puccinia andropogonis Schw. Aecia on Pentstemon gracilis.

Millston.

Huccinia perminuta Arth. On Agrostis perennans. Alma.

Puccinia impatientis Arth. On Elymus canadensis glauci-

folius. Maiden Rock.

Puccinia graminis Pers. Telia on Calamagrostis canadensis.

Madison. (E. T. Bartholomew.)

Puccinia bplleyana Sacc. Aecia on Sambucus racemosa.

Drummond. (L. S. Cheney)

Puccinia polygoni-amphib ii Pers. Uredinia and telia on

Polygonum- scandens. St. Croix Palls. Uredinia on Polygonum

acre leptostachyum. Madison.

Phragmidium disciflorum (Tode) James. ( Ph . americanum

(Pk.) Diet.) On Rosa humilis. Gaslyn and St. Croix Falls.

Pucciniastrum agrimoniae (Schw.) Tranz. Uredinia on Agri-

monia mollis. Prescott and St. Croix Palls.

Davis — Parasitic Fungi in Wisconsin — III . 261

Uredinopsis atkinsonvi Magn. Primary uredinia and telia on

Aspidium novel or acense. St. Croix Falls.

In the provisional list no aecia on Pinus were recorded save an

undetermined Peridermium on leaves of Pinus Banksiana. This

was not because of the absence of such rust forms from the state,

but because field work had not been done in the proper regions

at the proper time to detect them. Since the list was prepared,

however, some attention has been given them, the results of which

it may be of interest to summarize.

The leaf Peridermium on Pinus Banksiana was collected in

July 1907, near Gordon, Douglas County, and at Spooner, Wash¬

burn County. It is presumably connected with Coleosporium

but has not been observed since. Peridermium cerebrum Pk.

occurs throughout the range of Pinus Banksiana in the state and

is quite abundant in some localities. Its effects are serious only

when the infections are multiple or when it attacks the axis.

The distribution of Peridermium pyriforme Pk. is also probably

coextensive with the range of Pinus Banksiana in the state it

having been collected in Jackson, Douglas and Yilas Counties,

but it appears to be very sparsely distributed. In June, 1914,

when special attention was given to it, no more than one speci¬

men was collected in any locality. If this is connected with

Cronartium comandrae Arth. it is not nearly so abundant or

widespread as the telial form. Peridermium comptoniae Orton

& Adams on the contraiy, while it has been observed only in

Douglas and Yilas Counties, occurs in considerable abundance

both as to number of trees attacked and the extent of the out¬

break on the individual tree. As I have seen it this usually oc¬

curs on the trunk near the base while Peridermium pyrijorme

Pk. I have seen only on branches. This is contrary to the state¬

ment of Arthur & Kern ( Mycologia 6: 132). Besides its native

host, Pinus Banksiana , this rust attacked the young trees of PL

nus ponderosa in the plantation of the Board of Forestry in.

Yilas County with severity; a severity due in large measure,,

doubtless, to the fact that the rust does not occur in the native

habitat of this host and hence there has been no breeding out of

susceptibility. Of the European Peridermium fischeri Kleb.

which became thoroughly established on Pinus sylvestris in Door

County, I have written elsewhere. It is hoped that with the de¬

struction of the alternate host, Sonchus, this will disappear from

262 Wisconsin Academy of Sciences , Arts, and Letters.

the state. On our more valuable species of pine, Finns Strobus

and P. resinosa, no rust has been observed in Wisconsin, and it

is hoped that none will be introduced.

Filamentous processes from base to apex of the peridium oc¬

casionally occur in Feridermium comptoniae Orton & Adams.

Aecidium maianthae Schum. On Maianthemum canadense.

Solon Springs. This is probably the aecial stage of the rust on

Phalaris arundinacea that was recorded under the name Puc-

cinia sessilis Schn. in the provisional list. While collecting it a

single sorus, but a well developed one, was found on Streptopus

roseus. Under similar circumstances I once found a single sorus

on Habenaria hyperboi'ea. Such occurrences seem significant as

to the relation of the segregates from Puccinia sessilis Schn.

This was erroneously given the name Aecidium smilacinae

Schum. in the provisional list.

Aecidium ranunculacearum DC. In small quantity on Ran¬

unculus abortivus at Solon Springs where it occurred abund¬

antly on Anemone quinquefoila. The infection of the former

host seems to be very exceptional.

ADDITIONAL SPECIES.

Not reported in the Wisconsin lists.

Uncinula parvula Cke. & Pk. On Celtis Occident alis. Madi¬

son.

Exoascus betulinus (Rostr.) Sadeb. On Betula alba papyri -

fera. Solon Springs. This was seen on but a single tree and

confined to a single branch.

Exoascus communis Sadeb. On fruit of Primus cuneata.

Millston, Solon Springs and Boulder Junction. Abundant in

the former locality in 1914. Exceptionally this attacks petioles

and young leaves which are deformed thereby.

Taphrina flava Farl. On Betula alba papyrif era. Ellison

Bay. I am indebted to Dr. Farlow for authentic material of

this species for comparison. My notes of the Wisconsin collec¬

tion were as follows : Affected areas subcircular, indefinite, yel-

Davis — Parasitic Fungi in Wisco?isin — III. 263

lowish becoming brown below, light green above, about % cm.

in diameter, sometimes confluent; asci hypophyllous, broad and

truncate or somewhat rounded at base, obtusely rounded at apex,

often more or less coustricted in the middle, 30-36 x 18-26/a.

No stalk cell is produced. The constriction of the asci appears

to be caused by the pressure of the encircling edge of ruptured

cuticle and to be greater when the asci are scattered.

Phyllosticta populina Sacc. On Populus deltoides. Prescott.

The light grey spots are circular to subcircular with a narrow

brown margin, 4— 8mm. in diameter. It was associated with a

Septoria.

In Ellis & Everhart’s “The North American Phyllostictas 9 ’

under Phyllosticta grossulariae Sacc. reference was made to Wis¬

consin material on Ribes floridum (= R. americanum ) but it ap¬

pears not to have been placed in my herbarium and was not re¬

corded in the provisional list. I have found it on Ribes vulgare

(cult.) at Fountain City accompanying Cylindrosporium ribis

but mostly immature.

Phyllosticta crataegi (Cke.) Sacc. On Crataegus. Maiden

Rock. This appears to be very close to what has been called

Phyllosticta destruens Desm. on Prunus virginiana and Amel-

anchier.

Vermicularia liliacearum West. Specimens on leaves of

Streptopus roseus appear to be parasitic as one might perhaps

ex-pect from the relation of this fungus to Collet otrichum by rea¬

son of its imperfect pycnidia. The sporules are narrow

(3-4/a) in this collection.

Placosphaeria punctiformis ( Fckl.) Sacc. On Galium bone ale.

Bridgeport. The spermogonial state of Pseudopeziza repanda

(Fr.) Karst.

Ascochyta marginata n. sp. Spots circular to subcircular,

5-15 mm. in diameter, at first green becoming brown with a

paler central portion and a darker periphery and a distinct nar¬

row margin ; pycnidia epiphyllous, scattered, pale brown, irregu¬

larly globose, about 100/a in diameter with a thin cellular wall

and a dark ring around the pore ; sporules hyaline, ovoid to ob¬

long with rounded ends, some of them uniseptate, 6-12 x 2-3%/a.

264 Wisconsin Academy of Sciences, Arts, and Letters.

On Aralia nudicaulis. Phlox, Wisconsin, July 11, 1914. The

smaller and continuous sporules are probably immature but are

in the majority in the material examined. There is evidence

that the affected tissues fragment and fall away, for the most

part probably, before the full maturity of the fungus.

A collection on leaves of cultivated Phlox made at Racine,

Oct. 1, 1896, should probably be referred to Septoria phlogis

Sacc. & Speg. The spots have no colored border ; the epiphyllous

pyenidia are delicate; the sporules range from 25-75 x 1-1 %/a.

Apparently the spots are brown and angular at first becoming

white or sordid and more rounded in outline with maturity.

More or less of the distal portion of the leaves of Co-rices in

Wisconsin are often observed to be dead and on examination

scattered pyenidia are found. Specimens showing hyaline gut-

tulate sporules 10-13 x 2%-3/a were referred to Phyllosticta cari-

cis (Fckl.) Sacc. in the provisional list. Of a collection on Car ex

sp. indet. at Racine it was noted “sporules 10-13 x 3-5/a mostly

becoming uniseptate; some of the sporules germinate without

forming a septum. ’ ’ A collection from Gaslyn on Car ex pennsyU

vanica bears 1-2 septate sporules 15-16 x 4/a ; one from Spooner

on Car ex intumescens has biseptate sporules 16 x 4/a ; one from

Oakwood on Carex sp. indet. shows 1-2 septate sporules

12-18 x 4-5/a while in a collection on Carex pennsylvanica made

at Neopit the sporules are 18-26 x 4-5/a, 3-4 septate. These

seem to me to represent various degrees of maturity and devel¬

opment of a single fungus which is perhaps Stagonospora cari-

cinella Brun. The Neopit collection bears also a Septoria hav¬

ing pyenidia about 100/a in diameter which contain sporules

37-55 x i/2-l/A.

A specimen on Carex retrorsa collected at Athelstane agrees

with the description of Stagonospora paludosa (Sacc. & Speg.)

Sacc.

Septoria acerella Sacc. On Acer Negundo. Galesville. This

agrees with the description given by Dr. Martin in “Septorias

of North America’7. (Journ. Mycol. 3: 79.)

Septoria lophantlii Wint. On Agastacke scrophulariaefolia.

St. Croix Falls. In these specimens the sporules vary in length

up to 80/a.

Davis — Parasitic Fungi in Wisconsin — III. 265

/

Septoria cylindrospora n. sp.

Pyenidia scattered, black, globose to lenticular with cellular

walls, 125-200 in diameter; sporules hyaline, cylindrical,

straight or slightly curved, 18-30 x 2-3/a. On calyces, bracts,

leaves, and upper part of stems (especially on the south side) of

Pedicularis canadensis. Solon Springs, June 1914. Under io¬

dine the sporules show a median division. I have not seen Bhab-

dospora sceptri Karst, of which this may be only a form differing'

in the straight cylindrical sporules. This might be referred to

Ascochyta.

Septpria xanthiifolia Ell. & Kell. On Iva xanthif olia. Alma.

PJileospora celtidis Ell. & Mart. On Celt is occidentaiis. Wy-

alusing. In this collection the sporules attain a length of 100/a,

or more and become 8 or more septate.

Gloeosporium trifolii Pk. On Trifplium pratense. Minocqua.

Abundant at one station.

Collet otrichum graminicolum (Ces.) Wilson. On leaves of

E chinochloa crusgalli. Devils Lake. Setae 50-100 x 5-6/a, spor¬

ules 16-24 x 3 %-6/x. Collected also at Alma and Maiden Pock.

COLLETOTRICHUM SORD1DUM n. sp.

Spots on the upper surface of the leaves varying from orbicu¬

lar to irregular, light brown with a darker margin, 5-15 mm. in

diameter to indefinite and more or less confluent into indefinite

areas, cinereous above from loosened cuticle, on the lower leaf

surface indefinite ; acervuli epiphyllous, scattered, small, flat ;

sporules hyaline, cylindrical with rounded ends, straight, 21-33

x 6 /a ; setae dark brown to black, mostly incurved, 50-75 x 3-6/a.

with a septum near the base below which the seta is abruptly

dilated. The affected portions of the leaves become quite fri¬

able. On Menispermum canadense, Wisconsin river bottom op¬

posite Bridgeport, July 31, 1914. As there is a possibility that

this may connect with Gloeosporium sordidum Speg. of South

America I have used the same specific name.

Didymaria astragali (Eil. & IIol. ) n. comb. {Bamularia astrag¬

ali, Ell. & Hoi.) On leaves of Astragalus canadensis. St. Croix

Falls.

266 Wisconsin Academy of Sciences, Arts, and Letters.

Ramularia spiraeae Pk. On Physocarpus opulifolius. Maiden

Pock. Also observed at Dresser Junction, but too late in the

season to secure good specimens.

Ramularia icxnophila n. sp.

Spots at first indefinite, green, angular, becoming suborbicu-

lar to irregular and more definite but not margined; conidio-

phores hypophyllous, hyaline, fasciculate from a more or less

prominent stromatic base, straight or somewhat bent, continuous,

25-55 x 3-4/x ; conidia hyaline, apical or subapieal, cylindrical,

straight, 1-3 septate, 18-45 x 3-4/x. On Viola canadensis Phlox,

Wisconsin, July 1914. It may be that more knowledge of the

Ramularias occurring on violets will bring together this and sev¬

eral other described species. The apex of the conidiophore fre¬

quently grows beyond the point where the conidium is borne.

Cercosporella scirpina n. sp.

On elongate brown areas which become confluent; conidio-

phores in small tufts disposed in long intervenular lines, hya¬

line, continuous, subulate to cylindrical, often bent and denticu¬

late above, 15-22 x 4-7 /x ; conidia hyaline, straight or curved, ob-

clavate- cylindrical, obscurely septate, 50-122 x 3/x. On leaves

of Scirpus pedicellatus. St. Croix Falls, August 25th, 1914.

Cercosporella filipormis n. sp.

Spots linear, brown, immarginate, %-4 cm. x 1-2 mm. ; conid-

iophores amphigenous, fasciculate, hyaline, continuous, somewhat

lax, 10-15 x 1-2/x ; conidia apical, filiform, hyaline, more or less

curved and lax, sometimes pseudoseptate, 30-75 x 1-2/x. On

leaves of Anemone patens var. Wolfgangima. Millston, Wis¬

consin, June, 1914.

Cercosporella trichophila n. sp.

Effused over indefinite areas on the lower surface of the leaves

which are not discolored; mycelium hyaline, superficial, repent

and ascending the trichomes ; conidiophores racemose on the hy-

phae, hyaline, cylindrical to nodulose, straight, often oblique or

denticulate at the apex, 10-15 x 3-5/x ; conidia hyaline, obclavate-

cylindrical, curved, pluriseptate, 45-75 x 3/x. On Fraximus

pennsylvanica. Bridgeport, Wisconsin, August 1914. The conid¬

iophores and conidia develop especially on the trichomicolous hy-

Davis — Parasitic Fungi in Wisconsin-Ill.

267

phae. The leaf surface becomes dotted with black, globose to

hemispherical, sclerotioid bodies apparently connected with the

same mycelium. Microscopically this fungus suggests a young

Erysiphea. Its systematic position is not clear.

Cercospora camptosori n. sp.

Spots subcircular to angular, pale brown becoming dark brown

with age, immarginate, 3-7 mm. in diameter; conidiophores am-

phigenous, more or less fasciculate, brown, usually undulate, nod¬

ulose or bent, sometimes 1-2 septate, 18-57 x 3-4/a ; conidia hya¬

line, obclavate-cylindrical to flagelliform, straight, 40-100 x 3/a.

On Camptosorus rhizophyllus. Marquette State Park, Grant

County, Wisconsin. August 1st, 1914. This differs from Cer-

cospora phyllitidis Hume, as described, in the shorter conidio¬

phores.

Cercospora muhlenbergiae Atk. On Muhlenbergia sylvatica.

Kenosha County.

Cercospora comandrae Ell. & Dearn. On Comandra umbel-

lata. Trempealeau. Curved and nodulose conidiophores are

not infrequent.

Cercospora sanguinariae Pk. On Sanguinaria canadensis.

Phlox.

Cercospora erysimi n. sp.

Spots pallid, subcircular, 3-5 mm.; conidiophores amphigen-

ous, fasciculate, fuligineous, simple, straight or somewhat in¬

curved, 30-55 x 3-4/a ; conidia straight, obclavate, fuligenous

tinted, about 5- septate, 45-75 x 3-4/a. On leaves of Erysimum

cheiranthoides. Alma, Wisconsin, August 13th, 1914.

Cercospora condensata E. & K. On Gleditsia triacanthps.

Marquette State Park near Wyalusing. Conidia up to 110/a in

length were noted.

Cercospora negundinis Ell. & Evht. On Acer Negundo.

Galesville and Alma. In this collection the conidia are hypo-

phyllous ; the conidiophores range to 40-50/a and the conidia to

150/a in length. As many as 9 septa have been observed in the

latter. The conidiophores are mostly scattered or in twos and

threes. Also collected in Grant County with amphigenous

conidia and at Bridgeport.

268 Wisconsin Academy of Sciences , Arts , and Letters.

Cercospora corni, n. sp.

Spots indefinite, pale brown, becoming mottled with purple*

especially above; conidiophores hypophyllous, scattered, erect

or ascending, brown, septate, 25-40 x5-7y; conidia apical, ob-

clavate, bright brown, strongly pluriseptate, 70-160 x5-7y. On

leaves of C.ornus paniculata. St. Croix Falls, "Wisconsin, Au¬

gust 31st, 1914. The conidiophores sometimes spring from the

arch of a superficial mycelium and. are then shorter. The af¬

fected areas which are mostly 1 cm. in diameter finally be¬

come dark and dotted with small, black, globular, sclerotioid bod¬

ies which are perhaps young pycnidia or perithecia.

Cercospora arctostaphyli n. sp.

Spots circular, definite, sordid-arid with a narrow purple bor¬

der, sometimes confluent, 2-5 mm. ; conidiophores epiphyllous,

springing mostly from small, dark tubercles, subhyaline, straight,,

erect, 7-15 x 3 y ; conidia straight or slightly curved, acute,

30-50 x 1-1 y2/i. On Arctostaphylos Uva-ursi. Millston, Wis¬

consin, June, 1914.

Cercospora echinocystis Ell. & Mart. On Echinocystis lobata

and Sicyos angulatus. Maiden Rock. In these specimens the

conidiophores are scattered rather than fasciculate. Conidia up

to 185 x 4y were measured.

Cercospora effusa (B. & C.) Ell. & Evht. (?). On Lobelia

siphilitica. Alma. In this collection the lax, nodulose, tortuous,

septate conidiophores are 75-150y long; the conidia 30-45y long,

triseptate, becoming brown and constricted at the septa when,

old. Cladosporium effusum B. & C. was said to occur on Poly¬

gonum punctatum, Lobelia puberula and L. siphilitica and Nab-

alus altissimus but Berkeley stated that he had seen conidia only

on Polygpnum and that they are curved which is not true of the

fungus referred to here. On Fungi Columbiani 2505 (on Lob¬

elia in flat a) I find conidia like those in the Wisconsin collection

and also a few slender obclavate ones nearly 100, u long. That,

the latter were borne on the conidiophores I cannot say. The

Fungi Columbiani specimen examined appears to bear a parasite

producing small rod-like sporules in pycnidia. The conidio¬

phores of the Wisconsin collection give off a few branches at or

near a right angle.

Davis — Parasitic Fungi in Wisconsin — III. 269

Cercosppra ageratoides Ell. & Evht. On Eupatorium urticae-

folium. Galesville. In this collection the tufts are scattered

over indefinite, but slightly discolored, spots.

Cercospora grindeliae Ell. & Evlit. On Grindelia squarrosa.

•St. Croix Falls.

Cercospora absinthii (Pk.) Sacc. I am using this name to re¬

cord the occurrence of a Dematiaceous fungus on leaves of Arte¬

misia lud,oviciana at St. Croix Fails regarding which the follow¬

ing notes were made: eonidiophores amphigenous, scattered or

in tufts of 2-6, more or less flcxuous, pluriseptate, brown or

olivaceous, 90-160x4-7^; conidia apical, obclavate to obclavate-

cylindrical, fuligineous tinted, developing about 4 septa, 30-50 x

4-6/x,. The affected leaves show at first brown spots which be¬

come confluent into brown areas. The scattered distribution of

the eonidiophores and the wooliness of the leaves make this quite

inconspicuous.

Frpmyces astragali (Opiz) Sacc. I am using this name for

the purpose of recording the occurrence of uredinia on Astraga¬

lus canadensis at St. Croix Falls. That this American rust is

conspecific with the European one having its aecia on Euphorbia

Cyparissias is questionable. The Sydows in Monographia TJre-

dinearum state that it is not while Arthur in North American

Flora refers it to that species under the synonym Nigredo pustula

(Schroet.) Arth. ( TJromyces pustulatus Schroet.) together with

the Fredo on Oxytropis that has been known as Uredo oxytrop-

idis (Pk.) De Toni. The statement of the Sydows that the ure-

dospores on Astragalus in North America have 6-8 germ pores

Is :pot borne out by this material in which the pores are 3-4.

Aecidium lupini Pk. On Lupinus perennis. Millston. I am

indebted to Dr. J. C. Arthur for the determination. In the 4th

supplementary list mention was made of the occurrence of Tu-

berculina persicina (Ditm.) Sacc. on Lupinus perennis as evi¬

dence that the host bears an Aecidium in Wisconsin winch how¬

ever was not collected until 1914.

Aecidium liatridis Ell. & And. On Liatris scariosa. Solon

Springs.

University of VvTsconsin Herbarium, Madison, Wisconsin,

April, 1915.

270 Wisconsin Academy of Sciences , Arts, and Letters ,

\

INDEX TO HOSTS MENTIONED IN “NOTES” III.

Page

Acer negundo . 264, 267

Acer saccharinum . 258

Agastache scrophulariaefolia 264

Agrimonia mollis . 260

Agrostis perennans . 260

Alnus crispa . 258

Amelanchier . 262

Anemone patens . 266

Aralia nudicaulis . 264

Arctostaphylos Uva-ursi .... 268

Artemisia ludoviciana . 269

Aspidium noveboracense .... 261

Aster . 254

Aster laevis . 258

Aster sagittifolius . 258

Aster Shortii . 258

Astragalus canadensis . . .257, 265, 269

Betula alba . 262

Bidens connata . 256, 257

Calamagrostis canadensis.... 260

Callirhoe triangulata . 260

Camptosorus rhizophyllus ... 267

Cardamine bulbosa . . 257, 258

Carex . 264

Carex intumescens . 264

Carex pennsylvanica . 264

Carex retro rsa . 264

Carya cordiformis . 259

Celtis occidentalis . 262, 265

Comandra umbellata . 267

Cornus paniculata . 268

Crataegus . 262

Delphinium . 257

Echinochloa crusgalli . 265

Echinocystis . 253

Echinocystis lobata . . . 257,268

Elymus canadensis . 260

Erysimun cheiranthoides . 267

Euphorbia Cyparissias . 269

Euphorbia Geyeri . 260

Euphorbia glyptosperma . 257

Euphorbia heterophylla . 260

Euphorbia humistrata ....... 260

Euphorbia serpyllifolia . 259

Eupatorium urticaefolium ... 269

Fraxinus pennsylvanica . 266

Galium Aparine . 252

Page

Galium boreale . 263

Gleditsia triacanthos .... _ 267

Grindelia squarrosa . 269

Habenaria hyperborea . 262

Hieracium aurantiacum . 258

Hypericum majus . 260

Hystrix patula . 257

Iva xanthifolia . 265

Juglans nigra . 259

Liatris scariosa . . 260

Lobelia inflata . 268

Lobelia puberula . 268

Lobelia siphilitica . 268

Lupinus perennis . 269

Maianthemum canadense .... 262

Medicago . 251

Menispermum canadense .... 265

Muhlenbergia sylvatica . 267

Nabalus altissimus . 268

Pedicularis canadensis . 265

Penstemon gracilis . 260

Penstemon grandiflorus . 260

Phalaris arundinacea . 262

Physocarpus opulifolius . 266

Pinus Banksiana . . 252, 261

Pinus resinosa . . . 262

Pinus Strobus . 262

Pinus sylvestris . 261

Polygonum acre . 260

Polygonum punctatum . 268

Polygonum scandens . 259, 260

Populus deltoides . 262

Prunus Cerasus . 255

Prunus cuneata . 255, 262

Prunus domestica . 255

Prunus pennsylvanica . 255, 259

Prunus pumila . 258

Prunus serotina . 255

Prunus virginiana . 255,259,263

Quercus bicolor . 258

Quercus ellipsoidalis . 254

Quercus rubra . 254, 259

Ranunculus abortivus . 262

Ranunculus septentrionalis . . 259

Ribes americanum . 262

Ribes gracile . 259

271

Davis — Parasitic Fungi in Wisconsin — III.

Page

Ribes nigrum . . . 258

Ribes vulgare . . . 262

Rosa humilis . 260

Rubus occidentalis . 258

Rubus triflorus . 259

Rumex altissimus . . 259

Salix pedicellaris . 259

Salix rostrata . . 259

Sambucus racemosa . 258, 260

Sanguinaria canadensis . 267

Scirpus pedicellatus . 266

Setaria glauca . 257

Setaria viridis . 259

'/ ,, 1 v

Page

Sicyos angulata . 268

Smilax . 255

Solidago . 254

Solidago ulmifolia . 256

Sonchus . 261

Steironema ciliatum . 260

Streptopus roseus . 262, 263

Trifolium . . . 251

Trifolium pratense . 265

Ulmus fulva . 258

Viburnum acerifolium . 260

Vicia americana . 257

Viola canadensis . 257, 266

272 Wisconsin Academy of Sciences, Arts, and Letters .

SOME OBSERVATIONS CONCERNING THE BOTANICAL

CONDITIONS ON THE GALAPAGOS ISLANDS.

By Alban Stewart.

Introduction.

When I began the study and identification of the vascular

plants of the Galapagos Islands at the Gray Herbarium, some

seven years ago, I intended to include all of the results in a

single publication. After I had completed that part of the work

included in my paper entitled : A Botanical Survey of the Gala¬

pagos Islands* it was found that such a mass of manuscript

had accumulated that it would probably be better to pub¬

lish this part, and to reserve the general consideration of the

floras of the individual islands for a separate publication.

An attempt has been made in this paper to describe briefly,

and in a general way, the botanical conditions as I saw them up¬

on each of the islands visited. No attempt has been made, how¬

ever, to describe the floras of the different islands in a detailed

way, because, such a consideration would consume too much

space, and furthermore, as our stay in some of the localities

visited was very brief, there was not sufficient time available to

make a sufficiently detailed study of the flora for this purpose.

This is especially true in some of the larger islands, where we

were obliged to get as far into the interior as possible in a short

time, hurriedly collect material, with brief notes, and then start

back to the shore. Expeditions into the interiors of most of the

larger islands are extremely difficult to make. Not only is the

country very rough in most places, and covered with heavy vege-

Proc. Cal. Acad. Sci., fourth series, vol. I, pp. 7-288. 1911.

Stewart — Botanical Conditions on the Galapagos Islands . 273

tation, but there is also no water on the most of them; which

makes it necessary for one to carry a supply of water with him.

On this account it is practically impossible to make trips into

the interior lasting longer than three days.

Department of Botany,

University of Wisconsin,

Madison.

Abingdon Island.

With the exception of the two small islands, Culpepper and

Wenman, Abingdon is the most northern island in the group. It

is located about thirteen miles northwest of Blindloe, and is the

smallest one of the islands that supports an extensive mesophy-

tic flora. This condition is brought about by the fact that it

reaches an elevation of 1950 ft., and consequently it receives a

greater amount of moisture than the other small islands. This

island was visited during the month of September 1906. The

most of the work was done on the south side where good anchor¬

age was found for our vessel in a small bay.

The shores along the south side of the island are composed of

lowv lava cliffs and occasional sand-beaches. The shores become

steeper, however, towards the southwest side. On the west side

there are perpendicular cliffs which rise directly from the sea

to a height of over 1,000 ft. The north and east sides of the is¬

land were not visited, but judging from the appearance of these

parts as seen from the vessel while sailing in the vicinity of the

island, the shores are low, and the sides of the mountain are cov¬

ered with lava to a considerable elevation. The lava covering

the south side is mostly basaltic in character with occasional

beds of volcanic cinders intermingled. This lava is of compara¬

tively great age, and it has become stained to a redish-brown col¬

or through surface oxidation. There are extensive deposits of

volcanic cinders on the southeast side of rather recent origin, the

most of which have come from a small crater at an elevation of

1,000 ft. There are still slight evidences of volcanic activity

around the base of this crater, as there is a constant escape of

steam here, which is sometimes great enough to be seen from the

shore. There is still another small cinder cone on the recent

lava near the shore, from which the lava in its immediate vicin¬

ity must have come. Remains of several other small craters

18— S. A.

274 Wisconsin Academy of Sciences , Arts , and Letters.

occur on the old lava around an elevation of 500 ft., all of which

have probably been inactive for a very long time. The lava on

the south side has been thrown into ridges and folds in places,

and there are also occasional lava tunnels the tops of which have

fallen in. One of these is located near the shore and is filled

with sea water. The south side of the island slopes up gradu¬

ally to an elevation of about 500 ft., above which the slope is

steeper.

The top of the mountain was envelloped in fog at the time it

was visited so that a survey of the surrounding region could not

be made. There seemed to be no central crater present, however,

and the highest part may be the remains of a portion of the rim.

There is a range of hills, about two miles west of the summit,

which runs parallel with the coast line, and have an elevation of

about 1,200 ft. These hills rise abruptly from a broad, flat

plain just east of them, which has an average elevation of 900 ft.

It is possible that this plain may be the floor of an old crater, the

rim of which has been mostly removed.

A few herbaceous halophytes grow on the sand beaches near

where we anchored. There was also a low thicket of bushes of

Laguncularia racemosa bushes growing here. Other than these

no halophytes were found.

All of the vegetation of the south and southeast sides of the

island below an elevation of 450 ft. consists of species w^h are

usually found on the lower and dryer parts of these islands.

They are smaller and fewer in number, however, than is usually

the case, a condition that may be due to the very scanty soil on

these parts. The lava on this part of the island is bare in most

places, and the only soil to be found is in the lava crevices. In

consequence of this condition, a large part of the surface is not

suitable at present for the support of higher plants. The trees

of Bursera graveolens are small, seldom exceeding a height of 8

ft. Usually they are mere bushes. Besides the small Bursera

trees, Opuntia galapageia is the only other species which reaches

the size of a tree in these lower regions. It occurs here abund¬

antly, and has weak spines and closely arranged branches. Eu¬

phorbia viminea forma castellana is the most common bush in

this region. It is about the only one that occurs in mass, all

other bushes being scattered. This species seems to be better

adapted to maintain an existence under the sterile conditions

than most of the other species found here. Other bushes found

Stewart — Botanical Conditions ,on the Galapagos Islands. 275

growing in this region, less abundantly than the above are : Cor-

dia lutea, Castela galapageia, Croton Scouleri var. brevifolius,

Euphorbia articulata, Prosopis dulcis, Scalesia Hopkinsii, Tel-

anthera echinoeephala, and Waltheria reticulata. The lava

ridges are often found to be more or less covered with vines

of Ipomoea Habeliana, and in various other places on t the

lava, I. Kinbergi was found growing and in blossom at the time

of our visit. Such grasses as Aristida suspicata, Cenchrus platy-

acanthus, Leptochloa albemarlensis and L. Lindleyana grew

with more or less abundance in the lava crevices. On the older

beds of volcanic cinders there was very little vegetation other

than occasional bunches of Cereus nesioticus; the more recent

beds of cinders were bare.

A change, readily noticed, takes place in the vegetation at

about 450 ft. elevation evidently brought about by the greater

amount of moisture and a more abundant soil. There is a gen¬

eral thickening up and an increase in the size of the vegetation

above this elevation. There are good sized trees of Bursera

graveolens in this region, and also trees of Pisonia floribunda,

which first make their appearance. Many of the trees and

bushes, at this elevation and above, are heavily covered with

Alectoria sarmentosa and other epiphytic lichens. Bushes and

small trees of Zanthoxylum Fagara also occur, usually infested

with Phoradendron Ilenslovii. Other bushes which occur in this

region and a little higher are : Chiococca alba, Erigeron tenui-

folius, Lipochaeta laricifolia, and Lippia rosmarinifolia. Such

ferns as Chelianthes microphylla, and Polypodium squamatum

occur. Very little change takes place in the vegetation on the

southwest side of the island below an elevation of 700 ft., prob¬

ably due to the fact that this side receives less moisture than the.

south and southeast sides.

Practically all of the plants which occur below 500 ft. dis¬

appear by the time an elevation of 1,000 ft. is reached. The spe¬

cies that continue into this region from below, are, for the most

part, those which first appeared around an elevation of 500 ft,

and above. The region between 1,000 and 1,650 ft. elevation is

covered with forests, on the southeast side, which are made up

mostly of Pisonia floribunda, and Zanthoxylum Fagara. There

is much undergrowth in these forests, consisting of bushes of

Croton Scouleri var. grandifolius, Erigeron tenuifolius, Lippia

rosmarinifolia, Psychotria rufipes, Scalesia Hopkinsii, Tounie-

276 Wisconsin Academy of Sciences , Arts , and Letters.

fortia psilostachya, and T. rufo-sericea, many of which are coy-

red with vines of Cissampelos Pareira, and Elaterium cor datum.

There are also many ferns among* which are : Adiantum Henslo-

vianum, Asplenium cristatum, Nephrolepis biserrata, Polypo¬

dium pectinatum, P. squamatum, and Trachypteris pinnata.

Many herbaceous plants also occur in this region.

There are open areas in the vegetation on the south side of

the island between 1,000 and 1,300 ft. elevation, which are cov¬

ered with grasses and herbaceous plants. These areas extend in

a more or less direct way up the side of the mountain, and are

bordered by bushes which are heavily covered with a growth of

brown Hepatic, probably a species of Frullania. These areas

are succeeded above by a heavy growth of bushes and small trees.

Above 1,650 ft. elevation, extending to the top of the moun¬

tain, there is a heavy growth of ferns which are often five feet or

more in height. Among the ferns there are low stunted bushes

of Zanthoxylum Fagara heavily covered with epiphytes.

The dry region*' on this island extends to about 450 ft. on the

southeast side, and to about 700 ft. elevation on the southwest

side. Judging from the appearance of the vegetation as seen

from a distance, this region must extend to an elevation of about

1,000 ft. on the north side. The transition region extends to an

elevation of about 1,000 ft. on the south and southeast sides, and

probably to within a short distance of the top of the north side.

There is apparently but a narrow strip near the top, on this side

of the mountain, that is covered with dark green vegetation, such

as is usually found in the moist region of these islands. All of

the country above an elevation of 1,000 ft. is covered with the

plants usually found in the moist regions.

Albemarle Island.

Albemarle lies towards the west side of the archipelago and is

the largest island of the group. It is about seventy-five miles

long, and forty-five miles broad at its widest part, which is to¬

wards the southern end of the island. The island has the gen¬

eral shape of the letter L the long limb of which extends in a

general northwest and southeast direction. There are five large

* For a discussion of the botanical regions on these islands see : Stewart,

A Botanical Survey of the Galapagos Islands. Proceedings of the California

Academy of Sciences, fourth series, vol. I, pp. 206-211. 1911.

Stewart — Botanical Conditions on the Galapagos Islands. 277

mountains on the island which vary in height from 3,150 to prob¬

ably over 5,000 ft., and several mountains of lower altitude. All

of the larger mountains are extinct volcanoes each of which has

an immense crater at its top. These craters are all inactive at

the present time except the one northwest of Villamil, on the

south side of the island, Sulphur fumes and other vapors issue

from the floor of this crater at times, and there are also two small

active sulphur volcanoes in it, each of which is surrounded by a

large quantity of almost pure sulphur. There has been some

volcanic activity at Banks Bay during the last few years, from

small craters on the Avest side of the mountain. With these ex¬

ceptions there has probably been no volcanic activity of the is¬

land for a great many years. There are many low hills on vari¬

ous parts of the island, some of which are small craters or blow¬

holes, and others simply masses of volcanic debris.

Banks Bay.

Banks bay is a broad, open roadstead on the west side of the

island, near its northern extremity. The main mountain at this

place is a broad flattopped crater with steep sides, which prob¬

ably rises to a height of over 5,000 ft. There is still a smaller

mountain close to the north shore of the bay that has an eleva¬

tion of 2,360 ft. according to the chart issued by the Hydrogra¬

phic Office. There are also a number of smaller craters and

hills around the base of the main mountain, and along it sides,

which usually have an average height of less than 100 ft. The

base of the main mountain is separated from the shore by a

broad plain which is covered with beds of comparatively recent

lava on which there is very little vegetation. There are places,,

however, on this plain which were not covered by the more re¬

cent flows of lava, on which there is a considerable amount of

xerophytic vegetation.

Unfortunately this region was not explored botanically, so

that all of the information concerning its flora is due to the kind¬

ness of other members of the expedition who visited this place..

At least three botanical regions are represented here, viz.: the

dry, transition, and moist regions, and possibly a fourth, as the*

vegetation around the top of the mountain appeared from a dis¬

tance to be quite different from that lower down. With the ex¬

ception of the transition, no estimate was made concerning the:

extent in elevation of these regions.

278 Wisconsin Academy of Sciences, Arts, and Letters.

The shores support many of the smaller halophytic plants

found on these islands, and ‘ ‘ large mangroves ’ \ probably Lagun-

cularia racemosa, and Rhizophora Mangle, occur abundantly in

places. The older lava around the base of the mountain is cov¬

ered with forms usually found in the dry regions. There are

occasional specimens of an arborescent Cereus, and a low species

of Opuntia occurs abundantly around the base, and on the sides

of the mountain to an elevation of 1,600 ft. The trees in the

lower regions are mostly of Erythrina velutina and Bursera

graveolens, the last one of which was found to extend up to an

elevation of 1,700 ft. on the side of the mountain. Many bushes

and shrubs occur on the lower parts, but with the exception of

Lipochaeta lariciTolia, the names of these are not known. They

are probably all of species usually common on the lower parts of

these islands. Beds of “maidenhair fern”, probably Adiantum

Henslovianum, were found in a lava cavern at the base of the

mountain. All together six species of ferns were noticed on this

part of the island, but it is very likely that many more could be

found if careful collecting were done in this region. Croton

bushes are abundant, and occur to an elevation of 2,300 ft. as

high as this mountain was explored by the members of the party

who visited this part of the island. A “broad-leaved variety of

Croton”, probably C. Scouleri var. grandifolius, occurs high up

on the side of the mountain, and ‘ 1 small-leaved varieties of Cro¬

ton” occur around its base. There are one or more flows of re¬

cent lava down the west side of the mountain which are bordered

by a heavy growth of bushes and morning glory vin$s around an

elevation of 2,300 ft. Above this there are forests which are ap¬

parently made up of an arborescent species of Scalesia, and other

trees. Orchids, and “sword ferns” were also noticed in the up¬

per regions visited.

Tagus Gove.

Tagus Cove is located on the wnst side of the island about op¬

posite the northeast corner of Narborough Island. It has been

formed from and old tufa crater the southwest side of which has

'been removed leaving a small and well protected bay inside.

The inner walls of the crater form steep bluffs which surround it

on all sides except the one open to the sea_. In some places these

bluffs are 600 ft. high, but they are much lower than this towards

the north end of the cove at which place a small ravine enters it.

Stewart — Botanical Conditions fin the Galapagos Islands, 279

There are also two other tufa craters in this vicinity. The

smaller one of these is located just north of the cove and con¬

tains a miniature salt-water lake, while the larger is situated

about a mile south near the coast. This third crater has prob¬

ably formed a small bay some time in the past, as its walls are

broken down on the side next to the sea, similar in this respect

to the crater that forms Tagus Cove. The opening has been

closed, however, by a flow of lava across it, and is now filled with

salt water which comes in through the cracks in the lava from

the sea a short distance away. There are four islets in this lake,

one of which has a small crater on it. It is evident from the de¬

scription, given by Darwin in his Voyage of the Beagle, that this

is the salt water lake that he describes as being located near

“ Banks Cove”. The sides of all these craters are much cut up

by gullies which have been eroded in them. All three of the

tufa craters just described, are separated from the base of the

mountain by a plain, about one and a half miles wide, which is

covered with deposits of volcanic cinder northeast of the cove.

These cinder deposits extend along the base of the mountain

northward and are continuous with the lava beds in the Banks

Bay region. The cinder beds do not extend south of Tagus

Cove, however, as the country around the base of the mountain

in this direction, is covered with deposits of tufa, which extend

out to the coast and form cliffs 40-50 ft. high.

The mountain lies northeast of the cove and is the second one

of the three mountains that make up the northern part of the is¬

land. The mountain at Banks Bay is the one furtherest north,

and the one at Cowley Bay furtherest south in the chain. The

west side of the mountain, opposite the cove, is rather steep to

an elevation of 2,500 ft. There are extensive deposits of tufa be¬

low this elevation, in which deep canyons have been eroded, and

small gullies are common everywhere. There are two flows of

comparatively recent lava, covering the tufa, and extending

down the side of the mountain. They have evidently originated

from small craters near an elevation of 2,500 ft. Deep fissures

occur in these beds in places. The side of the mountain is covered

with extensive deposits of partly disintegrated lava, above an

elevaton of 2,500 ft., which is similar but much older than the

lava which has formed the flows down the side below this eleva¬

tion. The north side of the mountain seems to be entirely cov¬

ered with lava.

280 Wisconsin Academy of Sciences , Arts , and Letters.

The top of the mountain is 4,000 ft. above sea level so that this

is probably the third highest mountain on the island, and the

fourth highest in the archipelago, the mountains at Banks Bay,

Iguana Cove, and the one on Narborough Island exceeding it in

height. There is an immense crater at the top which is about

four miles long and three broad as nearly as could be estimated.

The inner walls of the crater are nearly perpendicular in places.

The floor forms a broad flat plain, possibly 500 ft. below the rim,

which is covered with volcanic ashes, and beds of basaltic lava,

and cinders. There is a somewhat smaller crater inside the

larger one.

The tufa hills in the vicinity of the cove are covered with a

sparse growth of low bushes the most common species of which

are : Acacia macracantha, Croton Scouleri var. Macraei, Euphor¬

bia diffusa, Lipochaeta laricifolia, and Waltheria reticulata.

There are also a few low trees of Bursera graveolens with

rounded crowns, and a considerable amount of Opuntia insularis.

There are many places where the ground is nearly bare of vege¬

tation, and although we visited here in March at the end of the

rainy season, the prospect was far from inviting. Several

grasses occur in these open areas among which are : Aristida sub-

spicata, Anthephora hemaphordita, Bouteloua pilosa, Cenchrus

granularius, and other herbaceous plants.

With the exception of an occasional specimen of Cereus sclero-

carpus the lava beds around the base of the mountain are bare of

vegetation except in protected places where a few grasses and

other small plants occasionally appear. On the fiat area south

of the lava beds, which is covered with tufaceous soil, the vege¬

tation is thicker than it is on the tufa hills but is made up largely

of the same species with the addition of a few others. Bushes

and small trees are common here especially along the edges of

the lava beds where they often grow to a larger size and occur in

greater numbers than elsewhere. A few ferns are to be found

in protected places in this region.

The tufa deposits above the base of the mountain are covered

to a considerable elevation with forms which are practically the

same as those on the plain below, except that the arrangement is

somewhat different, there being many open areas which are cov¬

ered with grasses and other herbaceous plants. The canyons

here often have a heavy growth of Croton and Gossypium bushes

which grow much taller than they do in the more exposed places

Stewart— Botanical Conditions pn the Galapagos Islands . 281

outside the canyons. There is no very marked change in the

character of the vegetation to an elevation of 2,500 ft., as far as

the tufa deposits extend, except that the vegetation thickens up

in places and such conspicuous forms as Pisonia floribunda,

Tournefortia rufo-sericea, and Zanthoxylum Fagara are added.

The two lava flows down the side of the mountain, which cover

the tufa deposits, are bare of vegetation except for occasional

bushes of Erigeron lancifolius, Euphorbia viminea, and Wal-

theria reticulata, while the only plant of tree-like proportions is

Cereus sclerocarpus. In the deep crevices of this lava, however,

there is a more abundant vegetation as trees and bushes of Bur-

sera graveolens, Cordia lutea, and Zanthoxylum Fagara are to

be found, as well as a few ferns, among which are Asplenium

formosum and Notholaena sulphurea.

The side of the mountain above an elevation of 2,500 ft. is

covered in most places with low bushes, the most common one of

which is Lipoehaeta laricifolia. This condition continues to

within about 150 ft. below the rim of the crater, where there is

a narrow zone covered with a dense, and almost an impenetrable

growth of Pennisetum exalatum. The rim of the crater is cov¬

ered with bushes of Cordia galapagensis, Croton Scouleri var.

Maeraei, Dodonaea viscosa var. spathulata, Lantana peduncu-

laris, Maytenus obovata, Telanthera nudicaulis, and Scalesia

microcephala. Opuntia myriacantha also occurs here but the

specimens are smaller and not so profusely branched as they were

lower down. There are a number of herbaceous plants and

ferns among the other vegetation. The vegetation on the floor

of the crater appeared to consist of occasional specimens of Cer¬

eus sclerocarpus and clumps of Dodonaea bushes.

It seemed impossible to divide this side of the mountain into

botanical regions, as was done at the most of the other places vis¬

ited. There is a great similarity in the vegetation all over this

side, and the forms which occur at the top of the mountain are

mostly of the same species which occur at or near the base. This

rather peculiar condition is probably due to the fact that this

side of the island gets very little of the moisture which is brought

to the opposite side in the form of fog banks. Mr. R.. H. Beck

visited the south side of this mountain, in his search for tortoises,

and reported it to be less sterile than the west side A small lat¬

eral crater, which occurs on the south side, appeared to be heav¬

ily covered with vegetation, when seen from the west side of the

mountain.

282 Wisconsin Academy of Sciences , Arts , and Letters .

Cpwley Bay.

Cowley Bay is located on the east side of the island near its

center. The shores around the bay and along the adjacent coast

north of here, are composed of low cliffs of pumice and occa¬

sional pebble and sand beaches. A plain, covered with partly

disintegrated pumice, extends inland from the shore to the base

of the mountain, a distance of about half a mile. The east side

of the mountain rises rather steeply to 2,000 ft., and is covered

to this elevation with partly disintegrated pumice, similar to

that on the plain below. Occasional ridges of lava protrude

through the pumice in places so it is likely that these deposits

are not very thick. The slope is more gradual above 2,000 ft.

and continues so to within a few hundred feet of the rim of the

crater. The sides of the crater’s rim are quite steep. Apparent¬

ly all of the mountain side above an elevation of 2,000 ft. is cov¬

ered with basaltic lava which has become partly broken down in¬

to soil through which lava boulders project forming a rather

rough surface in most places. The side of the mountain, a short

distance south of the bay, is covered with deposits of recent lava

to a considerable elevation. The west side of the mountain was

not visited, but it was noticed while sailing past this side of the

island, that the vegetation was quite heavy here so it is likely

that this side of the mountain is covered with lava and not pum¬

ice.

There are but few halophytic plants in this vicinity, possibly

due to the steep and unstable nature of the shores. A few

bushes of Laguncularia racemosa were noticed, aud a small grove

of trees of Rhizophora Mangle were noticed a mile or two fur¬

ther south.

The region near the shore is almost bare of vegetation in many

places, and with the exception of the more recent beds of lava

on some of the other islands, it is the most sterile place botani-

cally that we visited. What little vegetation there is here is

very much scattered and consists largely of low Bursera and

Croton bushes, and bushes of Cordia lutea, Discaria pauciflora,

Dodonaea viscosa, Lipochaeta laricifolia, Maytenus obovata and

Scalesia gummifera, all of which are rather stunted except when

they occur in protected places. We visited this place during the

dry season but it is not likely that conditions would be much

more inviting during the rainy season, because very few remains

of annual plants were found.

Stewart — Botanical Conditions on the Galapagos Islamds. 283

The lower slopes of the mountain are more heavily covered

with vegetation than is the plain just mentioned, but even here

there are often areas of a considerable size which have scarcely

any vegetation on them. The species which occur on the plain

also occur on the side of the mountain in greater number, and

many of them that were stunted on the lower part, reach their

normal size around an elevation of 1,000 ft. The increase in the

humidity1 of the atmosphere is shown at this elevation, and

above, by the large amount of Usnea longissima, and other lichens,

which cover the vegetation to such an extent as to give it a gray¬

ish appearance. A number of mesophytic plants were first seen

around 1,300 ft. elevation, the most noticeable of which were:

Pisonia floribnda, Psidium galapageium, and Scalesia micro-

cephala, the last of which forms a zone on this side of the moun¬

tain to an elevation of 1,650 ft. There are also a great many

Bursera trees at an elevation of 1,200 ft. and above. Below this

they were few in number. Ferns begin to appear at a slightly

higher elevation.

There is an abrupt change in the appearance of the vegetation

at an elevation of 2,000 ft. The character of the soil also

changes here from pumice to disintegrated lava mixed with vege¬

table mold, so that the change in the vegetation is due more to an

increase in the number and size of plants than to a sudden

change of forms. There are heavy forests here made up of

trees of Bursera graveolens, Pisonia floribunda, Psidium gala¬

pageium, and Zanthoxylum Fagara as well as many species of

bushes the most of which were found at a lower elevation al¬

though usually smaller in size. The trees are often covered with

vines/ of Cissampelos Pareira, and fruticose lichens continue to be

abundant. Ferns are also abundant, the common species being

Adianaum concinnum, Doryopteris pedata, Polypodium pectina-

tum, and Trachypteris pinnata. Small specimens of Opuntia

myriacantha were seen at this elevation and they continue to

within a few hundred feet of the top of the mountain according

to Mr. R. H. Beck, who visited this region.

The sides of this mountain were not explored botanically above

an elevation of 2,100 ft. From the top of a tree at this elevation,

the whole of the country beyond could be seen. The character

of the vegetation did not seem to change until the steep slope,

below the rim of the crater is reached. Just below the rim, in

several places, there were light green areas which, according to

284 Wisconsin Academy of Sciences , Arts , and Letters.

Mr. Beck, are covered with a tangled growth of bushes and morn¬

ing glory vines.

The conditions on this side of the mountain are such that it

is very difficult to determine the extent in elevation of the botan¬

ical regions, the nature of the soil being such as to cause xerophy-

tic plants to predominate higher up than would probably be the

case if the lower part of the mountain was covered with a more

suitable soil. The transition region evidently begins around an

elevation of 1,200 ft. and it probably extends up to within about

500 ft. in elevation, from the top of the mountain.

Iguana Cove.

Iguana Cove is a slight identation in the shoreline on the

southwest side of the island. It is somewhat protected from the

direct action of the swell, but owing to its small size, it does not

afford an anchorage inside. The anchorage is just outside the

cove, but owing to the fact that there are jagged rocks projecting

from the water a short distance from it, on which the swell

breaks heavily, it is dangerous to anchor here except in calm

weather. The shores are precipitous in this vicinity, being made

up of bluffs, which in places rise to a height of 200 ft. These

tall bluffs do not come down to the shore, however, except in one

place ; in other places there is a low flat plain intervening be¬

tween them and the shore. In the vicinity of Christopher Point, •

just north of the cove, the shores are made up of low lava cliffs,

and the country back of them is covered with rather recent lava

on which there is apparently very little vegetation. In the im¬

mediate vicinity of Christopher Point there are many small cra¬

ters and blowholes which rise fifty or more feet in height, and

which give the surrounding country a weird and grotesque ap¬

pearance. South of the cove the shores are made up of low

cliffs with occasional shelving beaches of gravel and sand.

The mountain northeast of Iguana Cove is a broad flat-topped

crater which probably rises to a height of 5,000 or more feet. As

the weather was very bad when this place was visited, no attempt

was made to reach the top of the mountain. The sides are very

steep here, and are covered with a considerable amount of soil,

composed of disintegrated lava and vegetable mold, which sup¬

ports a heavy growth of vegetation. The north side of the moun¬

tain is not so steep and is covered with beds of barren lava in

Stewart— Botanical Conditions ,c m the Galapagos Islands . 285

which there are occasional islands of older lava which are cov¬

ered with xerophytie vegetation.

The botanical conditions in the vicinity of Iguana Cove are

rather unique, as it is the only place on the islands where an ex¬

tensive mesophytic vegetation occurs near sea level. It is very

likely that the steep slope has something to do with this, as no

such conditions are found a short distance south of the cove

where the slope is more gradual.

The halophytic flora is of no consequence here due probably

to the steep shores. There are mangrove swamps, however,

along the north shore of this part of the island between Christo¬

pher Point and Elizabeth Bay, and on the south shore between

Essex Point and Cape Rose.

The flat area at the base of the cliffs, just south of the cove, is

covered with a heavy growth of bushes consisting for the most

part of: Cordia ITookeriana, Cryptocarpus pyriformis, Tourne-

fortia rufo-sericea, and Zanthoxylum Fagara. This condition

continues for some distance down the coast was found out by

some members of the party who attempted to come overland.

The sides of the cliffs just back of the cove, are pendicular so

that it is difficult to scale them except, in a few places where they

are somewhat shelving. In such places there are a few trees

and bushes, and a considerable number of herbaceous plants and

ferns. Above the top of the cliff there is a heavy growth of

vegetation consisting of trees of Bursera graveolens, Pisonia

floribunda, Scalesia Cordata, and Zanthoxylum Fagara, the last

of which is usually heavily covered with Phoradendron Hens-

lovii. There are usually thick tangled masses of bushes which

are heavily overgrown with such vines as Cissampelos Pareira

and Ipomoea Bona-nox. Many ferns, both epiphytic and terres¬

trial, occur here.

Owing to the rainy weather, while we were at this place, no

plants were collected above 500 ft. The conditions at this ele¬

vation were about the same as those near the tops of the cliffs

above the cove, except that there were occasional open places in

the vegetation, which were covered with ferns and grasses.

These areas get larger a little higher up. Mr. R. E. Snodgrass

climbed about half way up the side of this mountain when he

visited these islands several years ago. He has told me that ap¬

parently the rest of the way up, the vegetation is made up of

dense fern brakes with irregular rows of shrubs running through

286 Wisconsin Academy of Sciences , Arts , and Letters..

them. It was noticed from the shore that the vegetation had a

streaked appearance about half way up the side of the mountain.

It appeared to be made up of alternating light and dark bands,

and suggested that there might be flows of different kinds of lava

in this region, each of which presented conditions peculiarly

adapted for the growth of certain species of plants.

The country around the top of the mountain was examined

through a field glass on a clear day later in the season as we

were sailing past this part of the island. The vegetation ap¬

peared to be smaller than lower down and it was rather grayish

in color instead of dark green. It is very likely that the upper

part of this mountain receives less moisture than does the middle

part. The upper part of this mountain could be plainly seen

from the top of the mountain at Villamil when we visited there.

The lower part of the mountain, however, was entirely hidden

by the fog at this time. It was also noticed that much of the soil

on top of the Villamil mountain was dry while lower down it

was wet. The fog banks apparently hang low when they strike

the islands.

Turtle Cove.

Turtle Cove is on the south side of the island about six miles;

west of Villamil. The coast in this vicinity is low and rocky

with occasional sand beaches, while back of the coast the coun¬

try is low and covered with beds of basaltic lava for a consider¬

able distance inland. There are springs of comparatively fresh

water and pools of strongly brackish water in the vicinity of the

shore and farther inland.

There were large trees of Avicennia officinalis on the sand

beach where we landed, back of which there is a swampy area

covered with a dense growth of Conocarpus erectus and trees of

Rhizophora Mangle. Rhizophora occurs for some distance in¬

land, surrounding the pools of brackish water. It also occurs in

isolated patches on the open coast, but owing to the fact that the

surf breaks heavily here at times, none of these are extensive.

Thickets of Laguncularia racemosa are also to be found in places;

in the vicinity of the shore and there are quite a number of small

trees of Hibiscus tiliaeeus, and bushes of Tournefortia rufo-seri-

cea.

The vegetation on the inland country consists of plants usu-

aly found in the dry regions except that there is an unusually

Stewart— Botanical Conditions pn the Galapagos Islands. 287

large number of trees of Hippomane Mancinella. Other com¬

mon trees are those of Bursera graveolens and Opuntia myria-

cantha. The country adjacent to the coast, just east of this

place, is covered with dense thickets of Cryptocarpus pyriformi&

apparently to the exclusion of all other vegetation of any size.

West of here, in the vicinity of Cape Rose, there are extensive de¬

posits of volcanic cinders on which the vegetation is very open,

probably due to the fact that the lava has disintegrated but lit¬

tle. What few plants that do occur here are for the most part,

the ones that are commonly found in the dry regions of these is¬

lands.

All of the country which lies between the mountain at Iguana

Cove, and the one northwest of Villamil is low, probably in no<

place exceeding an elevation of 200 ft.

Vilamil.

Villamil is on the south side of the island about seven miles

northwest of Brattle Island. A settlement of about one hundred

and fifty people was established here some years ago, by Mr. An¬

tonio Gil of Guayaquil, Ecuador. A considerable industry in

hides, molasses, and sulphur is carried on, the products of which:

are sent to Guayaquil by means of a small vessel which makes

periodic trips to the mainland. A part of the settlement is lo¬

cated near the shore, but the most of it is about twelve miles in¬

land, at an elevation of 1,300 ft., where there are plantations of

sugarcane, bananas, and other tropical fruits and vegetables.

Villamil Bay is surrounded by low beds of basaltic lava, but

west of the settlement on the open coast, there are extensive sand

beaches. These are continuous with a broad sand-flat just back

of them, which extends back for about half a mile. The country

for several miles inland is flat and is covered with beds of basaltie

lava and volcanic cinders which usually lie almost horizontally.

There are numerous crevices in the lava, in some of which there

are pools and springs of nearly fresh water. Owing to the low

elevation of this part of the island, these springs usually occur

only a few feet below the level of the ground. There is a consid¬

erable amount of precipitation on the upper part of this island,

in the form of fog and rain. There is not sufficient soil to retain

this water, however, so it percolates through the lava and comes

out again near sea level. On this account the water is usually"

slightly brackish even at a considerable distance inland.

288 Wisconsin Academy of Sciences , Arts , and Letters.

The flat country extends inland for about four miles, to the

base of the mountain. Above this the slope is very gradual to

an elevation of 500 ft. So far as could be observed, this side of

the mountain is covered with volcanic cinder, which has become

slightly disintegrated and mixed with vegetable mold, forming a

scant soil. The slope is less gradual above 500 ft. and continues

so to an elevation of 2,400 ft. This part of the mountain side is

rather rolling and slightly terraced in places. There is suffi¬

cient soil over the upper part of the mountain to completely

cover the lava except on ridges and other exposed places. The

slope of the mountain side is quite abrupt above 2,400 ft. to the

rim of the crater, which has an elevation of 3,150 ft.

As near as could be estimated the crater is about seven miles

long and four miles broad, the greatest diameter being approxi¬

mately east and west. The floor of the crater is flat at its east¬

ern end and is filled with numerous crevices through some of

which vapors issue periodically. There is a prominent ridge

near the center of the crater which rises gradually in height un¬

til at its west end it is nearly as high as the rim. A small active

sulphur volcano is situated on the south side of this ridge, and

still another larger one at its west end around both of which

there are deposits of sulphur. It is from this place that the in¬

habitants obtain the sulphur which they export to Ecuador.

Small swamps of Khizophora Mangle occur in places around

Villamil Bay, and trees of Avicennia officinalis, and bushes and

trees of Laguncularia racemosa are to be found in several places

near the coast. Quite a grove of these occurs near the settle¬

ment. Along the sand beaches west of the bay, there are many

small halophytic and semihalophytic plants such as : Cryptocar¬

pus pyriformis, Heliotropium curassavicum, Ipomoea Pes-caprae,

and Scaevola Plumieri. The sand flat, back of the beach, is cov¬

ered with a dense growth of Sporobolus virginicus in which there

are small groves of Hippomane Mancinella trees, and bushes of

Cryptocarpus pyriformis. In places around the edges of the

sand-flat there are thickets of Conocarpus erectus, some of which

form trees twenty-five or more feet high.

There is a low area of limited extent about a mile west of Villa-

mil in which the soil is kept moist by the water which comes

down through the lava from the interior. There are quite a

number of mesophytic plants here. Vines of Argyreia tiliae-

folia and Cissampelos Pareira cover the rocks in places and there

Stewart — Botanical Conditions on the Galapagos Islands. 289

is a small grove of trees of Anona glabra and ferns. The in¬

habitants have planted a garden in this place which has been

quite successful as bananas and other tropical plants grow there.

The change from xerophytic to the mesophytic type of vegeta¬

tion is very abrupt here, as such pronounced xerophytes as Lan-

tana peduncularis, Opuntia myriacantha, and Prosopis dulcis

are found growing only a few feet away from the mesophytic

plants enumerated above. There are several low marshy areas,

filled with brackish water, in the vicinity of the settlement, in

which there is a heavy growth of Eleocharis mutata. The stems

of this plant are used by the inhabitants for making mats. The

higher land between these marshes is covered with low and rath¬

er open forests consisting of trees of Acacia macracantha, Bur-

sera graveolens, Hippomane Mancinella, and Opuntia myria¬

cantha, among which there are bushes of Chiococca alba, Clero-

dendron molle, Cordia lutea, Gossypium barbadense, and bushes

and small trees of Zanthoxylum Fagara on which Phoradendron

Henslovii is often found. In many of the lava crevices, which

are deep enough to reach the ground water, there are large

bunches of Cyperus ligularis.

On the broad plain some distance inland, there are beds of ba¬

saltic lava and volcanic cinder of a considerable width. The

basaltic lava is often heavily covered with vegetation and in one

place an entire flow is covered with a forest of Opuntia myria¬

cantha trees, underneath which there are low dense thickets of

Euphorbia viminea and occasional bushes of Acacia macracan

tha. Cyperus Mutisii was found growing abundantly in the

smaller crevices of the lava in this area. The vegetation on the

cinder deposits, however, is very open and consists mostly of oc¬

casional bushes, or small clumps of bushes, of Clerodendron

molle, Erigeron tenuifolius, Lippia rosmarinifolia, and Scalesia

gummifera on many of which there was a dense growth of vines

of Cardiospermum galapageium, and Passiflora subrosa. Be¬

tween the bushes the ground is often bare for some distance.

In one place, several miles inland, there is a low area which

had the general appearance of having been filled with water

at some time. There is much more soil here than in any place

in this vicinity. There are pools here which seem to contain wa¬

ter the most of the time, around which Cyperus laevigatus and

Sporobolus virginicus grow. Groves of Hippomane Mancinella

grow in this area, in the shade of which there are bushes of Cae-

19— S. A.

290 Wisconsin Academy of Sciences , Arts , and Letters.

salpina Bonducella, Cryptocarpus pyriformis, Discaria pauci-

flora, Scalesia gummifera, and Solanum verbascifolium. In the

more open places in this area there were large bunches of Pani-

cum fasciculatum and other herbaceous plants. On barren lava

beds and on exposed ridges in this vicinity, Cereus sclerocarpus

was the only plant that grew to any considerable size.

A change takes place in the vegetation between an elevation

of 100 and 200 ft. where many of the plants common below dis¬

appear, the most common of which are: Acacia macracantha,

Castela galapageia, Cereus sclerocarpus, Discaria pauciflora,

Euphorbia viminea, and Waltheria reticulata, while such promi¬

nent woodland plants as Pisonia floribunda, Psidium gala-

pageium, and Scalesia cordata begin to appear along with ferns

and other plants, which are found abundantly higher up. There

is a general thickening above an elevation of 200 ft. and f ruticose

lichens are very abundant on trees and bushes.

Sapindus saponaria was first seen around 250 ft. elevation.

There are only occasional trees of this species at this elevation,

the dense Saponaria forests not beginning for another hundred

feet or so in elevation. Scalesia cordata also increases in abun¬

dance so that the forest trees throughout the moist region con¬

sist mostly of these two species. There is a heavy growth of

bushes in these forests, increasing with the elevation, which

consist largely of the following species. Clerodendron molle,

Croton Scouleri var. grandifolius, Erigeron tenuifolius, Psycho-

tria rufipes, Tournefortia psilostachya, T. pubescens, and T.

rufo-sericea. There are many ferns both terrestrial and epi¬

phytic, the common epiphytic species being : Polypodium lanceo-

latum, and P. lepidopteris, while on the higher branches of many

of the trees there are large bunches of Lycopodium dichotomum.

Other common epiphytes in this region are Ionopsis utricular-

iodes, Peperomia galapagensis, P. Stewarti, and Tillandsia in-

sularis. There are a large number of trees of Hippomane Man-

cinella in the forests at an elevation of 600 ft. but none were

found below this, except near sea level.

A considerable amount of the forest has been cleared away be¬

tween 600 and 1,300 ft. elevation. Much of this area has since

been neglected and has grown up in bushes of Tournefortia rufo-

sericea which are heavily covered in places with vines of Argy-

reia tiliaefolia, and Ipomoea Bona-nox. There is also usually a

heavy growth of grass in between the bushes, and brakes of Pter-

Stewart — Botanical Conditions fin the Galapagos Islands. 291

is aquilina var. esculenta are not uncommon. The forest bor¬

dering the cleared area seems to be made up mostly of the same

forms found around an elevation of 600 ft., where the cleared

area begins, but the lower part of it was not carefully explored.

The vegetation becomes much thinner in the uncleared areas

above 1,200 ft. elevation and with the exception of an occasional

tree of Sapindus saponaria, there are no trees of large size. The

country is covered with open woodland made up largely of small

trees and bushes of Croton Seouleri var. grandifolius, Scalesia

cordata, Solanum verbascifolium, Tournefortia rufo-sericea,

IJrera alceaefolia, Zanthoxylum Fagara, many epiphytic plants

and ferns. There are many park-like areas in the woodland

which are covered with grasses. The trees become smaller and

more scattered to an elevation of 1,500 ft., where they end rather

abruptly.

The side of the mountain above 1,500 ft. elevation is somewhat

rolling and is covered with grassland on which large numbers of

cattle graze, which are slaughtered by the inhabitants of the is¬

land for their hides. Paspalum conjugatum is the principal

species of grass found in this region. This condition continues

to an elevation of 2,400 ft, above which there is a decrease in the

amount of grass and a large increase in the fern flora. Small

tree ferns, Hemitelia multiflora, and other large species of ferns

are common from here to the top of the mountain.

There is a great difference in the vegetation of the outer and

inner sides of the southern rim of the crater, where the most of

the collecting around the top of the mountain was done. The

outside of the rim at this place is mostly covered with small vege¬

tation consisting of ferns, club-mosses, and small herbaceous;

forms, all of which lie close to the ground, and it is only in places

which are protected from the wind that plants of any size are to>

be found. Just over the rim of the crater, however, there is a

considerable growth of bushes of Duranta repens, Erigeron lan-

cifolius var. glabriusculus, Solanum verbascifolium, Zanthoxy¬

lum Fagara, and other bushes. Hemitelia multiflora also occurs

here in large numbers and such other ferns as Asplenium Serra,

Dryopteris parasitica, Elaphoglossum muscosum, Polypodium

aureum, and Polystiehum aculeatum abound.

A gradual change from a mesophytic to a xerophytic vegeta¬

tion can be readily noticed as one travels around the southern rim

of the crater towards the northwest side, but as our time was lim¬

ited when this region was visited, no collections were made.

292 Wisconsin Academy of Sciences , Arts , and Letters.

The floor of the crater is 400 ft. below the rim, and was exam¬

ined near the west end, in the vicinity of one of the active vol¬

canoes. Here were found patches of Sporobolus indicus cover¬

ing considerable areas in places while other areas were covered

with Gnaphalium luteo-album, the gray color of which caused

them to stand out prominently when the floor was viewed from

the rim. There were also occasional specimens of an arbores¬

cent species of Cereus, and low stunted specimens of Opuntia

myriaeantha. An occasional tree of Zanthoxylum Fagara was

seen, usually close to the crater’s wall. There is a considerable

growth of stunted bushes in places consisting mostly of : Clero-

dendron molle, Dodonaea viscosa, Euphorbia equisetiformis, and

Lipochaeta laricifolia. There were brakes of Pteris aquilina var.

esculenta in one place, but outside of this, ferns are few at this

end of the crater.

The northwest side of the mountain seems to be covered with

grassland, which is much drier than are the south and southeast

sides. At least it appeared to be as far down on this side as we

could see from the top of the mountain. The soil on the rim of

the crater was also much dryer on this side than it was on the

southeast side.

The dry region is confined largely to the broad flat plain at

the base of the mountain at this place. As near as could be de¬

termined it does not extend above an elevation of 150 ft. Above

this many of the mesophytic forms appear, and there is a large

amount of fruticose lichen on the vegetation, indicating a greater

humidity. The transition region forms a narrow belt along the

base of the mountain, the upper limits of which reach to an ele¬

vation of about 350 ft. The lower part of the moist region is

covered with dense forests of Saponaria, and Scalesia trees,

while on the upper part the vegetation consists mostly of bushes,

and small trees with open spaces between them at intervals.

Near the upper limit of this region, around an elevation of 1,500

ft., there are only low bushes. The grassy region extends from

1,500 ft. nearly to the top of the mountain.

Barrington Island.

Barrington is situated ten miles southeast of Indefatigable,

and twenty-six miles west of Chatham Island. It is one of the

smaller islands of the group, and it is of low altitude the most

Stewart — Botanical Conditions on the Galapagos Islands. 293

of it not reaching an elevation of over 350 ft. There is a hill

near the northwest side, however, which attains an elevation of

650 ft. This hill ends abruptly at the top of a tall bluff: which

drops almost straight downward into the sea. The shores of the

island are made up of low lava cliffs for the most part, but there

is a small bay on the northeast side, wThich is surrounded by

sand-beaches. This bay is sheltered by a small islet and a reef,

Although this bay can not be entered by vessels, it nevertheless

affords an excellent landing place for boats.

Topographically the island is made up mostly of alternating

ridges and valleys which have a general trend towards the south¬

east. The ridges, in a general way, are 100 ft. higher than the

valleys, and are covered with tumbled masses of lava. The val¬

leys, on the other hand, usually have a considerable amount of

soil in them, the most of which has probably been formed on the

sides of the ridges and washed down. The soil varies from a

light brown to an ochre color, and is very light in texture. The

lava all seems to be basaltic in character and is evidently quite

old as it has become stained to a redish-brown color.

The only plants found on the sand-beaches surrounding the

bay, were bushes of Cryptocarpus pyriformis, and mat-like

growths of Sesuvium Edmonstonei both of which are not exclu¬

sively halophytic in their habits. The shores on other parts of

the island are too steep to support halophytes. A short distance

inland from the beach there are low thickets of Discaria, and

Maytenus bushes.

Owing to a low altitude, all of the vegetation in the interior

of the island is very xerophytic in character, and about the only

noticeable change that takes place in the vegetation towards the

higher parts is the greater abundance of fruticose lichens. The

most noticeable plants are the large trees of Opuntia myriacan-

tha which grow in great numbers over the most of the island.

Small trees of Bursera graveolens also occur, much infested with

lichens. There was a fair growth of bushes in most places, con¬

sisting for the most part of such species as : Cordia lutea, Gor¬

ton Scouleri, Gossypium barbadense, Lantana peduncularis, Tel-

anthera echinocephala and Scalesia Helleri, the last one of which

was the only conspicuous green plant to be found on the interior

of the island at the times we visited it. In the valleys between

the ridges there were small areas which are covered with a

growth of Euphorbia viminea forma barringtonensis. In other

294 Wisconsin Academy of Sciences, Arts, and Letters.

places in these valleys where the soil is loose there is an abun¬

dance of Coldenia fusca.

This island was visited during the months of July and Octo¬

ber, in consequence of which the annual vegetation, which comes

on during the rainy season, was missed entirely. Very few re¬

mains of such plants were found on either of our visits as goats

have been introduced upon this island during the last few years

which eat up all of the edible vegetation as fast as it grows.

Even the trunks of the large Opuntia trees do not escape their

ravages.

Bindloe Island.

This island is the largest of the group of three which lie some

distance north of the main part of the archipelago. It is eight

miles long, six and one-half miles broad, and its highest part at¬

tains an elevation of 800 ft. according to the chart issued by the

Hydrographic Office. When seen from a distance the island ap¬

pears to be made up of numerous small peaks which vary in ele¬

vation. The greater part of the island is covered with beds of

recent lava which consist mostly of volcanic cinder. The whole

of the north side, along which we sailed, is covered with such de¬

posits, on which there are occasional exposures of older lava

which support a considerable amount of vegetation.

We anchored on the northeast side of the island near where

a broad strip of country, covered with deposits of tufa, extends

down to the shore. This area is covered with vegetation, appar¬

ently the largest continuous body of such on the island.

A few small green patches, evidently halophytic plants of

of some kind, were noticed along the north shore of the island

as we were sailing past it. With this exception, no halophytes

were seen, as the shores are too steep in most places to support

them.

The vegetation is arranged in irregular clumps in the vicinity

of the shore with broad open lanes between them. The vegeta¬

tion here is made up mostly of low bushes of Euphorbia amplexi-

caule, E. articulata, Castela galapageia, and low thickets of

Opuntia. This open arrangement disappears inland and the

country is heavily covered with xerophytic vegetation. The

only trees found here are those of Bursera graveolens which

grow quite large considering the very dry conditions which pre¬

vail. They are sometimes covered with vines of Ipomoea Habel-

Stewart — Botanical Conditions pn the Galapagos Islands. 295

iana, the only place on the islands where this species assumes the

climbing habit in such a pronounced way.

The edges of recent flows of lava are often bordered with

bushes of Cordia lutea, and Waltheria reticulata forma interme¬

dia, both of which occur in other places but less abundantly.

No collecting was done on the upper part of the island, but

according to Mr. Beck, who visited this part, the country is cov¬

ered with beds of recent lava which .have but little vegetation on

them. There are a few moist places, in the vicinity of steam-

vents, in this region, around which such ferns as Ceropteris tar-

tarea, Nephrolepis biserrata, and Polypodium squamatum grow

to some extent. The whole of the island may be included in the

dry region.

Brattle Island.

Brattle is a small island, that is situated about four miles off

the south side of Albemarle Island near its eastern end. It is a

semilunar in general outline and is the remains of an old tufa

crater the south and west sides of which have been eroded away,

except in two places, where there are small islets. The top of

the island is 275 ft. above sea level, and the sides are very steep

and much cut up with gullies and ravines. Owing to the steep

nature of the shores landing is difficult, and can only be done

with safety on the north side, when the water is comparatively

still.

The greater part of the surface of the island is bare of vegeta¬

tion, a condition that is probably due to the steep sides and the

loose soil, which is composed of volcanic ashes and small bits of

lava loosely cemented together.

The most common plant on the island is a low bush which is

covered with thick, succulent leaves, and which forms thickets

around the top in various places. This plant was neither in

flower or fruit at the time the island was visited so it could not

be identified with certainty. Bushes of Croton Scouleri occur

along the sides to some extent, but they are stunted and the

leaves are smaller here than is usually the case with this species.

Three species of herbaceous plants: Coldenia fusca, Ipomoea

Kinbergi, and Tribulus cistoides were found at the top, as well

as the remains of several grasses. Two lichens, Ramalina com-

planata and Rocella peruensis are also found.

296 Wisconsin Academy of Sciences, Arts, and Letters.

Charles Island.

With the exception of Hood, this is the most southerly island

in the group. It is located about thirty -seven miles south of In¬

defatigable Island. The island is ten miles long and eight miles

broad. It reaches an elevation of 1,780 ft. at its highest point.

Geologically it is probably one of the oldest islands in the group,

and volcanic activity upon it has evidently long since ceased.

There are no deposits of even comparatively recent volcanic ma¬

terial upon it.

In approaching the island from the south, one is impressed

with the number of large craters on it. Fourteen of these were

counted, seven of which were larger than the rest. The tops of

the most of the craters are evenly rounded, and it was found out

later that the southeast sides of many of them were broken down.

The slope is quite gradual from the shore to the central region,

on all sides but the east. This side was not visited, but in sail¬

ing along the shore, the slope appeared to be rather steep, and

was covered with xerophytic vegetation among which were a

large number of Cereus.

There is a fair amount of soil in most places, composed of vol¬

canic ashes and bits of lava. There are exposures of lava, how¬

ever, on which there is but little soil, but they are less common

than on other islands visited. The central part of the island is

covered with a plateau, several miles square, which has an aver¬

age elevation of 1,000 ft. Several large tufa craters are located

on the plateau^ which usually rise 500-800 ft. above it. Springs

occur around the base of one of these, and a considerable amount

of water is afforded by one of them. Such domesticated animals

as : Cattle, hogs, goats, cats, and dogs have been introduced upon

the island. The inhabitants from Chatham island often come

here to dry beef for the use of the laborers on that island.

Black Beach Road.

Black Beach Hoad is located on the west side of the island and

was the port for the settlement which was located on this island

many years ago. A good trail leads inland from here so the cen¬

tral region is more accessible than on the most of the other un¬

inhabited islands.

The shores are low and rocky in the vicinity of Black Beach

Road, against which the surf breaks heavily at times. On this

Stewart — Botanical Conditions on the Galapagos Islands, 297

account there are no halophytes to speak of except a small bunch

of rather stunted mangroves a short distance south of the land¬

ing place.

The region north and east of this place is covered with a fair

amount of ashy soil through which the lava seldom appears.

South of here, however, there are exposures of lava, covered for

the most part with Croton bushes. Just back of the landing

place there is a flat area covered with bushes and small trees of

Maytenus obovata and Prosopis dulcis. Another small area

occurs a few hndred yards north of the landing place near the

coast, which is covered with tumbled masses of lava among which

Cereus galapagensis, Lecocarpus pinnatifidus, Mentzelia aspera,.

and Scalesia decurrens grow.

The larger vegetation to an elevation of 450 ft. consists of trees

of Bursera graveolens, and Opuntia galapageia. In the vicin¬

ity of the shore there are also trees of Cereus galapagensis. The

vegetation is all rather open but there are a considerable number

of bushes of Cordia lutea, Croton Scouleri var. Macraei, Lantana

peduncularis, Maytenus obovata, Gossypium barbadense, and

Vallesia pubescens. Acacia macracantha and Prosopis dulcis

also occur in this region to some extent but they assume the size

of trees around 450 ft. There are remains of an old settlement

at this elevation which is marked by a grove of Geoffroea striata

and other trees, as well as by a few other domesticated plants of

smaller size. There was evidently a spring of water here at

some former time but it was dry at the times this place was vis¬

ited.

A decided change takes place in the vegetation above an eleva¬

tion of 450 ft. For possibly the first 200 ft. there are large

bunches of bushes of Clerodendron molle, in between which are

grasses and smaller plants. This is succeeded above by more

open country on which there are occasional Bursera trees and

bushes, the most common of which are, Capraria biflora, and Lip-

ochaeta laricifolia. Perennial grasses grow between the bushes,,

to which a considerable number of annual forms are added dur¬

ing the rainy season.

The plateau region, around an elevation of 1,000 ft., is covered

with stretches of rather open woodland, and meadow. The

woodland usually occurs where the lava is exposed or reaches

nearly to the surface of the ground. In these areas trees of Scal¬

esia pedunculata are common but they do not grow to as large a

298 Wisconsin Academy of Sciences , Arts , and Letters.

size as they do on some of the other islands where this species oc¬

curs. Other trees in the woodland besides those that have evi¬

dently been introduced are: Pisonia floribunda, and Zanthoxy-

lum Fagara, the last one of which is often heavily covered with

Phoradendron Henslovii, as often happens when this tree grows

where there is a considerable amount of moisture. Common

bushes in the woodland are: Capraria biflora, Croton Scouleri

varieties brevifolius and grandifolius, Erigeron tenuifolius, Psy-

chotria rufipes, Tournefortia psilostachya, and T. rufo-sericea.

The soil is ashy in the meadows, with small fragments of lava

scattered through it. Grasses and herbaceous plants occur here,

the common species being: Aristida subspicata, Eleusine indica,

Eragrostis ciliaris, Acalypha parvula, Lippia canescens, Malvas-

trum americanum, Plumbago scandens, and Stachytarpheta

dichotoma. Many evidences of former habitation appear in the

flora throughout the plateau region as such introduced species as

Ambrosia artemisiaefolia, Bixa Orellana, Datura Tatula, Inga

edulis, Spondians purpurea, orange, lemon, and lime trees grow

there in greater or less abundance. The lime trees are the most

common of these, and there are areas of considerable size which

are covered with them. They seem to spread mostly in the open

meadows as there are but few of them in the woodland. Prob¬

ably in time they will cover all of the open country in the plateau

region. The limes and lemons are of good quality and many

• tons of them rot on the ground each year.

The main crater rises to a height of 1,780 ft., and is covered

on the outside with a heavy growth of bushes of Lipochaeta lari-

cifolia, to within about 400 ft. of the top. This condition is

found on all sides but the south and southeast. The bushes

gradually disappear towards the south side of the crater and

their place is taken by a heavy growth of Stachytarpheta dicho¬

toma. Above an elevation of 1,450 ft. the outside of the crater is

covered with low bushes of Capraria biflora, occasional bushes

of Tournefortia rufo-sericea, grasses and other herbaceous plants

the last of which are dried up during the greater part of the

year, giving this part of the mountain a very barren appearance

the most of the time. One will be greatly surprised if he should

climb the west side of this crater during the dry season. On this

side Opuntia galapageia grows to an elevation of 1,300 ft. The

other plants which occur here and above are of a xerophytie

type and continue so to the top of the mountain. Upon going

Stewart — Botanical Conditions on the Galapagos Islands. 299

over the rim, into the interior of the crater this condition changes

and the plants which occur, there are decidedly mesophytic

Small trees of Acnistus ellipticus grow there which are covered

with such epiphytes as: Lycopodium taxifolium, Polypodium

lanceolatum, Peperomia ramulosa and leafy Hepatics. Perns

and herbaceous plants are also common in this vicinity. The

sudden change in the character of the vegetation within such a

short distance is due to the fact that the moist winds strike the

inner side of the crater and keep the vegetation damp the most

of the time, while they pass directly over the top so that the mois¬

ture seldom descends far upon the leeward side. The inside of

the crater is covered, a short distance below the top, with bushes

and occasional small trees of Zanthoxylum Fagara all of which

bear numerous epiphytic plants. Ferns are common in the crev¬

ices of the lava. The floor of this crater is covered with trees

of Scalesia pedunculata and bushes.

There are several other craters in the upper regions which do

not reach as great an elevation as the one described above. For

the most part, they are covered with a heavy growth of lime

trees and bushes on their leeward sides while the windward sides

are covered with low bushes and herbaceous plants above an ele¬

vation of 1,300 ft.

It seems impossible to make out distinct botanical regions her?

as can be done on some of the other larger and higher islands of

the group. For some reason or other, this island apparently does

not receive as much moisture as do the other islands of similar

elevation. In consequence of this the upper part, including the

plateau, is covered with a mixture of both xerophytic and meso¬

phytic forms, the last of which, however, are more abundant than

they commonly are in the transition regions on the other islands.

There are a few places in which there is a suggestion of a moist

region but these are very limited in extent, dependent upon some

very local condition or conditions. The country below an eleva¬

tion of 1,000 ft. can be divided into open woodland below and

bushy country above, the line of separation between the two be¬

ing at about 450 ft. elevation. /

Cormorant Bay.

Cormorant Bay is an open sheet of water situated on the north

side of the island a short distance east of Post Office Bay. There

are several sand beaches along the shore here on which Batis

3-00 Wisconsin Academy of Sciences , Arts , and Letters.

rrxaritima, Lycinm sp., Scaevola Plumieri, Sesuvium Edmon-

stonei, and S. portnlacastrnm grow quite commonly, while at the

west end of the bay there is a small swamp of Rhizophora

Mangle. A short distance back of the bay there is a small lake,

the water in which has become saturated with salt and a layer

has crystalized out which is thick enough in places to support

one’s weight. A number of trees of Avicennia officinalis grow

in the water on the edge of this lake. There is a heavy growth

of pneumatophores and mats of Sesuvium Edmonstonei in the

water beneath these trees. There are some exposures of lava

in this vicinity on which there is little vegetation besides Cacti

and a few bushes. Around the base of a small crater, near the

east end of the bay, there are bushes of Acacia macracantha, Cor-

dia lutea, Cryptocarpus pyriformis, and Scalesia villosa. There

is another small crater, about a half mile inland, the sides of

which seemed to be covered with Bursera trees and Croton

bushes.

Post-Office Bay .

Post Office Bay lies about two miles west of Cormorant Bay*

just mentioned. This bay derives its name from the fact that

the British Warship Leander placed a barrel on a post there

many years ago, in which vessels which visited this place could

deposit letters. The next vessel that called was supposed to

take them out and carry them to their next port of call. Sever¬

al of the members of the party, including the writer, mailed let¬

ters here which reached their destination some eighteen months

later.

The interior region, on the north side of the island, was visited

from this place, and was found to be much rougher than was the

region near Black Beach Road on the west side of the island.

There are several small craters on this part, the highest one vis¬

ited having an elevation of 700 ft. There are broad valleys be¬

tween the craters in some of which there were low rocky areas

which might have contained water at some time as there were

rounded boulders in them which had the appearance of having

been water-worn.

The coast, along the south side of the bay, is rocky with occa¬

sional sand beaches on which there are a few patches of Rhizo¬

phora Mangle, and trees of Avicennia officinalis. This, by the

way, is the only place on the islands where Avicennia grows so

Stew art— Botanical Conditions pn the Galapagos Islands. 301

'close to the open sea that its roots were covered with water at high

tide. There are also dense thickets of Laguncularia racemosa

in one or two places near the shore. In the sandy soil near the

shore there are thickets of Cryptocarpus pyriformis, and Dis-

caria pauciflora.

The low areas in between the craters in the interior region,

are usually covered with such a thick growth of bushes of Pro-

sopis dulcis as to render traveling difficult. In such places

Castela galapageia, Maytenus obovata, Parkinsonia aculeatat,

Yallesia pubescens, and Waltheria reticulata forma intermedia

are also found. In addition to these a considerable number

of herbaceous plants occur among which are : Abutilon cris-

pum, Aristida suspicata, Bidens refracta, Cyperus Mutisii,

and Tetramerium hispidum. The sides of the craters are rather

steep and support a more open vegetation than the valleys

surrounding them. Plants which commonly occur on the sides

of these craters are : Acacia macracantha, Chiococca alba,

Scalesia affinis, and S. villosa. The interior of some of these

craters were filled with a heavy growth of Parikinsonia acu-

leata. No ferns or other distinctly mesophytic plants were

found on this side of the island, to an elevation of 700 ft., as

high as it was explored. *

Chatham Island.

Chatham is the most eastern island of the group. It is the

fifth in size, being exceeded in this respect by Albemarle, Inde¬

fatigable, Narborough, and James Islands. It is twenty- three

miles long and about ten miles broad, the greatest diameter of

which extends northeast and southwest. Geologically the island

is probably very old as there are few evidences of recent volcanic

activity, except in the vicinity of Sappho Cove, on the west side

of the island.

The southern part of the island is most visited as there is a

settlement located several miles inland, from which has a good

wagon road leads down to Wreck Bay. This part of the island

is rather flat for some distance inland, and is covered with a con¬

siderable amount of soil through which lava boulders project.

There are several low lava hills on this part. The northeastern

and eastern parts of the island were not visited, but a general

survey of these was made from the vessel, and from higher parts

302 Wisconsin Academy of Sciences, Arts, and Letters.

in the interior of the island. The region around Terrapin Road,

and for a considerable distance south, is covered by a broad plain

which slopes gradually upward towards the southwest. There

are several steep hills on this portion of the island some of which

probably rise to a height of 500 ft. The color and general ap¬

pearance of these hills indicated that they were composed of

tufa. This part of the island is covered with forests, apparently

made up of the forms usually found on the dryer parts of these

islands.

There is a strip of country south of the wooded arear which is

covered with beds of basaltic lava on which there is apparently

little or no vegetation. This is followed on the southwest by

still another area covered with vegetation, which extends down

to within possibly three miles of Finger Point. East and south

of Finger Point the country is again covered with lava, a portion

of which is evidently volcanic cinders. There are many small

craters on the lava in this vicinity, fifty of which were counted

while sailing past this part of the island.

The country around the northeastern end of the island, and

for some distance southwest along the east coast, is apparently

very similar to that around Terrapin Road. We did not get

very close to this part of the island so no very exact observations

could be made of its close features. The country around Fresh

Water Bay, on the south side, is quite steep and is heavily covered

with dark green vegetation well down towards the shore. A

stream of water is said to enter the sea at Fresh Water Bay, but

as there is no good anchorage here it was not visited.

The interior part of the southern half of the island is a broad

plateau which varies in elevation from 900-1,600 ft. The pla¬

teau is rolling and there are numeros small ravines which have

streams of water in them during a part of the year. Small

marshes are formed in some of the lower places in this region

during the rainy season, but they quickly dry up soon after the

dry season sets in again. The main central mountain rises to a

height of 2,100 ft. and is composed of bits of volcanic cinders

and other fragmentary material all of which is very much decom¬

posed. There is no indication of a crater here, the mountain ap¬

parently being a huge pile of volcanic debris. A small crater

lake is located on this plateau but was not visited for botanical

purposes.

This island is probably better watered than any other one of

Stewart — Botanical Conditions pn the Galapagos Islands. 303

the group. There are several large springs on the plateau, the

water from which is piped down to the settlement, located at an

elevation of 900 ft. Sufficient water is obtained in this way to

supply a settlement of about four hundred and fifty people, and

to run a large sugar mill. Streams also occur on the lower parts

in the vicinity of Wreck Bay, during the rainy season, some of

which have a considerable amount of water in them at times.

The plateau is covered with a heavy coating of yellow clay-like

soil which is mixed with vegetable mold in the wooded areas.

This island was visited from Basso Point, Sapho Cove, and

Wreck Bay.

Basso Point.

Basso Point is on the west side of the island about five miles

northeast of Wreck Bay. The point shelters a broad bay which

lies southwest of it, around which there are sand beaches and

rocky shore. The sand beaches support a few halophytic plants

and in the immediate vicinity of these beaches there are thickets

of bushes of Cryptocarpus pyriformis, Discaria pauciflora, and

Maytenus obovata. The country in the interior is quite rough

and there are many exposures of lava on which there is scarcely

any vegetation. The country rises gradually to an elevation of

about 1,100 ft. above which the ascent is more abrupt, leading

up to the top of a range of hills which run parallel with the coast

in a northeasternly direction.

There are low dense forests on this part of the island which

are made up mostly of trees of Bursera graveolens and Piscidia

Erythrina, both of which are smaller than they are in the region

around Wreck Bay. There is not as much cactus here as is usu¬

ally the case in the lower regions. Cereus galapagensis occurs

to some extent near the shore but no specimens of Opuntia were

seen below an elevation of 800 ft., and they were not abundant

even there.

There is a heavy growth of bushes in the forest in most places

the most common species of which are: Croton Scoulera var.

albescens, Discaria pauciflora, Gossypium barbadense, and Lan-

tana peduncularis. The Discaria bushes grow so thickly in

places as to form impassable barriers by the interlocking of their

thorny branches.

This part of the island was not visited above an elevation of

900 ft. With the exception of a few specimens of Polypodium

304 Wisconsin Academy of Sciences , Arts, and Letters.

squamatum, all of the other plants which occur at this elevation

grow abundantly lower down. Apparently the same sort of

vegetation continues to the base of the hills mentioned above,

about 200 ft. higher. The northwest sides of these hills are cov¬

ered with forests, apparently made up largely of Scalesia trees.

The southeast sides are treeless, however, as was noticed from

the settlement near Wreck Bay, later in the season.

Sappho Cove.

Sappho Cove is also situated on the west side of the island

about four miles northeast of Basso Point. The bay is small and

almost entirely land-locked, but owing to the fact that it is very

shallow, only small vessels can anchor in it. The shores sur¬

rounding the bay are made up of basaltic lava and sand beaches

on which there are small groves of trees of Rhizophora Mangle

and thickets of Laguncularia bushes in places. A short distance

back of the beach in the vicinity of salt water pools, there are

trees of Avicennia officinalis. The sand in the vicinity of the

pools, and on the beaches, is covered in places with a heavy

growth of Batis maritima, Sesuvium Portulacastrum, and Spor-

ooblus virginicus. In several places along the open coast,

in this vicinity, the sand has been thrown up into long ridges, as

a result of the action of wind and w'aves. These ridges are cov¬

ered, on the side next to the land, with a heavy growth of Con¬

ocarpus ereetus, Cryptocarpus pyriformis, Discaria pauciflora,

Maytenus obovata, Scaevola Plumieri, and Yallesia glabra. The

roots of these bushes prevent the sand from shifting inland too

rapidly.

The country is very flat between Finger Point and Sappho

Cove, and is covered with beds of basaltic lava. In the immedi¬

ate vicinity of the cove this lava is covered with a tolerably dense

growth of xerophytic vegetation which gradually becomes thin¬

ner farther north until it is practically bare in the vicinity of

Finger Point. As the vegetation decreases in amount it also be¬

comes smaller, and such species as Bursera graveolens, which

grow to the size of trees around Sapho Cove and further south,

are mere bushes as Finger Point is approached. This place

illustrates the gradual invasion of lava by higher plant life bet¬

ter than any other visted upon the islands. The most common

plants on the lava beds here are : Aristida subspicata, Borreria

Stewart — Botanical Conditions pn the Galapagos Island 305

ericaefolia, Bursera graveolens, Cardiospermum corindum, Cas¬

sia picta, Cereus galapagensis, Coldenia Darwini, Cordia gala¬

pagensis, C. lu tea, Cryptocarpus pyriformis, Discaria pauciflora

Euphorbia articulata, E. viminea forma chathamensis, Gossyp-

ium barbadense, Lycopersicum esculentum var. minor, Mollugo

gracillima, Pectis tenuifolia, Phoradendron Henslovii, Polygalla

galapagensis var. insularis, Porophyllum ellipticum, Scalesia

divisa, Tephrosia cinera, and Waltheria reticulata forma inter¬

media. All of these plants grow from the crevices of the lava in

which there is usually no appearance of soil at the surface.

The country south of Sappho Cove is covered with much

older lava than is the country north. This lava is very rough in

places and has deep fissures in it. The lower parts here are cov¬

ered with a dense growth of xerophytic vegetation, very similar

to that found in the vicinity of Basso Point, except that many

trees of Hippomane Mancinella are found around an elevation

of 500 ft. These trees usually grow along what appeared to be

an old stream bed, as there were water-worn boulders in it.

There are several small craters near an elevation of 800 ft., cov¬

ered with forests of Bursera graveolens, on which there was an

abundance of fruticose lichen.

Wreck Bay .

Wreck Bay is a rather open sheet of water, somewhat protect¬

ed by reefs, which is situated at the southwest end of the island,

and is the port for the settlement in the interior. The shores

around the bay are composed of steep sand beaches, and low

cliffs of lava. The country is low for some distance back of the

shore, and it is probably of marine origin. There is a moder¬

ately steep ascent at the end of the flat area near the shore, which

leads up to a broad plain, covered with masses of lava and soil

which slopes gradually upward towards the interior to an eleva¬

tion of 500 ft. There are a few small lava hills and craters at

various places on this plain. There is rather a steep slope from

500-800 ft. elevation which leads up on to the rolling plateau re¬

gion covering the central portion of this end of the island.

The settlement is located on this plateau at an elevation of 900

ft. There is a considerable amount of land under cultivation

surrounding the settlement, on which coffee, and sugar-cane are

produced. There is also a large garden in which many of the

20— S. A.

306 Wisconsin Academy of Sciences , Arts , and Letters.

common vegetables and tropical fruits are grown. The pro¬

ducts of the settlement are shipped to Gyayaquil, Ecuador, by

means of a small schooner which usually makes monthly trips

to the mainland. The plateau region has several high hills and

craters on it.

There are no distinctly halophytic plants around the bay so

far as was observed. The beaches here are too steep and the

wave action at times is so strong that such plants would hardly

be able to maintain a hold. The flat back of the beach is covered

with Prosopis trees and bushes.

The country is covered with low dense forests below an eleva¬

tion of 600 ft., which are made up mostly of trees of, Bursera

graveolens, Piscidia Erythrina, Psidium galapageium and Zanth-

oxylum Fagara. In many places in the forest there is a dense

growth of bushes under the trees, which are mostly of the spe¬

cies usually found on the lower parts. Places occur in the for¬

est, however, where the trees are so closely arranged that there

is very little undergrowth. On rocky hills and craters, in the

lower part of this region, there is a considerable growth of Cer-

eus galapagensis. In low places along the side of the road lead¬

ing to the settlement there are also low groves of Hippomane

Mancinella trees. On the steep slopes between 600-800 ft. ele¬

vation there are fewer trees and more bushes than lower down.

The bushes that are commonly found here are : Croton Scouleri

var. grandifolius, Clerodondron molle, Lipochaeta laricifolia and

Psychotria rufipes. In some places on these hillsides, however,

there are small trees of Scalesia pedunculata, and Hippomane

Mancinella, the last of which is sometimes covered with Tilland-

sia insularis. Many of the bushes disappear higher up and those

that remain are very much scattered. There is a heavy growth

of grasses and sedges in this region consisting of the following

Species: Cyperus rubiginosus, Digitaria sanguinalis, Leptoch-

loa virgata, Panicum geminatum, Seleria pterota, Setaria setosa,

and Stenotaphrum secundatum. Ferns which occur in shady

protected places in this region are: Asplenium formosum, A.

sulcatum, Doryopteris pedata, and Dryopteris furcata. The

plateau region above 800 ft. elevation is covered in most places

with_ grasses, the most common species of which is Paspallum

conjugatum, while in low and protected places there are a few

trees and bushes. In temporary pools of water, formed during

the rainy season, such aquatic and semi aquatic plants as Azolla

Stewart — Botanical Conditions pn the Galapagos Islands. 307

caroliniana, Eleocharis capitata, E. fistulosa, Jussiaea repens,

Lemna minor, and Polygonum acre are found. There are

hedges of coffee and lime trees, near the settlement, and occa¬

sionally one of these is found in the open country, probably an

escape from cultivation. There are also deep ditches which were

evidently dug at an earlier day to take the place of fences.

These have become filled in many places with a heavy growth of

bushes of Miconia Robinsoniana, and on the moist perpendicular

walls of the ditches there are many small ferns.

The plateau rises gradually in a northeasterly direction, at¬

taining an elevation of 1,700 ft. at the base of the main moun¬

tain. Around the base of this mountain there are extensive

brakes of Pteris aquilina var. esculenta, and in protected places,

similar brakes of Gleichenia linearis and small trees of Telan-

thera rugulosa occur. The sides of the main mountain, which

rises 400 ft. above the plateau, are covered in many places with

a heavy growth of Lycopodium clavatum and ferns. On the lee¬

ward side of the mountain and other protected places, this

growth is 2-3 ft. high, but where exposed to the constant action

of the wind, it is usually much lower. Occasional tree ferns,

Hemitelia multiflora also occur on the side of the mountain.

Other plants common here are: Cyperus grandifolius, Hibiscus

diversifolia, and Polygonum galapagense. Exposed places near

the top are covered with Sphagnum, while at the top there are a

few bushes and ferns. The soil and rocks in this region are cov¬

ered with lichens, associated with which are large gelatenous

masses which are probably Nostoc colonies.

A good general view of the plateau region was obtained from

the top of this mountain. Except in ravines and other low

places, it is covered with an unbroken stretch of grass-land from

the northeast side around to the south side. On the remaining

sides there are occasional trees scattered over the grassland.

All of the lower part of the island, that is covered with forests

in the vicinity of Wreck Bay, seems to constitute the dry region.

This extends up to about 650 ft. elevation, above which the

transition forms a narrow belt extending to the grassy region

which begins at 800 ft. elevation. The transition region here is

more of a transition from woodland to grassland than from a

xerophytic to a mesophytic vegetation. The moist region is ap¬

parently not well represented here unless we accept certain small

areas, where local conditions are such as to permit a heavier

growth of vegetation, as representing this region.

308 Wisconsin Academy of Sciences , Arts, and Letters.

Culpepper Island.

Culpepper is located farther north than any other island in

the group, and it is also one of the smaller of the islands. De¬

posits of basaltic lava, and tufa make up the island and form

perpendicular cliffs along its sides. The cliffs descend directly

into the sea and render landing impossible except at one place on

the north side where they have become broken down and formed

a large mass of talus. A gently rounded plateau covers the un-

per part of the island and has an elevation of several hundred

feet. This plateau is inaccessible but is heavily covered with

vegetation, which appeared from the vessel to be composed

mostly of Bursera trees and Croton bushes. In addition to the

above, there are a considerable number of small bushes, very

similar in size and appearance to Scalesia Snodgrassii, which oc¬

curs on Wenman Island about twenty miles to the southeast.

Thickets of a species of low Opuntia also occur here. On the

talus slope, near the landing place on the north side, a few

bushes, of Croton Scouleri var. brevifolius, and Telanthera

Helleri occur. Bright red patches of vegetation were seen at

various places along the sides of the cliffs a short distance above

the water. These are probably composed of Sesuvium Edmon-

stonei as it forms similar patches elsewhere. One member of the

party reported having seen the remains of sedges, the first time

that these plants have been reported from this island.

Daphne Island.

Daphne Island is a small tufa crater, about half a mile in di¬

ameter, and 200 ft. high, which is located five miles north of In¬

defatigable Island. The sides of the island are steep, and with

the exception of a few small trees of Bursera graveolens, there is

but very little vegetation on them. The only plants collected

here were : Abutilon crispum, Euphorbia amplexicaule, and Tri-

bulus cistoides, all of which were taken by Mr. F. X. Williams

from the inside of the crater.

Duncan Island.

Duncan is a small island, about three miles in diameter, which

lies between Albemarle and Indefatigable Islands. The shores

are steep and are made up of tall cliffs on all sides but the north

Stew art —Botanical Conditions pn the Galapagos Islands. 309

and northeast. There is a small cove on the northeast side, shel¬

tered by 1a small islet, which affords a good landing place for

boats. Good anchorage can be obtained off the month of this

cove, but as none is marked on the charts of these islands, vessels

should take soundings from a small boat before attempting to

come to anchor here.

The sides of the island are steep in most places, and all of the

lower part is covered with loose fragments of lava among which

there is a scanty soil. There are two old craters in the central

part of the island which directly join one another there being no

distinct rim separating the two. The crater to the north is the

smaller, and its side walls are very steep and redish in color as

is the soil which covers its floor. The southern crater forms a

broad basin the floor of which is 850 ft. above sea level, and 400

ft. above the floor of the northern crater. There are some irreg¬

ularities in the floor of the larger crater, but the most of it forms

a rather flat plain, with occasional low places in it, some of which

appeared to have been recently filled with water. There is a

considerable amount of soil in this crater, which is light gray in

color and loose in texture.

There is a high ridge to the east of the larger crater which in

one place attains an elevation of 1,300 ft., the highest point on

the island. The west side of this ridge is steep and there are

cliffs in some places of a considerable height. The east side of

this ridge is not so steep, however, but slopes downward to the

tops of the cliffs along the eastern shore of the island. The up¬

per part of this frdge is irregular and is covered with soil in

places made up of disintegrated lava and vegetable mold.

Halophytic plants are but poorly represented on the island,

and so far as was observed, consist of a single tree of Avicennia

officinalis, a few Laguncularia bushes, and a few small stunted

trees of Rhizophora Mangle, all of which grow at the end of the

cove on the northeast side of the island. Sesuvium Edmonstonei

also occurs here but not under halophyitc conditions. It is

found in various places on the northeast side of the island to an

elevation of 450 ft.

Unfortunately we were not able to get to this island during

the rainy season, so at the times we visited here the vegetation

was in the resting condition. The lower parts had more the ap¬

pearance of a winter landscape in temperate region than that of

a region within only a few miles of the equator. With the ex-

310 Wisconsin Academy of Sciences , Arts , and Letters.

ception of the few mangroves, which grow near the shore, there

was no other green vegetation worth mentioning. There were

no trees of any kind, even the Bursera trees, which occur so

abundantly in dry places on the most of the other islands, were

entirely absent. This part of the island is covered with open

thickets of bushes consisting of Cordia lutea, Gossypium barba-

dense, Lantana peduncularis, and Prosopis dulcis. Opuntia

galapageia occurs abundantly above 450 ft. elevation, on the

northeast side, but it appears to be almost absent from the north

side. The branches of this species are more loosely arranged

than usual, and the plants have a rather sickly stunted appear¬

ance. Those which occur high up on the island are often heav¬

ily covered with fruticose lichens as is the most of the other vege¬

tation in this region. The vegetation thickens up considerably

around 500 ft. elevation, on the north and northeast sides, and

consists mostly of Croton bushes.

The east side of the island is very rough and covered in places

with long ridges of rough broken lava, many of which extend

down nearly to the tops of the cliffs above the sea. The vegeta¬

tion on these parts consists mostly of Croton and Prosopis

bushes to an elevation of 750 ft. Above this elevation the vege¬

tation is more open, however, and there are areas which are cov¬

ered with small plants, and occasional bushes of Acacia macra-

cantha, Prosopis dulcis, and Zanthoxylum Fagara. The south

sides of many of the large lava boulders in this region are cov¬

ered with Polypodium squamatum, while the north sides are

bare. This is probably due to the fact that the south sides of

these boulders are bathed by the moist winds during several

months of the year, while the north sides are not. The south

side of the island was not visited but from a high point it ap¬

peared to be covered with a dense growth of bushes, many of

which were covered with a brown colored epiphyte, probably a

species of Frullania.

The interior of the smaller cf the two craters was not visied,

but the inner walls appared to be covered with Croton and other

bushes, all of which were heavily covered with lichens. The

floor had a few bushes on it but the growth is not heavy enough

to hide the soil in most places. The floor of the larger crater,

to the south of the smaller one, is sparingly covered with Opun¬

tia galapageia, and bushes of Prosopis dulcis, and Zanthoxylum

Fagara. In low places around dried pools there was an abun-

Stewart — Botanical Conditions son the Galapagos Islands . 311

dant remains of Cyperus rubiginosus. The inner side of the

ridge, to the east of this crater, is covered with a dense growth

of Zanthoxylum and other bushes all of which were covered with

lichens and leafy Hepaties. Along the top of this ridge, above

an elevation of 1,200 ft., there were bushes and small trees of

Acnistus ellipticus, Chiococca alba, Croton Scouleri var. brevi-

folius, Erigeron tenuifolius, Pisonia floribunda, Scalesia Baurii,

Tournefortia psilostachya, T. Pubescens, and Zanthoxylum Fa-

gar a. Exposed rocks in this region are often covered with a

thick growth of ferns which have formed a considerable amount

of vegetable mold upon them. Tillandsia insularis, and two epi¬

phytic species of ferns, Polypodium lanceolatum, and P. lepidop-

teris occur to some extent upon the bushes in this region. Sev¬

eral grassy areas extend down the east side from the top of this

ridge on which there are also small specimens of Opuntia gala-

pageia.

No large trees occur around the top of this island, although

Pisonia floribunda, and Zanthoxylum Fagara, which grow here,

usually attain the size of large trees at similar elevations on

the other islands where these species occur. The absence of

large trees is probably due to the strong winds which strike the

top of the island during a greater part of the year, and thus

prevent the growth of large vegetation.

Botanical regions are not well marked here but practically all

of the plants which occur below an elevation of 900 ft. are forms

typical of the dry regions, above this elevation many of the

plants commonly found in the transition region make their ap¬

pearance.

Gardner Island, Near Charles Island.

This island is situated about four miles off the east side of

Charles Island. It is the smaller one of the two Gardner Is¬

lands, which occur in this group, and consists of an immense mass

of lava several hundred feet high. The shores are steep in most

places and are made up of tall cliffs some of which are perpen¬

dicular. Landing is dangerous, and can only be done with safety

when the surrounding water is comparatively still.

The writer did not land upon this island so that the only

plants collected were by other members of the party. They are

few in number and in no way represent the entire flora. The is¬

land appeared from the vessel to be covered with low bushes

312 Wisconsin Academy of Sciences, Arts, and Letters.

which had a grayish color similar to those found on the lower

parts of the most of the other islands. There were many speci¬

mens of a low species of Opuntia. Lichens seemed to be abun¬

dant on the vegetation.

Gardner Island/ Near Hood Island.

This one of the Gardner Islands is situated about a mile off the

north shore of Hood Island. The water in Gardner Bay, be¬

tween the two islands, is quite shallow, so it is likely that the

two islands have been connected at some past time. The island

is quite small, and is made up of very old lava, some of which

has broken down in places forming a light covering of soil mixed

with small lava fragments. The east and south sides are rather

flat and sand beaches occur along the shores on these sides. The

remainder of the island is rough, however, and the shores are

bordered by tall cliffs. A small bay is surrounded by these on

the north side.

Low bushes of Cryptocarpus pyrif ormi occur in several places

near the shore. The only trees on the island are those of Bur-

sera graveolens and Opuntia galapageia. Bushes of Cordia

lutea, Lantana peduncularis, and Prosopis dulcis are quite com¬

mon.

Hood Island.

Hood is the most southern island of the group, being located

five miles further south than Charles Island, thirty-six miles

west of it. It is also one of the smaller and lower of the islands

as its greater diameter is about eight miles and its highest point

has an elevation of 640 ft. So far as was observed, the shores

are high and rocky on all sides but the northeast where there are

long stretches of sand-beach and low rocky shore. The sides

slope up gradually from the shore at Gardner Bay, to a some¬

what flat central region on which there are several rocky hills,,

some of which rise possibly a hundred feet above the surround¬

ing country. There is no distinct crater on this island. There

is, however, a broad flat plain, about half a mile south of Gard¬

ner Bay, which may be the floor of a crater, the surrounding

hills being all that is left of the rim.

The highest point is towards the southwest side of the island,

and consist of a flat-topped hill of lava. A considerable amount

Stewart — Botanical Conditions pn the Galapagos Islands . 315

of soil occurs in various places which has resulted from the

breaking down of the lava. The most of the soil has probably

been washed from above as it usually occurs in low places. The

most of the isalnd has but very little soil on it, and the surface is

strewn with fragments of lava.

The beaches are rather steep here in consequence of which

there is not a great amount of halophytic vegetation. Bushes of

Cryptocarpns pyriformis occur on the sand beaches, and at vari¬

ous other places near the shore. Patches of these bushes form

about the only green vegetation during the greater part of the

year, and they stand out sharply when the island is examined

from the top of one of the hills in the interior. Sesuvium Ed-

monstonei grows here but usually not under halophytic condi¬

tions. The mangrove vegetation is confined to a small thicket

of bushes of Rhizophora Mangle which occur on the northeast

side of the island below Gardner Bay. The beach, in the vicin¬

ity of these mangroves, was strewn with pieces of bamboo, co-

coanut husks, and other drift, showing that this area receives a

more or less constant supply of such material.

Other plants which occur on or near the beaches are : Cacabus

Miersii, Cenchrus distichophyllus, Coldenia fusca, Discaria pau-

ciflora, Maytenus obovata, and Vallesia glabra.

The only trees of any size on the island are those of Bursera

graveolens and Opuntia galapageia. The Opuntias have rather

low thick trunks and closely arranged branches. Goats, which

have been introduced upon this island within the last few years,

eat off all of the lower Opuntia branches, and they even eat into

the thick trunks in some instances. As these plants form their

only suitable food and probably their only source of water,

for several months of the year, there is danger of this species

being exterminated on this island in time.

The most of the vegetation on the island consists of bushes, the

most common of which are: Acacia macracantha, which forms

small trees in protected places, Cordia lutea, Croton Scouleri,

Gossypium barbadense, Lantana peduncularis, Parkinsonia acu-

leata, and Prosopis dulcis. These bushes occur in patches in

many places in between which there are open spaces which are

probably covered with grasses and annual herbaceous plants

during the rainy season. By following these open spaces one can

travel over the most of the island without much difficulty.

There are some indications of a greater amount of moisture

314 Wisconsin Academy of Sciences, Arts, and Letters.

around 600 ft. than lower down, as Polypodium squamatum

grows from the lava crevices at this elevation. There are also a

considerable number of small trees of Zanthoxylum Fagara in

this vicinity which is near the base of the high hill on the south¬

west side of the island. The top of this hill is covered with

bushes of Cordia galapagensis, Cryptocarpus pyriformis, Ly-

cium geniculatum, Tournefortia psilostachya, and Vallesia gla¬

bra. Ipomoea Habeliana covers the. rocks here to a considerable

extent.

Indefatigable Island.

Indefatigable is the second largest, and with the exception of

Duncan, is the most centrally located island in the group. It is

roughly circular in outline, and appears to have a large central

crater when seen from the south side. There is probably no

larger crater present, however, because in sailing around the is¬

land towards the west side it is seen that the upper part is cov¬

ered with many small craters, twenty-one of which were counted.

Small lateral craters also occur at various places along the sides,

the largest one of which is on the southeast side of the mountain

a short distance below the top.

The shores along the south, southeast, and east sides of the is¬

land are bordered by low rocky cliffs of lava in most places,

while on the other sides the shores are low with occasional sand

beaches. There is a large bay on the south side which we chris¬

tened it Academy Bay in honor of the California Academy of

Sciences.

The sides of the island slope up very gradually on all sides so

that it is necessary for one to travel several miles inland in order

to get into the region where collecting is good. All of the lower

parts are covered with the usual xerophytic forms so common on

these islands, but which are not in proper condition for collect¬

ing through a greater part of the year. The lower parts are

covered with lava on which there is but little soil, but in the in¬

terior there is an abundant soil, and in places there is said to be

water, although we were not fortunate enough to find it. The

interior of this island is very fertile, and with proper cultivation

is capable of supporting quite a large population, but up to the

time we visited it no attempt had been made to establish a

settlement there.

We visited the island at Academy Bay, at two places on the

Stewart— Botanical Conditions pn the Galapagos Islands . 315

north side, the northeast side, the northwest side at a point a

little south of Conway Bay, and on the southeast side.

Academy Bay .

Academy Bay is a small body of water, partly surrounded by

-cliffs, on the south side of the island. It is marked by a small

islet which lies on the east side of the entrance. Small vessels

can find good anchorage in this bay but care should be taken in

anchoring in the western part of it as there are hidden rocks

present there. This part of the bay is the best protected from

the southeast swell, so we anchored there on our first visit to

this place, and were unfortunate enough to get aground on two

occasions. The best landing place for boats is at the north end

of the bay where there is a small sand beach, and a low flat area

covered with bushes and grass, back of which there is an old

trail leading into the interior. Two springs of brackish water

occur here, each of which are marked by a bunch of small trees

of Hibiscus tiliaceus. The country is very rough for a mile

or more back from the beach and is covered with low ridges of

lava, many of which run in a direction nearly parallel with the

coast line. There are also many crevices in this lava, some of

which are evidently quite deep. The lava has disintegrated but

little on this part so there is very little soil to be seen on the sur¬

face, but nevertheless it is heavily covered with vegetation.

To the north of the rough area just mentioned, there is a line

of cliffs, about 75 ft. high, above which the lava is evidently

much older, as it has broken down in many places into soil,

through which boulders of lava protrude at intervals. The

amount of soil increases farther inland, completely covering the

lava in most places above an elevation of 500 ft. The soil at this

elevation, and above, is composed largely of vegetable mold

which has been formed from the decay of the abundant vegeta¬

tion in this region.

Small swamps of Rhizophora Mangle occur at several places

around the shores of Academy Bay and around a small lagoon

which empties into it. There are also clumps of Laguncularia

bushes along the shore, and occasional trees of Avicennia offici¬

nalis around salt water pools in the vicinity of the shore. Back

of the beach, at the north end of the bay, there is a small area

that is thickly carpeted with Sporobolus virginicus. Thickets

316 Wisconsin Academy of Sciences , Arts , and Letters.

of Cryptocarpus bushes also occur here covered with Passiflora

foetida. Occasional trees of Hippomane Mancinella also grow

on this area. In many places along the west and south sides of

the bay the growth consists of low bushes, with a considerable

number of Cereus sclerocarpus and Opuntia myriacantha trees

scattered among them.

After leaving the immediate vicinity of the shore, at the north

end of the bay, one encounters dense jungles of xerophytic

plants which extend inland a mile or more to the base of the

cliffs mentioned above. This jungle is composed largely of trees

of Acacia macracantha, Bursera graveolens, Erythrina velutina,

Opuntia myriacantha, and Piscidia Erythrina. The specimens

of Opuntia are very large, some of them attaining a height of

thirty or more feet. In general it may be said, that nearly all

of the species which occur in this area either attain a larger size,

or grow more abundantly than they usually do so near to sea

level. There is a dense growth of bushes underneath the trees

consisting of Cordia lutea, Croton Scouleri varieties, brevifolius

and Macraei, Discaria pauciflora, Gossypium barbadense, Lan-

tana peduncularis, Maytenus obovata, Parkinsonia aculeata,

Prosopis dulcis, Scalesia gummifera, Telanthera echinocephala,

Tournefortia pubescens, and Zanthoxylum Fagara. Such ferns

as Polypodium squamatum and Trachypteris pinnata grow on

the sides of the cliffs at an elevation of 75 ft. Above these cliffs

there is a considerable area which is covered with Prosopis and

other bushes of an xerophytic character, but the arrangement of

these is more open than below the cliffs. The trees of Opuntia

myriacantha are numerous and very large in this area, and form

a portion of the continuous zone of Opuntia trees which extend

around the south side of the island. They are so numerous here

that their redish-brown trunks give this color to the surrounding

landscape when seen from a distance.

There is a general thickening up of the vegetation further in¬

land, but there is not much change in the species of plants pres¬

ent below 350 ft. Around this elevation such forms as Cordia

lutea, Croton Scouleri var. brevifolius, Discaria pauciflora,

Opuntia myriacantha, Parkinsonia aculeata, Piscidia Erythrina,

Prosopis dulcis, and Telanthera echinocephala disappear. The

most of the vegetation, around this elevation, is heavily covered

with fruticose lichens. There are dense forests here made up

largely of trees of Pisonia floribunda, Scalesia pedunculata, and

Stewart — Botanical Conditions pn the Galapagos Islands, 317

■ Zanthoxylum Fagara, the first and last of which occur near the

shore as bushes. The trees are often covered with a heavy growth

of Cisampelos Pareira, which usually put down large numbers of

aerial roots, forming tangled masses, rendering traveling dif¬

ficult. Projecting ridges of lava occur in some places in this

region, which are usually covered with dense mats of Polypo¬

dium squamatum, ands such herbaceous plants as Peperomia gali-

oides, P. galapagensis, P. Stewarti, and other forms. The trunks

and branches of many of the trees, especially those of Pisonia

floribunda, are heavily covered with epiphytic plants such spe¬

cies as: Asplenium sulcatum, Ionopsis utricularioides, Lycopo¬

dium dichotomum, Polyp odium lanceolatum, P. lepidopteris,

Peperomia galapagensis, and Tillandsia insularis being the most

common. Phoradendron Henslovii also occurs in this region

and higher up, but it grows much larger than it does lower down.

Owing to the dense shade there are fewer bushes, but more herb¬

aceous forms than at a lower altitude. Ferns are also common.

The region above an elevation of 500 ft. on this side of the is¬

land consists of two distinct parts as far as the vegetation is

concerned. The country immediately north of Academy Bay,

above this elevation, is covered with dense forests which extend

towards the east side of the island. In some places north of

Academy Bay these forests probably begin a little lower down,

but farther east they eyidently begin somewhat higher, as they

were not encountered at an elevation of 650 ft. when the south¬

east side of the island was visited. It might be said in this con¬

nection, that all of the botanical regions gradually ascend to¬

wards the east side of the island here, a condition which can be

readily seen from the shore by the difference in color of the dif¬

ferent regions. The forests, just mentioned, were not visited

but from theirappearance they must be made up largely of trees

of Scalesia pednculata, and other species common in the Scalesia

forests of these islands. Northwest of Academy Bay there are

extensive areas covered with bushes on which there is a heavy

growth of Argyreia tiliaefolia and other vines. The vegetation

m this region was denser than in any other place visited upon

the islands. Traveling was very slow and difficult here it being

necessary at times to lift one member of the party up and let him

fall at full length into the bushes and other vegetation in order

to mash them down, for it was almost impossible to cut one’s

way through this vegetation, loaded down with water and food

318 Wisconsin Academy of Sciences , Arts 9 and Letters.

as we were. The principal bushes in this region are : Psychotria

rufipes, Tournefortia rufo-sericea, Urera alceaefolia, and some

others, the time spent in this region not being sufficient to make

as complete collections as was desirable. There are many ferns

and herbaceous plants in this region. Many of the herbaceous

forms which occur here also occur lower down but are much

smaller in size. The most noteworthy of these are: Crotalaria

setifera and Fleurya aestuans, the last of which has fewer sting¬

ing hairs than at the lower elevations where it occurs. Groves

occur occasionally in the bushy areas, made up of the trees usu¬

ally found in the moist regions. Some of these groves are quite

large, but usually they are small. Isolated trees are not at all

uncommon.

No plants were collected above an elevation of 650 ft. on this

side of the island, but other members of the party who succeeded

in getting farther inland, report that there is a decrease in the

number of vines and an increase in the size of the bushes higher

up. Messrs. Williams, Ochsner, and Gifford succeeded in reach¬

ing an elevation of 1,100 ft. here, reported that the country, a

short distance beyond where ther discontinued their journey, ap¬

peared to be covered with low spreading trees or bushes on

which there was a large amount of “brown moss.” A large part

of the country, above the Scalesia forests, and bush areas, had a

distinctly brown color in which there are patches of green. The

brown color is possibly due to a heavy growth of one or several

species of leafy Hepatics, and is confined to the south side of the

island, none of it appearing on the other sides. The top of the

mountain is covered with green vegetation, but it is likely that

there are no trees present there, because none appeared in the

sky-line when the top of the mountain was viewed with a field

glass on a clear day.

As near as could be ascertained, the dry region extends to an

elevation of about 350 ft., the transition region to 500 ft. while it

is likely that the moist region extends to at least 1,500 ft. and

possibly higher.

Judging from the appearance, the upper part of this island

ought to be very interesting botanically, as it is apparently en¬

tirely different from the upper part of any other island visited.

It would probably require a week or more to explore this re¬

gion properly, and in order to do this, it would be necessary to

cut a good trail into the interior, and a camp established, where

Stewart — Botanical Conditions /on the Galapagos Islands . 319

supplies of food and water could be brought in each day. It

would probably be more economical to have this work done by

laborers, as they could probably be secured from Albemarle Is¬

land at a small cost. Unfortunately our expedition was not fi¬

nanced in a way to make this possible.

Southeast Side.

The place visited on this side of the island, is situated about

seven miles east of Academy Bay, the region just described. It

is dangerous for vessels to anchor here, except during calm

weather, as there is a strong swell at other times. We visited

this place during the month of October, and were greatly incon¬

venienced by the violent rocking of the vessel at times. There

are several hidden reefs between the anchorage and the shore, on

which the swell breaks heavily at times. One has to use care in

going in and out in a small boat when there is even a slight

swell, because it is liable to break in unexpected places. A small

crater stands near the shore at this place. Broad sand beaches

border the shore, back of which there is a sandy basin containing

pools of brackish water. There is supposed to be fresh water in

this vicinity but we were unable to find it.

With the exception of a few ravines in the vicinity of the

shore, the country is comparatively smooth for some distance

inland, and is covered with a fair amount of soil. Farther in¬

land, above an elevation of 200 ft., there are beds of more re¬

cent lava which has been piled up in places forming low ridges

and valleys. Several small craters are located about four miles

inland, between 400 and 500 ft. elevation, which rise on an av¬

erage of about 100 ft. above the surrounding country. The

country to the right of these craters is rough and covered with

irregular masses of lava, while to the left it is comparatively

smooth. The country was not visited beyond these craters, but

it appeared from the top of the one fartherest inland, to have no

pronounced irregularities in it as it sloped gradually up to th$

base of the craters in the central part of the island.

Rhizophora Mangle occurs in isolated patches along the shore,

but there are no large swamps of it, probably due to the fact

that the surf breaks here much of the time. Quite a number of

other species of plants grow on the beaches among which are

Coldenia Darwini, Cryptocarpus pyriformis, Heliotropium cur-

320 Wisconsin Academy of Sciences , Arts , and Letters.

assavicum, Scaevola Plumieri, Sesuvium Portulacastrum, and

Vallesia glabra. The low sandy area back of the beach is cov¬

ered in some places with a heavy growth of Sporobolns virgini-

cus, while in other places, where the sand is encrusted with salt,

Ipomoea Pes-caprae grows very abundantly and of great length,

individual plants sometimes being fully one hundred feet long.

Cyperus laevigatus is very common around the pools of brackish

water, which have a strong odor of Sulphurated Hydrogen.

Bordering this sandy area are low dense groves of Avicennia offi¬

cinalis, Hippomane Mancinella, and Laguncularia racemosa.

There are several other low areas and ravines in the vicinity of

the shore, above high tide mark, which are filled with thickets

of Discaria pauciflora, and Parkinsonia aculeata. The remainder

of the lower part of the island at this place is covered with

bushes, Bursera trees, and cactus. Cereus sclerocarpus occurs

commonly in the vicinity of the shore but was not seen above an

elevation of 100 ft. Opuntia myriacantha, on the other hand,

occurs abundantly on the lower parts, and to a considerable ex¬

tent to about 500 ft. At an elevation of 600 ft., however, it is

scarce, and much smaller than lower down. The specimens seen

at this elevation, were of about the same size as those which oc¬

cur at an elevation of 350 ft. at Academy Bay a few miles west

of here. The Bursera trees are larger and more abundant above

an elevation of 350 ft. than they are lower down. They are

usually heavily covered with fruticose lichens. Piscidia Ery-

thrina is another common forest tree in this region.

There is a very noticeable decrease in the number of many

of the forms common on the lower parts, between 350 and 450

ft. elevation. Some of the species disappear here among which

are : Croton Scouleri, Discaria pauciflora, Maytenus obovata,

and Telanthera echinocephala. A few stunted specimen of Cheil-

anthes microphylla were found in the lava crevices at an eleva¬

tion of 350 ft., and were the first ferns ever collected on this

island.

There are fewer trees in the region of the craters, at 400 ft.,

elevation, than lower down, and the country is covered with

Lantana bushes four to five feet high with a few trees scattered

through them. The sides of the craters are covered with low

bushes of Euphorbia viminea, and other species, while at the

top there are bushes, ferns, and grasses. The country to the

north of these craters, is heavily forested with Bursera and

Stewart — Botanical Conditions on the Galapagos Islands . 321

other trees, under which there are tangled thickets of Zanth-

oxylum bushes.

Low forests cover the country between 450 and 650 ft. eleva¬

tion, which are made up of a mixture of xerophytic and meso-

phytic forms. Bursera trees are common on the lower part of

this area, but they decrease in number higher up, where the

forests are composed largely of Pisonia floribunda, Psidium

galapageium, and Scalesia pedunculata. The Scalesia trees

are smaller, and fewer in number here than they are in the

Scalesia forests higher up. Quite a number of ferns grow in

the vegetable mold, and on exposed rocks in this region, such

species as Polypodium pectinatum, P. squamatum, and Trachy-

pteris pinnata being the most common. A few epiphytes grow

on the trees and bushes among which are : Ionopsis utrieular-

ioides, Peperomia galapagensis, Polypodium lepidopteris, and

Tillandsia insularis. Common bushes in this region are : Chio-

cocea alba, Erigeron tenuifolius, Gossypium barbadense, and

Tournefortia strigosa. The region above an elevation of 650

ft. was not visited, but it appeared to be covered with Scalesia

forests a short distance above this elevation.

The dry region extends to an elevation of about 400 ft. at

this place. We did not reach the upper limit of the transition

region, but as near as could be estimated, it extends to about

800 ft., the elevation at which the Scalesia forests probably be¬

gin.

No effort was made to get far into the interior at this place,

as we made our first visit to the island here, and expected to

find better places for doing this elsewhere later in our trip.

After having failed to accomplish this at other places, it now

seems probable that this would have been the best place to

have made the attempt after all. It is very likely that one

would have no difficulty in reaching the lower edge of the

Scalesia forests in a half day’s journey, if no collecting were

done on the way. From this point it is probably not over three

or four milesi to the base of a large lateral crater, on the south¬

east side of the mountain just below the top. Judging from

its appearance, this crater would present about the same botan¬

ical conditions as occur around the top of the mountain. The

advantages to be gained from exploring the interior of the is¬

land from this side are ; the slope is not so gradual, so that ele¬

vation could be gained by traveling shorter distances, and,

21— S. & A.

322 Wisconsin Academy of Sciences , Arts , and Letters.

there are heavy forests in the moist region which would prob¬

ably have less undergrowth in them than the bushy areas en¬

countered in this region at Academy Bay. Should another

party ever attempt to reach the interior of the island from

there, they should keep well to the south of the group of cra¬

ters, about four miles inland, as the country is not so rough,

nor is the vegetation so dense as it is north of these craters. A

low monument of lava boulders was built by the side of the

trail where We came into it in coming back from the interior.

It is likely, however, that the trail will be entirely obliterated

before another party attempts to explore this island.

Northeast Side.

The island was visited on this side at a point about three miles

west of Gordon Rocks which are situated a short distance off

shore. The coast is bordered by low cliffs in this vicinity, which

rise abruptly fifteen or more feet above the water. There are oc¬

casional small sand beaches, however, so that landing from boats

can be easily accomplished. The country is very flat for some

distance inland, and for the most part, is covered lightly with a

reddish colored soil. Farther inland there are rough deposits of

lava. We had expected to try to reach the interior of the island

from this place,- but we did not attempt it after we had dis¬

covered the rough character of the country.

There are no true halophytes along the shores in this vicinity,

a condition that is accounted for by the lack of extensive sand

beaches, such as occur on the other sides of the island where there

is a more or less extensive halophytic flora. Such plants as Cryp¬

tocarpus, pyriformis, Discaria pauciflora, and Maytenus obovata,

which sometime grow under semihalophytic conditions, occur,

however, at this place.

The country is very barren in the vicinity of the shore, and

there are no trees present unless a few stunted specimens of Bur-

sera, and Prosopis could be called such. The country farther in¬

land, however, appeared to be covered with forests of Bursera

trees, but as we visited this region during the dry season, the

general appearance of these forests was too uninviting to tempt

one to venture far to examine them. All of the vegetation of

any considerable size, leans in a northwestern direction showing

the influence of the strong southeast winds during the growing

Stewart — Botanical Conditions on the Galapagos Islands. 323

season. The strong winds, and the loose character of the soil,

are the probable causes for the small amount of vegetation here.

The ground in most places is covered with a sparse growth of

grass, the two common species of which are, Aristida subspicata7

and Panieum hirticaulum.

Nprth Side.

The shores along the north side of the island are made up of

sand beaches and low rocky coast, which in most instances slope

down gradually to the edge of the water. There are but few

cliffs along the shore on this side, a condition that is probably due

to the fact that this is the leeward side of the island and is con¬

sequently not subject to strong wave action as are the more ex¬

posed sides.

For a considerable distance inland the country is flat and cov¬

ered in many places by beds of basaltic lava on which there is

but little soil. There are small areas, however, which are cov¬

ered with a light gray ashy soil in among the deposits of lava.

Rough lava ridges are common, and there are several low lava-

hills, and small craters scattered around at various places on this

side.

The sand beaches for the most part bear the usual herbaceous

plants, and in the vicinity of the shore thickets of Cryptocarpus

pyriformis, and Yallesia pubescens bushes are common. Bushes

of Maytenus obovata line the shore in places, some of which grow

so close to the water’s edge that their roots are covered at high

tide. In such places the trunks are more gnarled, and the leaves

more succulent than is usually the case. Mangrove swamps oc¬

cur in several places, the largest one being located about two

miles west of the lower end of South Seymour Island. So far as

was observed this is the most typical mangrove swamp on the is¬

lands. A shallow bay occurs here which has a narrow opening

into the sea and affords much quieter water than in most places

on these islands. There are several small islets in the bay, and

these as well as the shores of the bay are heavily covered with;

mangroves. One is able to get through the swamp in a small!

boat by following the deeper channels at high tide. We found it

to be an excellent place for capturing sea turtles. Trees of Avi-

cennia officinalis and Laguncularia also occur in this swamp, but

Rhizophora makes up the bulk of the vegetation. All three spe-

324 Wisconsin Academy of Sciences, Arts, and Letters.

cies occur at various other places along the north shore, and Avi-

cennia was found in one instance to be growing in a sunken

place, a short distance inland, which was apparently filled with

sea water at high tide.

The interior is barren in many places where there are expos¬

ures of lava. With the exception of a few low stunted trees of

Bursera graveolens, a low Opuntia, and scattered bushes these

beds bear no other vegetation. The species of Opuntia is possibly

a new one as it differs quite markedly from the other species in

this genus which occur on these islands. It also occurs on South

Seymour Island, a few miles away, but at neither place were the

specimens in good shape at the times these were visited. Occa¬

sional bushes of Acacia macraeantha, and thickets of Croton

bushes occur in small patches between the lava beds, where there

is a sufficient amount of soil to support them. Grassy areas oc-

43ur, between the patches of bushes, which are usually covered

with Aristida subspicata. These areas continue at intervals, to

an elevation of 1,000 ft. as high as this side of the mountain was

explored. Very little change takes place in the character of the

vegetation at this elevation, except that some of the species grow

to a larger size than they do lower down. This is especially true

of Bursera graveolens, and Piscidia Erythrina, both of which are

quite small near the shore but form trees in the interior.

The dry region extends to possibly an elevation of 1,500 ft. on

this side of the mountain, as the appearance of the vegetation

did not appear to change for several hundred feet above where

exploration was discontinued. The moist region seems to form

.a narrow zone just below the top on this side of the mountain.

Northwest Side.

This side of the island was visited at a place about two miles

south of Conway Bay, which is marked by a small tufa crater

near the shore. Our reason for stopping here instead of at Con¬

way Bay, the usual place for landing on this part of the island,

was because we had learned at Chatham Island that an old trail

led into the interior from this point.

The coast is low in this vicinity, and is made up of sand

beaches and low cliffs. A flat area surrounds the tufa crater,

mentioned above, covered with a soil composed largely of volcan¬

ic ashes, which have evidently originated from the crater. This

Stewart — Botanical Conditions ,on the Galapagos Islands. 325

condition is local, however, as the most of the soil on the lower

parts is composed of disintegrated lava with small boulders and

bits of lava intermixed. There are also lava ridges in this vicin¬

ity without soil.

The slope is very gradual to 700 ft. and in places the country

is slightly terraced. Above this elevation the ascent is quite

steep to 1,000 ft. beyond which there is a broad valley, three or

four miles wide, extending in to the base of the craters on the

west side of the island. There is a heavier growth of vegetation

here than on the north side of the island, probably due largely

to the fact that there is more soil.

About the only vegetation on the beach, where we landed, was

a sparse growth of Sporobolus virginicus, but back of the beach

around the base and on the sides of the cliffs there were bushes

of Cryptocarpus pyriformis, Discar ia pauciflora, and Maytenus

obovata. Grassy areas occur back of the shore, where the soil is

composed of ashes, which at the time of our visit were covered

with a heavy growth of Aristida subspicata. There is probably

quite a growth of annual plants in addition to the above, during

the rainy season, as the remains of quite a number of these were

found. Patches of bushes occur in the grassy areas which are

made up of such species as Acacia macracantha, Clerodendron

molle, Cordia lutea, Croton Scouleri var. brevifolius, Gossypium

barbadense, and Waltheria reticulata forma intermedia. Low

trees of Bursera graveolens and Piscidia Erythrina occur among

the bushes. A few specimens of Opuntia grovT in the vicinity of

the shore but whether or not these are O. myriacantha or the un¬

described species from the north side of the island, was not de¬

termined, as no specimens were collected. They are more abun¬

dant farther inland.

The character of the vegetation changes but little to an eleva¬

tion of 300 ft. except that the grassy areas soon stop and the

country is covered with Bursera forests very similar to those on

the lowrer parts of other islands. Trees of Erythrina velutina

also occur here in considerable number. Cissampelos Pareira

appears first at about 300 ft. elevation, but it becomes more

abundant higher up where it often covers trees and bushes. A

few of the more xerophytic species of ferns appear around an ele¬

vation of 400 ft. We experienced much difficulty with the

thickets of Furcraea cubensis at an elevation of 450 ft. and

above, as they often cover areas of several acres in extent in this

326 Wisconsin Academy of Sciences, Arts, and Letters.

region. This species was brought to this island many years ago

by the tortoise hunters who made a temporary settlement here,

and who introduced it among other domesticated plants. They

were planted along the trail, leading into the settlement, by

Captain Thomas Levick of Chatham Island, on one of his period¬

ic visits to this place. He hoped to permanently mark the trail

by this means, but judging from the difficulty we had getting

around these thickets and finding the trail again, his method of

marking it was rather too effective. They grow so thickly that

they have driven out the smaller vegetation in places.

The country around 650 ft. elevation is heavily forested with

trees of Bursera graveolens, Piscidia Erythrina, Pisonia flori-

bunda, and Zantlioxylum Fagara. This forest is open in places

and the ground is covered with occasional bushes and ferns.

Above 700 ft. elevation the forms which occur abundantly on the

lower parts, disappear. The forests above this are made up of

large trees of Pisonia floribunda, Psidium galapageium, Scalesia

pedunculata, and Zantlioxylum Fagara, the last of which forms

trees 25-30 ft. high, usually heavily covered with mistletoe. The

undergrowth is usually dense in these forests and is composed

largely of bushes of Erigeron tenuifolius, Psychotria rufipes,

Tournefortia rufo-sericea, and several species of ferns and herb¬

aceous plants. Epiphytes are common, consisting of ferns and

orchids, Ionopsis utricularioides. Conditions similar to the

above, continue to an elevation of 1,000 ft. as high as this side of

the island was explored. From a tree at this elevation, the coun¬

try appeared to change but little for several miles further in¬

land.

We visited this place during July, and as it was the last time

that we expected to stop on this island, we made a rather deter¬

mined effort to get farther into the interior than we had done be¬

fore. We hoped to follow the trail in as far as the old settlement

where it is reported that water can be found, and where there

are a number of domesticated plants growing which are suitable

for food. We expected to camp at the settlement and try to

work inland from there. We lost the trail at 1,000 ft., however,

and had to turn back as our supply of food and water was not

sufficient to justify us in going farther. The only evidences of

former habitation found, was a number of lime trees near where

we lost the trail. We learned afterwards that these were but a

short distance away from the settlement, but as everything was

Stewart — Botanical Conditions on the Galapagos Islands. 327

so overgrown with vegetation in this region, it probably would

have been impossible for us to have found it unless the trail had

led directly to it.

The botanical regions are fairly well marked here. The dry

region seems to extend to about 450 ft. and the transition region

to about 700 ft. elevation. The moist region seems to be evenly

forested and has none of the open areas covered wTith bushes and

vines, such as was found at Academy Bay on the south side of

island.

James Island.

James, the fourth largest island in the group, is located nine

miles northeast of Cowley Bay, Albemarle Island, and twelve

mile north by west of Indefatigable Island. The general shape

of the island is a parallelogram the length of which is about

twenty miles, and extends east and west. With the exception of

a few sand beaches, the shores are rocky and are bordered in

most places by low cliffs of lava. The eastern part of the island

is low, and slopes up gradually to a broad central plateau which

extends, with a gentle slope, to the base of the main crater, lo¬

cated towards the west end of the island. This crater has an ele¬

vation of 2,850 ft., and can be more easily reached from James

Bay than from any other point. Many other craters occur on

the island, but with one or two exceptions, they are all small.

There are a number of these in the vicinity of Sullivan Bay, and

along the south side. Deposits of basaltic lava, and volcanic

cinders, cover the greater part of the island, the most of which is

quite old, and has become very much oxidized. In many places

in the interior, it has become entirely broken down on the sur¬

faces, into soil, which is mixed with quite a large amount of

vegetable mold. There are, however, some very recent deposits

of lava on the south side, some of which have recently come from

a small crater which has been active within the last few years.

Deposits of tufa occur on the west side, but they are very local

in their distribution.

Northeast Side.

This side of the island was visited about six miles northwest

of Sullivan Bay. There is a small salt-water lagoon here, appar¬

ently the only one on the island. The shores are low and rocky

328 Wisconsin Academy of Sciences, Arts, and Letters.

in most places in this vicinity, but short sand beaches occur occa¬

sionally. Th e country is covered with beds of rough basaltic

lava and cinders all of which is very old and has become stained

with a dark brown color. This lava has not become broken down

to any extent, so consequently there is but little soil in this re¬

gion. The ascent is very gradual here so that it is necessary to

go about four miles inland in order to reach the plateau region,

which covers the central part of the island. The eastern part of

this plateau has a general elevation of about 700 ft., but it slopes

up gradually, towards the west, to the base of the mountain.

There is more soil on the plateau than lower down, but it is

mostly in low places so that the surface of the ground is usually

strewn with lava fragments. Several old craters are located on

the plateau at an elevation of 700 ft., all of which are low in al¬

titude.

The sand beaches support many of the smaller plants which

are usually found in such situations on these islands. Among

these are: Batis maritima, Cryptocarpus pyriformis, Disearia

pauciflora, and Sporobolus virginicus. Rhizophora Mangle is

the only one of the mangroves that grows on the open coast at

this place. It also grows to some extent around the shores of the

salt water lagoon, mingled in places with Avicennia officinalis.

In the vicinity of the shore, the country is covered in places with

thickets of Disearia, and Maytenus bushes, but there are no

trees except those of Opuntia myriacantha. There are a few

specimens of Bursera, but they are mere bushes, and do not grow

to the size of a tree for some distance inland. The crowns of the

Bursera trees are usually much flattened, due to the action of

the wind. There are quite a number of species of bushes farther

inland, all of which grow from the crevices of the lava. Among

these are: Alternanthera rigida, Cordia lutea, Croton Scouleri

var. albescens, Euphorbia articulata, and Scalesia atractyloides.

With the exception of the halophytes along the shore, the Scale¬

sia bushes were about the only plants that presented a green ap¬

pearance at the time of our visit. There were many other plants

in leaf at this time, but the leaves were either small, or covered

with a dense coating of hairs so that the green color was ob¬

scured. There are small stretches of lava near the shore on

which there is practically no vegetation.

The vegetation becomes more abundant between 100 and 200

ft. elevation where there are extensive thickets of Lipochaeta

Stewart — Botanical Conditions pn the Galapagos T stands, 329

bushes, which also occur still higher up on this side of the island.

This species is infested with Phoradendron Henslovii as are

some other forms in this region and above. The vegetation is

further added to, between 300 and 400 ft. elevation, by the ap¬

pearance of small trees of Erythrina velutina, and thickets of

Zanthoxylum bushes, the last of which apparently does not at¬

tain tree size on this side of the island. Opuntia galapageia oc¬

curs in this region and continues to above 700 ft. elevation.

The plateau, around 700 ft. elevation and above, is covered

with much the same sort of vegetation as the lower country, ex¬

cept that it is thicker in places, and some of the species attain a

larger size. No ferns or other distinctly mesopliytic plants were

found if occasional small trees of Pisonia floribunda be expected.

The small craters on the plateau are covered with bushes, and

trees of Cereus sclerocarpus and Opuntia galapageia all of which

have a considerable amount of fruticose lichen on them. A good

view of the surrounding country can be had from the tops of these

craters. The country to the south and east was barren in the ex¬

treme and appeared to be covered with much the same sort of

vegetation which occurs in the region explored on the north side

of the island. The country to the west appeared almost as Un¬

inviting, for possibly 700 ft. higher, and the vegetation all had

the distinctly gray color characteristic of the dry regions of these

islands. It was noticed, while we were sailing along the north

shore, that similar conditions to the above are present nearly to

the top of the mountain on this side.

James Bay.

The conditions at James Bay, at the wrest end of the island,

were much more inviting than they were on the northeast side.

A sand beach extends along the east side of the bay and affords

a good landing place for boats the most of the time. The north

side of the bay is bordered by cliffs, which rise in height towards

the northwest and terminate in a tall perpendicular cliff about

opposite Albany Island, which is situated a short distance off

shore. The coast is rocky for some distance south of the bay,

and is made of low up cliffs of recent lava. Back of the sand

beach, just mentioned, there is a stretch of flat sandy country, in

which there is a small salt water lake. The flat country extends

to the base of the mountain which rises quite abruptly to an ele¬

vation of about 1,400 ft., beyond which the slope is more gradual.

330 Wisconsin Academy of Sciences , Arts , and Letters.

and the country is in the nature of a table land to the base of

the main crater at about 2,200 ft. elevation. The sides of this

crater are steep and the top has an elevation of 2,850 ft. To the

south and southwest of the main crater there are deep valleys

in between other craters and hills. The whole of the south side

of the island is steep, above 900 ft. elevation, for several miles

east of James Bay. Below this elevation, however, the slope is

more gradual and the country is covered with recent flows of

lava, the most of which has probably come from one or more of

the small lateral craters around 900 ft. elevation. The lava

fields are comparatively smooth near the shore, but higher up

they become rough, and in places the lava has cooled enclosing

gas bubbles with only a thin shell of lava above, through which

one breaks in walking.

The region around Sugarloaf mountain, towards the south¬

west side of the island, is covered with tufaceous deposits, which

have probably come from the tufa craters in this vicinity. One

of the smaller tufa craters, encloses a salt water lake, on the bot¬

tom of which there is a layer of apparently nearly pure salt sev¬

eral inches in thickness. The people from the inhabited islands

used to come here for their supplies of salt many years ago.

Except on the recent lava and on the steeper sides of the

mountain, there is a considerable amount of soil to be found all

over this part of the island. The soil is composed of disinte¬

grated lava, which on the higher parts is mixed with vegetable

mold. No springs occur on this island but there are a few small

stream beds in the upper region, which appeared to have con¬

tained water at some time, as there were water-worn stones and

pebbles in them.

There are but few halophytic plants on the sand beaches around

James Bay. Batis maritima occurs in a few places and there is a

considerable growth of bushes of Conocarpus erectus bordering

the shore. A heavy growth of bushes and small trees surrounds

the salt-water lake, just back of the beach, which consists of the

following species : Avicennia officinalis, Cryptocarpus pyriformis,

Discaria pauciflora, and Maytenus obovata. The remainder of

the sandy area, at the base of the mountain, is covered with open

woodland made up largely of trees of Bursera graveolens, and

Erythrina velutina, the last of which were in blossom when we

visited this place in December. There are also open grassy areas

in the woodland covered mostly with Setaria setosa, bushes of

Stewart — Botanical Conditions pn the Galapagos Islands. 331

Telanthera echinocephala, and Lantana pednncnlaris. The last

one of these also occurs on the north and south sides of the bay,

where it often forms dense thickets five or six feet high, covered

in shady places with vines of Asclepias angustissima, and Passi-

flora linearloba.

Except for an occasional bunch of Cereus nesioticus, and a

few other plants, all of which occur in protected places, the re¬

cent lava beds south of James Bay, are practically bare of vege¬

tation below an elevation of 500 ft. Along the edges of these

beds, next to the older lava, however, there are thickets of bushes

which are made up almost entirely of Scalesia atractyloides,

which does not seem to grow in any other situations, here. Bor-

reria ericaefolia is another plant which is found in similar situ¬

ations to the above, and also for some distance out on the lava

beds. Above an elevation of 500 ft. there is a considerable

amount of small vegetation on the recent lava, made up mostly

of Asclepias angustissima, and Polypodium squamatum. This

increases in amount with the ascent and at an elevation of 900

ft., there are small trees of Cereus sclerocarpus, and bushes of

Dodonaea viscosa, and Lipochaeta larcifolia. Quite a number

of ferns are to be found in the old craters and lava caverns

around this elevation among which are : Asplenium, cristatum, A.

formosum, A. sulcatum, Ceropteris tartarea, Nephrolepis bisser-

ata, and N. pectinata. Many of the species which were common

at a lower elevation, occur along the edges of the recent lava beds

here, associated with such mesophytic forms as bushes of Erig-

eron tenuifolius var. tomentosus, and Psychotria rufipes, and

trees of Pisonia floribunda. The presence of ferns and other

mesophytic plants associated with such xerophytes as Bursera,

and Cereus, indicates that the region around an elevation of

1,000 ft., on this side of the island, lies within the transition re¬

gion.

There are several islands of vegetation on the lower part of

the south side of the island which are surrounded by beds of re¬

cent lava. None of these were visited, but they appeared from

a distance to be covered with the usual species found in the dry

regions. Several small mangrove swamps occur along the south

shore.

The tuf aceous region in the vicinity of the Sugarloaf Mountain

is covered with small Bursera trees, Croton bushes, and other

dry-region forms. The Sugarloaf mountain is a large tufa

332 Wisconsin Academy of Sciences, Arts , and Letters.

crater with steep sides, which rises to a height of 1,200 ft. The

sides of the mountain are covered with dry-region forms and ap¬

parently there is but little change in the character of the vege¬

tation from the bottom to the top. The appearance of the moun¬

tain was so uninviting that the top was not visited. The interior

of one of the smaller tufa craters in the vicinity of the Sugar-

loaf was visited, however, and a few halophytic plants, and trees

of Hippomane Mancinella were found growing around the salt

water lake in its interior.

The sides of the mountain east of James Bay are covered with

forests, which to an elevation of 1,000 ft. are composed largely

of trees of Acacia tortuosa, Bursera graveolens, Erythrina velu-

tina, and a few trees of Hippomane Mancinella. The under¬

growth is usually rather open in this region and is made up

mostly of bushes of, Castela galapageia, Cordia lutea, Croton

Scouleri var. brevifolius, Telanthera eehinocephala, Tourne-

fortia strigosa, and Waltheria reticulata forma intermedia.

Occasional trees of Scalesia pedunculata begin to appear

around an elevation of 1,000 ft., but they become larger and

more abundant higher up. Ferns are found abundantly above

1,300 ft. elevation such species as Doryopteris pedata, Polypo¬

dium lepidopteris, P. pectinatum, and P. squamatum being the

most common. Such epiphytes as Peperomia galapagensis, and

Tillandsia insularis are also found. The undergrowth, which is

made up largely of Tournefortia strigosa, becomes thicker than

lower down.

The rolling plateau, which extends from 1,400 ft. to the base

of the main crater at 2,200 ft. elevation, is covered with forests

of Pisonia floribunda, Psidium galapageium, Scalesia peduncu¬

lata, and Zanthoxylum Fagara. The Scalesia trees are the most

abundant in this region, and form true Scalesia forests, as on

some of the other larger and higher islands of the group. The

trees of Psidium galapageium are smaller than at similar eleva¬

tions on other islands where this species occurs. Bushes of

Tournefortia strigosa continue into this region and such other

bushes as Brachistus pubescens, Croton Scouleri var. grandifo-

lius, Erigeron tenuifolius var. tomentosus, Phytolacca octandra,

Psychotria rufipes, Tournefortia rufo-sericea, and Urera alceae-

folia are commonly found, especially towards the upper part of

this region. There are many open areas, in the deeper valleys

between the hills and craters, which are covered with a heavy

Stewart — Botanical Conditions pn the Galapagos Islands. 333

growth of Paspalum eonjugatum. In his Voyage of the Beagle,

Darwin mentions that he found beds of Cyperus, on the upper

part of this island, in which there was a species of water rail.

The Ornithologists of the expedition succeeded in capturing sev¬

eral specimens of this rather rare bird, but in each instance they

were found to be hidden in the thick growth of Paspalum grass,

no beds of Cyperus having been found.

The forest trees, and bushes are heavily covered with ej>i-

phytes around 2,200 ft. elevation and above. Ferns are common

among these, Polypodium aureum, and Nephrolepis pectinata

being the most abundant. Other epiphytes worthy of mention

are: Epidendrum spicatum, Lycopodium taxifolium, and Peper-

omia galapagensis, besides mosses and lichens. There are many

species of terrestrial ferns in shady places and on moist banks.

The Scalesia forests extend nearly to the top of the mam crater

on the leeward side, but on the windward side, which is bathed

almost constantly by the strong southeast trade winds for sev¬

eral months of the year, the trees begin to thin out a short dis¬

tance above the base of the crater and there are none at the top,

although Zanthoxylum persists here as small gnarled bushes.

Bushes of Psychotria rufipes are very common on this side and

around the top. There are many ferns around the top, among

which is Hemitelia multiflora, the only tree fern on the islands.

The top of this mountain was heavily covered with fog at the

time it was visited so that no general survey of the surrounding

region could be made. As near as could be determined the dry

region extends to about an elevation of 1,300 ft. on this side

while the transition region extends to possibly 2,000 ft. varying,

however, at different places.

The steep sides of the mountain, above the lava fields on the

south side of the island, are covered with the usual species found

in the transition regions, a condition which extends up to above

1,600 ft. elevation. There are many ravines extending down this

side, which broaden out occasionally and enclose park-like areas.

In most places these ravines are filled with bushes and trees.

Fruticose lichens are very abundant upon the vegetation here.

Jervis Island.

Jervis lies about four miles oft the south shore of James Is¬

land. It is a small island, not over two miles in diameter, which

rises to a height of 1,050 ft., in consequence of which the sides of

334 Wisconsin Academy of Sciences , Arts, and Letters.

the island are very steep. A pebble beach extends along a por¬

tion of the north side, near which there is a small salt water

lake surrounded on the sides by a small area of level land.

There are three peaks on the island which vary in height from

950-1,050 ft. The flat area near the shore is covered with dark

red soil mixed with bits of lava, but the steep sides have very

little soil on them, except in low places and in crevices of the

lava. The sides in places are strewn wTith small lava boulders.

Low bushes of Cryptocarpus pyriformis grow along the beach,

while around the lake there are small trees of Avicennia officin¬

alis, and Laguncularia racemosa. Tangled thickets of Discaria

pauciflora with very long stiff spines, and bushes of Maytenus

obovata occur just above the lake on the side next to the land.

There are many small trees of Bursera graveolens, and Opuntia

myriacantha scattered over the lower part of the island. The

first of these species gradually disappears higher up, while the

second becomes very much reduced in size, appearing at the top

in a more or less decumbent form. The steep sides of the island

are covered with bushes which consist chiefly of Croton Scouleri,

and Waltheria reticulata. All of the vegetation, above an ele¬

vation of 450 ft., is heavily covered with fruticose lichen indicat¬

ing a somewhat greater amount of moisture than lower down.

The vegetation has a decidedly stunted appearance around the

summit, and with the exception of a single species of fern, Poly¬

podium squamatum, all of the other plants are distinctly dry-

region forms. There are no trees at the top, and the bushes

which occur there, are usually low and more or less prostrate.

Narborough Island.

Narborough, the third largest island in the group, is situated

just west of Albemarle Island from which it is separated by a

shallow channel about two miles wide at its narrowest point.

The northern end of the island is nearly opposite Tagus Cove on

Albemarle Island. A large crater, probably over 5,000 ft. in

height is situated somewhat north of the center, and there is a

gradual slope upward from the shore to the base of it on all sides

but the north. The outer walls of the crater rise abruptly on

the north side, and are almost perpendicular in places. There

is a broad flat plain of old lava at the base of the crater on this

side.

Stewart-Botanical Conditions fin the Galapagos Islands. 335

This island shows more evidence of recent volcanic activity

than any other island in the group. The sides are covered in

most places with deposits of basaltic lava and volcanic cinder

which are practically bare of vegetation in most places. There

are, however, occasional islands of older lava, which were not

covered by the more recent flows, on which there is a consider¬

able amount of vegetation. Some of these islands are quite

large. One was visited on the north side, and was found to ex¬

tend from the base of the crater to the shore, while on the east

and west sides of it there were extensive beds of volcanic cinder

of more recent origin. The older lava was deeply stained

through weathering. There were occasional lava tunnels in the

older lava, the tops of which had fallen in in places leaving

openings into long narrow caverns high enough in places for one

to walk through them in comfort. There was but little soil on

this part of the island in consequence of which the vegetation

was very scanty.

The island was visited for botanical purposes on the north,

and northeast sides. The shores are bordered by low cliffs,

along the north side, which are almost perpendicular in most

places. On this account landing is difficult and even dangerous

except in calm weather. As there is no suitable anchorage on

this side, we visited it in boats, and it was necessary for one

member of the party to remain in the boat and keep it off shore,

while the remainder of the party did their collecting. On this

account our stay here was more limited than it would have other¬

wise been.

Botanical conditions were very discouraging. There are no

trees of any kind on this part unless the few small Burseras

which occur here, could be designated as such. Neither Cereus

or Opuntia is to be found although they both occur on the south

side of the island. There is a scattered growth of bushes con¬

sisting of : Castela galapageia, Cordia Hookeriana, Euphorbia

diffusa, Lippia rosmarinifolia, Scalesia narbonensis, and Wal-

theria reticulata forma intermedia. Vines of Asclepias angus-

tissima, and Cissampelos Pareira were found to be growing on

the bushes in places. Several species of grasses grow from the

lava crevices among which are, Bouteloua pilosa, Cenchrus gran-

ularis, Eragrostis ciliaris, and Paspalum canescens. A single

fern, Notholaena sulphurea, was found at an elevation of about

500 ft.

336 Wisconsin Academy of Sciences, Arts, and Letters.

The part of the island visited on the northeast side, is about

opposite Tagus Cove. There is a small bay at this place around

which there are small mangrove swamps, as there are at several

other places along the east and south shores of the island. Bot¬

anical conditions were even more discouraging here than they

were on the north side, as all of the country in this vicinity was

found to be covered with beds of recent lava on which there was

apparently no other vegetation than occasional specimens of Cer-

eus nesioticus and Cyperus Mutisii.

Mr. Beck succeeded in reaching the top of the mountain, when

he visited the island from the south side somewhat earlier in the

season. He reported it to be heavily covered with vegetation,

among which were ferns and other mesophytic plants. He also

reported a heavy growth of tall grass around the top, which

from his description, must have been Pennisetum exalatum.

There are two lakes inside the crater which are probably 2,000

ft. or more below the rim. The inner walls were covered with

recent lava.

Seymour Islands.

The Seymour Islands, three in number, lie off the northeast

corner of Indefatigable Island of which they probably formed a

part some time in the past. The islands are all low and are

separated from each other by relatively narrow and shallow

channels. South Seymour, the only one of the three visited, is

the largest and lies closest to Indefatigable from which it is sep¬

arated by a channel about one half mile in wddth.

The shores of this island are steep and formed by low cliffs,

except in two places on the west side, where there are sand

beaches. One of these beaches borders a rather large bay which

affords good anchorage for vessels. Back of this bay there is a

flat sandy area of some extent, but otherwise the surface of the

island is covered with large irregular boulders of lava in be¬

tween which there is a scanty light red soil.

The densest vegetation on the island occurs on the sandy area

mentioned above, where thickets of bushes made up of Crypto¬

carpus pyriformis, Discaria pauciflora, and Maytenus obovata

are to be found. In front of these thickets, there is a consider¬

able area along the beach which is covered with Ammophila

arenaria, the only place on the islands where this species has

been found. The remainder of the island is covered with small

Stewart— Botanical Conditions on the Galapagos Islands. 337

Bursera trees, bushes of Castela galapageia, Gossypium barba-

dense, small specimens of Opuntia, Parkinsonia aculeata, and

Prosopis dulcis. Some of the Bursera trees are evidently B.

malacophylla, an endemic species collected on the Seymour

Isands some years ago by Snodgrass and Heller, but whether

or not they were all of this species could not be determined as

the Bursera trees were in the resting condition both times this

island was visited by our party. A few dried leaves was all that

could be obtained of this species.

There is a small salt-water lake near the west side around

which there are a few small trees of Avicennia officinalis. Man¬

grove swamps are probably present along the shore of the chan¬

nel separating the island from Indefatigable, but as they were

only seen from a distance the extent and composition of them

could not be determined. Goats have been introduced upon this

island during the past few years, by the inhabitants of one of the

other islands. They manage to gain a miserable existence from

the scanty vegetation and render the island even more barren

than it would otherwise be.

North Seymour is next in size to South Seymour. It is appar¬

ently covered with a dense growth of Croton, and other bushes.

The middle island is low and small. There were large red-col¬

ored patches of vegetation on it which had the appearance from

a distance of being composed of Sesuvium Edmonstonei.

Tower Island.

This is one of the smaller islands which lies about twenty-eight

miles east of Bindloe. The shores are bordered by cliffs about

forty feet high on all sides except the south where there is a

small bay and sand beaches. The island is only four miles in

diameter and is low in altitude, the highest point on it not being

over 200 ft. above sea level. There is a crater, nearly a half

mile in diameter near the center, but which can not be seen from

the shore or from the surrounding ocean as there is no rim pro¬

jecting above the surrounding country. There is a small salt¬

water lake at the bottom of this crater on the shore of which

there is a grove of trees of Rhizophora Mangle. Small blow¬

holes occur at other places on the island.

Outside of the mangroves just mentioned, there are no other

trees on the island except those of Bursera graveolens, which are

22 — S. & A.

338 Wisconsin Academy of Sciences , Arts , and Letters.

small and heavily covered with lichens. Low thickets of Opun-

tia Helleri occur along the tops of the cliffs and for some distance

back from them, and Cereus nesiticus is to be found in several iso¬

lated spots on the island. The most common bushes are those

of Cordia lutea, Croton Scouleri, Lantana peduncular is, and

Waltheria reticulata forma Andersonii. A few grasses, sedges,

and other herbaceous plants have been reported from this island,

but as our party visited it during the dry season in September,

none of these were found.

Wenman Island.

With the exception of Culpepper, Wenman is the most north¬

ern island of the group. It is nothing more than an immense

rock, about a mile in diameter, which lies seventy six miles north¬

west of Abingdon Island. The main island is surrounded by

perpendicular cliffs on all sides but the north where they are

somewhat broken down so that a landing can be effected and the

upper part reached. In many places the cliffs are several hun¬

dred feet high, and some of them reach practically to the highest

part which probably has an elevation of about 800 ft. There is

a smaller island, to the north of the main one, and separated

from it by a narrow channel. This, however, was not visited by

our party. The channel between the islands is comparatively

shallow and it is likely that an anchorage could be made here.

It was not attempted by our party.

We were unable to remain on this island for more than a few

hours in consequence of this the higher parts were not visited,

botanical collecting being confined to a shelf about 250 ft. above

sea level. The remainder of the island rises several hundred

feet higher.

The only trees found on the island were those of Erythrina

velutina, a small grove of which occurs on the northeast side.

Bushes of Croton Scouleri var. brevifolius occur in dense thick¬

ets and some of the specimens are several feet high approaching

the size of small trees. Low thickets of Opuntia Helleri are to

be found along the tops of the cliffs and hanging over the sides.

Ipomoea Kinbergi occurs commonly on trees. Other plants

found rather abundantly are : Scalesia Snodgrassi, and Telanth-

era Helleri var. obtusior. A few ferns were seen growing in inac-

cessable places on the sides of the cliffs. By shooting into a

Stewart — Botanical Conditions pn the Galapagos Islands . 339

bunch of these enough material was brought down to show that

Nephrolepis biserrata occurs here, but whether or not there are

any other species of ferns can not be said. The small island, to

the north of the main one, appeared to be covered with Croton,

and other bushes.

340 Wisconsin Academy of Sciences , Arts, and Letters.

SOME OBSERVATIONS CONCERNING THE BOTANICAL CONDI¬

TIONS ON THE GALAPAGOS ISLANDS.

INDEX.

Page

Introduction . ' . 272

Abingdon Island . 273

Albemarle Island . 276

Barrington Island . 292

Bindloe Island . 294

Brattle Island . 295

Charles Island . 296

Chatham Island . 301

Culpepper Island . 308

Daphne Island . 308

Duncan Island . 308

Gardner Island, near Charles Island . 311

Gardner Island, near Hood Island . 312

Hood Island . 312

Indefatigable Island . 344

James Island . 327

Jervis Island . 333

Narborough Island . 334

Seymour Islands . 33®

Tower Island . 337

Wenman Island . 338