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(U. S. DEPARTMENT OF AGRICULTURE. of 
BUREAU OF PLANT INDUSTRY -BULLETIN NO. 81. = Gt 


B. T. GALLOWAY, Chief of Bureau. 


EVOLUTION OF CELLULAR STRUCTURES. 


BY 


O. F. COOK anp WALTER T. SWINGLE. 


VEGETABLE PATHOLOGICAL AND PHYSIOLOGICAL 
INVESTIGATIONS. 


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Issurp Aucust 4, 1905. 


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WASHINGTON: 
GOVERNMENT PRINTING OFFICE. 
WOR. <4 Bo-b 


BUREAU OF PLANT INDUSTRY. 


B. T. GALLOWAY, 
Pathologist and Physiologist, and Chief of Bureau. 

VEGETABLE PATHOLOGICAL AND PHYSIOLOGICAL INVESTIGATIONS. 
ALBERT F. Woops, Pathologist and Physiologist in Charge, Acting Chief of Bureau in Absence of Chief. 
BOTANICAL INVESTIGATIONS AND EXPERIMENTS. 
FREDERICK V. COVILLE, Botanist in Charge. 

GRASS AND FORAGE PLANT INVESTIGATIONS. 

W.J. SPILLMAN, Agriculturist in Charge. 

POMOLOGICAL INVESTIGATIONS. 

G. B. BRACKETT, Pomologist in Charge. 

SEED AND PLANT INTRODUCTION AND DISTRIBUTION. 

A. J. Preters, Botanist in Charge. 

ARLINGTON EXPERIMENTAL FARM. 

L. C. Corsett, Horticulturist in Charge. 
EXPERIMENTAL GARDENS AND GROUNDS. 

E,. M. Byrnes, Superintendent. 


J. E. ROCKWELL, Editor. 
JAMES E. JONES, Chief Clerk. 


VEGETABLE PATHOLOGICAL AND PHYSIOLOGICAL INVESTIGATIONS. 


SCIENTIFIC STAFF. 
ALBERT F. Woops, Pathologist and Physiologist in Charge. 


ERWIN F. SMITH, Pathologist in Charge of Laboratory of Plant Pathology. 
GEORGE T. Moore, Physiologist in Charge of Laboratory of Plant Physiology. 
HERBERT J. WEBBER, Physiologist in Charge pf Laboratory of Plant Breeding. 
WALTER T. SWINGLE, Physiologist in Charge of Laboratory of Plant Life History. 
NEWTON B. PIERCE, Pathologist iit Charge of Pacific Coast Laboratory. 

M. B. Waite, Pathologist in Charge of Investigations of Diseases of Orchard Fruits. 
MARK ALFRED CARLETON, Cercalist in Charge of Cereal Investigations. 
HERMANN VON SCHRENK, in Charge of Mississippi Valley Laboratory. 

bP. H. Rovers, Pathologist in Charge of Subtropical Laboratory. 

C. O. TOWNSEND, Pathologist in Charge of Sugar Beet Investigations. 

P. H. Dorsett,@ Pathologist. 

T. H. KEARNEY, Physiologist, Plant Breeding. 

CoRNELIUS L. SHEAR, Pathologist. 

WILLIAM A. ORTON, Pathologist. 

W. M. Scorr, Pathologist. 

JOSEPH S. CHAMBERLAIN,? Physiological Chemist, Cereal Investigations. 


Ernst A. Bessey, Pathologist. ; 
FLORA W. PATTERSON, Mycologist. as S 
CHARLES P. HARTLEY, Assistant in Physiology, Plant Breeding. 


KARL F. KELLERMAN, Assistant in Physiology. 

DEANE B. SWINGLE, Assistant in Pathology. ; 
Jesse B. Norton, Assistant in Physiology, Plant Breeding. ho $ pes O 
JAMES B.RORER, Assistant in Pathology. “ ‘ 
LLoyD S. TENNY, Assistant in Pathology. 

GEORGE G, HEDGCOCK, Assistant in Pathology. 

PERLEY SPAULDING, Scientific Assistant. 

P. J. O'GARA, Scientific Assistant, Plant Pathology. 

A. D. SHAMEL, Scientific Assistant, Plant Breeding. 

T. RALPH ROBINSON, Assistant in Physiology. 

FLORENCE Hepces, Scientific Assistant, Bacteriology. 

CHARLES J. BRAND, Assistantin Physiology, Plant Life History. 

HENRY A. MILLER, Scientific Assistant, Cereal Investigations. 

ERNEST B. Brown, Scientific Assistant, Plant Breeding. 

Lesiirt A. Firz, Scientific Assistant, Cereal Investigations. 

Leonarp L. Harter, Scientific Assistant, Plant Breeding. 

JOHN O. MERWIN, Scientific Assistant. 

W. W. Copsey, Tobacco Expert. 

JOHN VAN LEENHOFF, Jr., Expert. 

J. ArtTuur Le Cierc,e Physiological Chemist, Cereal Investigations. 

T. D. BeckwitnH, Expert, Plant Physiology. 


«Detailed to Seed and Plant Introduction and Distribution. 
» Detailed to Bureau of Chemistry. 
¢ Detailed from Bureau of Chemistry. 


2 


LETTER OF TRANSMITTAL. 


U. S. DEPARTMENT OF AGRICULTURE, 
BurREAU OF PuANntT INDUSTRY, 
OFFICE OF THE CHIEF, 
Washington, D. C., May 31, 1905. 
Str: I have the honor to transmit herewith, and to recommend for 
publication as Bulletin No. 81 of the series of this Bureau, the accom- 
panying technical paper entitled ‘* Evolution of Cellular Structures.” 
This paper was prepared by Messrs. O. F. Cook and Walter T. 
Swingle, and has been submitted by the Pathologist and Physiologist 
with a view to its publication. 
The accompanying plate is necessary to a complete understanding 
of the text of this bulletin. 
Respectfully, B. T. GaLLoway, 
Chief of Bureau. 
Hon. JAMES WILSON, 
Secretary of Agriculture. 


Bisa, _ 


Cis 


PREPAC E. 


Ever since the epoch-making discovery of Charles Darwin there has 
been a steadily increasing influence of the theory of evolution on the 
scientific study and practical utilization of the plants and animals on 
which agriculture is based. The present paper marks a step in 
the further working out of the doctrine of descent, inasmuch as it 
embodies results of an association of the data won in two very different 
fields of investigation; one making the cell its object of study, the 
other occupied with the species. The results herewith presented open 
new views as to the nature of higher animals and plants which can 
not fail to stimulate research and which promise to have great 
economic significance in the determination of the actual and latent 
capacities of the organisms utilized by man. 

A. F. Woops, 
Pathologist and Physiologist. 
OFFICE OF VEGETABLE PATHOLOGICAL 
AND PHYSIOLOGICAL INVESTIGATIONS, 
Washington, D. C., May 8, 1905. 


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SM RMSNE ts. nl Peres se cepcet ia sastsaccesnece ded. od 


29810—No. 81—05——2 


ILLUSTRATIONS. 


PLATE. 


Piare I. Diagram illustrating the network of descent, succession of genera- 
tions, alternating phases, and expansion of the fertilized egg 


TEXT FIGURES. 


Fic. 1. Diagram showing the different types of cell structures and their posi- 
tion in the life history of organisms =. <-.:-..<-.-1.2285 62 eee 

2. Diagram showing the relative importance of the paragamic, apaylo- 
gamic, and haplogamic phases in the life history of various groups 

of organisms 


, 
5 


Page 


26 


18 


a 


B. P. I.—161. V. P. P. 1.—135. 


EVOLUTION OF CELLULAR STRUCTURES. 


INTRODUCTION. 


The value of a new point of view lies in the fact that it permits new 
relations to be perceived. By means of the kinetic theory of evolution 
it has become possible to understand that organic development has 
been carried forward through gradual improvement of the methods of 
descent rather than by environmental causes. Instead of there being 
a law of heredity which tends to keep the individuals of a species 
uniform, or exactly alike, the tendency, especially among higher 
plants and animals, is to maintain, inside the species, a diversity of 
form and structure, most conspicuously manifested in the phenome- 
non of sex. 

This intraspecific diversity is neither accidental nor incidental, but 
of great physiological and evolutionary importance. The interweav- 
ing of distinct lines of descent is necessary to sustain the strength and 
vital efficiency of the individual organisms, and to continue the evo- 
lutionary progress by which the species adapts itself to changing 
environments or enters new ones. Interbreeding is as indispensable 
for the species as for the individual, or even more so, for seedless 
plants continue their individual existences after the coherence of the 
specific group has been lost. 

Normal and long-sustained evolutionary progress is not accomplished 
on single or narrow lines of descent, but is possible only for large 
companies of interbreeding individuals; that is to say, for species. It 
is thus no mere accident, but a fundamental necessity, which brings 
about the association of organic individuals into species and determines 
what might be called the specific constitution of living matter. 
Species are sexual phenomena; they have come where they are only 
through symbasis; that is, as groups of interbreeding individuals, 
traveling together along the evolutionary pathway. 

This interpretation of familiar biological facts is supplemented and 
confirmed by the study of the processes of cell conjugation, which are 
the means of symbasic interbreeding. Among simple organisms con- 
jugation is a periodical incident in the multiplication of equal and 
independent cells. Higher stages of organization were reached by 
the production within the same species of many kinds of cells and the 
building of these into large colonies or compound individuals. There 
was, however, a very early limit to the structures which could be 


9 


10 EVOLUTION OF CELLULAR STRUCTURES. 


built of the primitive, simple type of cells, as illustrated by the fila- 
ment of the lower alga, the vegetative mycelium of the fungus, and 
the thallus of the liverwort. The plant series would have culminated, 
apparently, with the leafy axis of the moss if the basis of organiza- 
tion had not been changed from the primary or simple type of cells to 
the double or sexual type. 

In undifferentiated unicellular or equal-celled (isocytic) organisms 
the successive generations of cells may be thought of as joined into a 
network by an occasional conjugation. The cells at the knots of the 
network are, as we know, double, being formed from the association 
of two nuclei and the accompanying protoplasms. They are often 
strikingly different from the remainder of the cellular fabric of 
descent, and have been given special names, such as oospore, zy go- 
spore, and resting spore. In the first or lowest category of sexual 
organisms only one cell in each generation is double; there is only one 
large bead at each node of the genealogical network. (See Pl. L) 
A second type of organic structure was initiated when an organism 
attained the art of forming two or more of these double cells by divi- 
sion.” It is of such double cells that all the higher plants and animals 
are built. The new type of organization was not merely supplemen- 
tary to the old; it was a new biological invention, giving rise to a new 
category of vitality, which not only outstripped the old type of struc- 
tural organization, but even caused it to be abandoned and eliminated 
as a worse than useless impediment. 

Organisms which were farthest ahead on the primitive basis have 
fallen far behind since the new course of development was opened. 
In such groups as the liverworts, mosses, and ferns the diversity of 
the two types of organic structure is strikingly obvious, and has 
received extensive study for years past under the name of ‘‘ alterna- 
tion of generations.” Ample homologies have been found in the 
highest groups of plants to show that the so-called alteration of gen- 
erations was everywhere in ancestral condition, and that all have 
followed essentially the same history in having abandoned the simple 
type of cell for the double as the basis of structural development. 


“The terminology followed in this paper presupposes for convenience the exist- 
ence of the cellular type of organization common to most animals and plants. The 
conclusions here reached apply with equal force, however, to organisms such as the 
Infusoria among the protozoa, the Siphonocladiacese among the alge, and the 
Saprolegniales and Mucorales among the fungi, in all of which groups considera-. 
ble structural differentiation is attained without any division of the organism into 
cells, Such forms as Caulerpa and Acetabularia among the Siphonocladiaceze reach 
a considerable size and even show a well-marked differentiation into the analogue 
of stem and leaf, rhizome and root, without the enormously expanded thallus being 
divided up into cells at all, although very numerous nuclei arise by subdivision and 
are scattered throughout the cytoplasm. These nuclei could be double, in which 
case such plants would be directly homologous to the double-celled organisms 
described in the following pages. 


INTRODUCTION. ¥4 


That these converging data pointed to something of fundamental 
evolutionary significance has been confidently believed since the publi- 
cation a decade ago of Strasburger’s epoch-making essay entitled ‘* The 
Periodic Reduction of the Number of the Chromosomes in the Life- 
History of Living Organisms,”* but a new evolutionary standpoint 
was required before the larger import of the facts could be perceived. 
The reduction of chromosomes is indeed a striking and unique phe- 
nomenon in the life history of organisms, and it naturally became the 
focus of interest in the rapidly developing science of cytology. A 
new point of view was the more necessary, however, because of an 
unfortunate choice of terms which has undoubtedly tended to prevent 
the perception of the true relations of the facts, as it now interferes 
with a correct description of them. We refer to the characterization 
of the higher, double-celled, spore-bearing ‘* generation” as ** asexual.” 
Appreciating the primitive character of such structures as the pro- 
thallus in ferns, Strasburger asserted that a new ‘‘asexual generation” 
had been intercalated into the life history of organisms. It is now 
perceived that for cytological purposes this is not the whole truth, 
and that for evolutionary purposes it is not true at all. The new 
‘* veneration” was not merely intercalated into the /7fe history of the 
organism; it was intercalated into the sexual process. It is, therefore, 
not asexual, but sexual, and in a higher degree than the so-called sex- 
ual generation. The latter bears, it is true, the cells which conjugate, 
but the former is produced during the actual process of conjugation. 
Organic perfection has been attained, not through the development of 
an ‘‘asexual generation,” but by the lengthening out of the sexual 
process itself; not by abandoning or avoiding sexuality, but directly 
by means of it. 

Among the lower plants the single cell formed by conjugation accom- 
plishes in a brief space of time all the cytological processes which in 
the higher plants come between fertilization and chromosome reduc- 
tion. Sexual fusion is immediate and complete, and takes place during 
a brief period of interruption of the growth and subdivision of vegeta- 
tive cells. If the vegetative fern prothallus is to be termed sexual 
because it produces antheridia and archegonia, the sporophyte is sex- 
ual to the second degree, for it is built of conjugating cells, contain- 
ing, until synapsis and the subsequent reducing divisions, a double 
number of chromosomes, the parental chromatin elements being still 
unfused. However important chromosome reduction may be, it is, 
after all, only a corollary or sequel of the dowbling conjugation. It 
was not the reducing division, but the long postponement of the reduc- 
tion division, which permitted higher types of organisms to be developed 
by means of double, sexual cells. 

A special evolutionary significance was ascribed to the chromosome 


“Strasburger, Edward, Annals of Botany, 1894, 8: 281-316. 


12 EVOLUTION OF CELLULAR STRUCTURES. 


reduction because cytology was approached from the standpoint of the 
somatic tissues of the higher plants and animals. This current imter- 
pretation reverses, however, the historical course of events. The 
reducing division was not an expedient incidental to the adoption of a 
process of sexual reproduction by organisms previously sexless. /¢ 
was not the reduction to fewer chromosomes, but the retention of the 
double number, that constituted the important step in sexual reproduction 
and made possible the evolution of complex higher organisms. It is, 
therefore, not the reducing division, but the doubling conjugation, 
which should constitute the datum point or base line for tracing cyto- 
logical homologies. 


THE ELIMINATION OF THE SIMPLE-CELLED PHASE. 


Chromosome reduction brought about by synapsis, or the fusion of 
the chromatin elements, followed by two special nuclear divisions, is 
not, historically speaking, the beginning of the sexual process, but 
the end of it. Chromosome reduction stands in no special causal rela- 
tion to the subsequent conjugation. The number of cell generations 
formed between synapsis and conjugation differs greatly in the various 
natural groups, and merely shows how far the organism still adheres 
to its old simple-celled life history. Fecundation and synapsis,-as the 
beginning and the end of the sexual process, would seem to be directly 
comparable in all organisms which have developed a double-celled 
sexual phase, . 

From the physiological standpoint, it may be an advantage to dis- 
pense with the simple-celled phase and thus shorten the period between 
the chromosome reduction which marks the end of one conjugation 
and the cell fusion which begins another. Synapsis relieves organic 
fatigue by means of new nuclear configurations, and has been thought 
of as a stimulant of vital activity or energy of growth, the benefit of 
which can be secured for the new double-celled structure by very 
prompt conjugation, as occurs in all the higher plants and animals. 
This consideration would help to explain the organic inferiority of such 
a group as the ferns, which, although they have developed a double- 
celled phase, continue to waste the energy derived from synapsis on a 
worse than useless simple-celled structure. 

In all animals above protozoa this reduction of the simple-celled 
phase has gone so far as to result in its complete elimination, for the 
two peculiar nuclear divisions which occur in rapid succession imme- 
diately after synapsis constitute an essential part of chromatin reduc- 
tion. That these phenomena noted are indissociably connected stages 
in the process of chromosome reduction has been emphasized recently 
by Farmer and Moore,“ who propose the convenient term maiosis to 


«Farmer, J. B., and Moore, J. E. 8. On the Maiotie Phase (Reduction Divisions) 
in Animals and Plants, in Quarterly Journal of Microscopical Science, n. s., No, 192 
(vol. 48, No. 4), Feb., 1905, pp. 489-557, pl. 34 


ALTERNATION OF STRUCTURAL TYPES. 13 


include synapsis and the subsequent heterotype and homotype nuclear 
divisions.“ 
ALTERNATION OF STRUCTURAL TYPES. 


‘‘Alternation of generations” is an expression borrowed from zool- 
ogy; its application to the archegoniate plants has introduced endless 
complexities, and can be justified, after all, only by false analogies. 
Alternation of generations was discovered by Chamisso in a species of 
Salpa, a marine animal belonging to the Tunicates; but here, as well 
as in the traditional zoological example of the Aphides, or plant lice, 
the phenomena have entirely different evolutionary significance from 
the so-called antithetic alternation of generations in the archegoniate. 
Generations or individual life cycles of Salpa and of plant lice, which 
were originally alike, have become different, so that now partheno- 
genetic generations alternate with sexual generations. To make the 
archegoniate plants a parallel instance, it would be necessary to assume 
that what is now called the sporophyte was originally another thallus, 
or something that corresponded to one, which later on became modified 
into the sporophytic ‘‘ generation.” To state the case in this way may 
seem quite superfluous, since nobody has made such a suggestion. 
Strasburger and others have repeatedly declared that the so-called 
asexual generation had been intercalated—that is, added anew—and 
not substituted for something else. This, however, makes it only the 
more obvious that the sporophyte is a generation only in a very loose 
and inaccurate sense, and not because it corresponds to or takes the 
place of any other generation. The simple fact is that, instead of form- 
ing merely one oospore as the result of fertilization, the archegoniates 
have come to form a whole sporophyte or double-celled structure by 
the multiplication and progressive sterilization of potentially sporoge- 
nous tissue, as Bower has shown.? 

Bower’s generalization is, however, only a half truth, since the sterili- 
zation, or, better, the arrest of spore formation of some of the cells, is 
conditioned on the possibility of continued subdivision and growth of 
the fertilized egg, and this can cccur only when there is a definite 


«To recognize, however, as Farmer and Moore do, these two cell generations as a 
distinct ‘‘maiotic phase’’ in the life history of Metaphyta and Metazoa does not 
seem warranted, since chromosome reduction is apparently a mechanical necessity 
resulting from sexual conjugation and is consequently brought about in a practically 

identical manner in all symbasic organisms, from the lowest to the highest. Maiosis 
is rather a connecting link at the node in the network of descent than a distinct 
phase subject to expansion or contraction as organisms mount in the scale of evolu- 
tionary progress. On the other hand it isclear that the two peculiar cell generations 
occurring during maiosis can not properly be classed with the double-celled phase 
that usually precedes or with the simple-celled phase that usually follows, but con- 
stitute a transition stage marking the end of one generation and the beginning of 
another. 

b Bower, F. O. A Theory of the Strobilus in Archegoniate Piants. Annals of 
Botany, 8:343-365. 1894. 


14 EVOLUTION OF CELLULAR STRUCTURES. 


postponement of some stage of sexual fusion, for if the final stage 
is once reached and the chromatin fuses, no further growth is possible, 
and a new generation is inaugurated automatically. When sexual 
fusion is immediate and complete, i. e., when nuclear fusion follows 
close on cell conjugation and is in turn at once succeeded by chromatic 
fusion, no development of the oospore can occur; it simply breaks up 
into four spores. Such was once the fate of the eges of all organisms, 
and such is still their fate in the lower plants. All development of the 
fertilized egg other than a simple splitting into four spores is due 
to an arrest of the process of sexual fusion which permits its expansion 
into a mass of double cells, such as constitute the bodies of higher 
animals and plants.“ It is, however, clear that the effect of such an 
arrest in the process of sexual conjugation and consequent intercala- 
tion of a double-celled phase in the life history of the organism is to 
lengthen the life cycle; it lessens the number of generations instead 
of making more of them. 

Notwithstanding half a century of endeavor, botanists and zoolo- 
gists have not yet found in the higher animals any definite homologue 
of the so-called antithetic alternation of generations discovered by 
Hofmeister? in the archegoniate plants. The whole idea of alternat- 
ing generations must, however, be abandoned and emphasis placed 
instead on the expansion of the oospore or fecundated egg into a 
double-celled phase that comes to occupy a larger and larger part of 
the life cycle as organisms mount higher in the scale of evolutionary 
progress. It then becomes evident that in higher animals (Metazoa) 
the expansion of this phase has gone so far that the simple-celled stage 
has been completely suppressed, and in consequence their life history 
is as free from alternating phases as that of the lower plants, though 
for a very different reason. The lower groups show no expansion of 
the fertilized egg. The higher animals consist of nothing else.¢ 


«1t is clear that the expansion of the fertilized egg could occur in siphonaceous 
algze and fungi without any cross walls forming between the nuclei as they arise by 
subdivision. The mature thallus of an Acetabularia is obviously the enormously 
expanded syngamete and may or may not contain double nuclei. On the other hand, 
the Infusoria may be found to consist of one double cell, the successive cell genera- 
tions not adhering to form a tissue. 

» Hofmeister, W., Vergleichende Untersuchungen der Keimung, Entfaltung und 
Fruchtbildung héherer Kryptogamen und der Samenbildung der Coniferen. Leip- 
zig. 1851. 

¢This fact is obscured, but not negated, by the splitting up during chromosome 
reduction of the egg and sperm mother cells of animals into four gametes, which are 
simple cells, but which are no longer capable of further development unless they 
conjugate. As previously noted (p. 13, footnote a), these two cell generations occur- 
ring during chromosome reduction constitute a transition stage between the old and 
the new generations and can not properly be classed with the simple-celled phase. 

The occurrence of alternating phases in the life history of an organism indicates 
that it is in an unstable evolutionary condition, sinee it has not yet attained the 


SEXUALITY A MECHANISM OF EVOLUTION. 15 


That there are two unicellular stages in the life history of an organism 
should not be allowed to introduce any confusing technicalities. For 
genealogical purposes the spore is quite as much the descendant of the 
antherozoid and the egg cell as it would be if the other tissues of the 
sporophyte had not been intercalated. From chromosome reduction 
to chromosome reduction, from spore to spore, or from egg to egg is 
one generation, and not two. The prothallus is no more mysterious 
than any other piece of ancient history. The ferns were originally 
liverworts, the capsules of which had the good fortune to get roots 
into the ground and keep on growing, but they have not yet learned 
to dispense with their first vain attempt at building a structure on a 
simple-celled basis. 


SEXUALITY A MECHANISM OF EVOLUTION. 


Stress has also been laid upon this supposed alternation of ‘* sexual” 
and ‘‘asexual” generations in the belief that a clew was here to be 
gained regarding the nature of sex and of attendant ‘‘ mechanisms of 
heredity.” But since only one generation is really involved instead of 
two, and since the phase of existence which has been termed asexual 
is in reality the more strongly sexual, it is not surprising that these 
expectations have not been realized. Sexuality facilitates interbreed- 
ing and makes it the more effective by distributing new variations 
throughout the species; it is, in short, a mechanism of diversification and 
of evolution, a fundamental and universal fact which stands squarely 
in the way of the alleged law of heredity under which organisms would 
breed true and be exactly alike. This notion of a uniform and unchang- 
ing heredity,” or of any natural tendency to such a condition of organic 


most effective type of organization. The persistence of a clearly two-phased condi- 
tion in the vascular cryptogams and of a reduced alternation of phases, even in the 
highest algze and flowering plants, is a proof of the extreme slowness of the evolu- 
tionary progress of the plant kingdom. Animals seem to have passed through the 
diphase period of their existence before the dawn of geologic history, and appear in 
the oldest fossil-bearing strata, not only as completely double-celled organisms but 
nighly differentiated ones at that. Not only are there no traces of the two-phased 
progenitors which must have gone before the lowest known fossil organisms, but up to 
now zoologists have not realized the need of postulating such forms at all, and have 
been content to derive the higher animals from merely simpler but always com- 
pletely double-celled ancestors, which, of course, are not primitive. It seems not 
imprebable that the completely double-celled condition has been reached inde- 
pendently by different groups of higher animals, just as it has been approximated, 
though not attained, by the Fucaceze and the phanerogams among plants. The 
animal kingdom does not contain, so far as is now known, a single species that 
shows alternating phases in its life history; it has no counterparts of all that wealth 
of forms which in the plant kingdom bridge the interval from the protophytes to 
the flowering plants. 

a “The modifications introduced into palingenesis by kenogenesis are vitiations, 
strange, meaningless additions to the original, true course of evolution.’’—Haeckel, 
Evolution of Man, vol. 11, p. 460, note 9. 


16 EVOLUTION OF CELLULAR STRUCTURES. 


stagnation, can well be relegated to the limbo of hypotheses which 
have not proved useful. Heredity is not a mechanism or a force; it 
is merely another name for the property of organic continuity or sue- 
cession. ‘There is no more heredity in an organism at one stage of its 
life history than at another. 

Sexual and other diversities inside specific lines are not useless 
morphological complexities or mere failures in the execution of a 
fundamental plan of complete uniformity. Diversity, interbreeding, 
and evolution are physiological factors of the highest importance in 
maintaining vital efficiency. 

Morphologically speaking, sexuality is a specialization of the inter- 
nal diversity of the species, and among plants, at least, it has been 
attained independently in a large number of unrelated natural groups. 
There are grades of sexual differentiation just as there are of organic 
structures. Moss plants and fern prothalli may be sexually differen- 
tiated and the differentiation may occur farther back in the spore 
itself, or even in the sporophyte or double-celled phase, as in the 
flowering plants and the higher animals. Thus in the same species 
there may be two sexualities, one in the simple-celled stage and 
another in the double, and these may have no homology or causal con- 
nection, except as they both serve the same purpose of promoting 
more efficient symbasis. Indeed, the sexuality of the highest types of 
organization is not merely double, but threefold; the individual has 
sex, as a whole; the double cells of which the body is composed are a 
part of a sexual process, and the simple cells which it produces for the 
initiation of a new generation are sexually differentiated. 


TWO TYPES OF DOUBLE-CELLED STRUCTURES. 


That organisms are everywhere associated in species is not because 
of some undiscovered principle or mechanism of heredity; it is simply 
because the interweaving of the lines of individual descent is being 
maintained, without which the specific association would be dissolved 
into indefinite radial divergence and degeneration, as among the varie- 
ties of bananas and other plants long propagated from cuttings. Many 
explanations have been conjectured for the supposed absence of sexual 
reproduction among the higher groups of fungi. From the standpoint 
of a symbasic evolution, however, it becomes evident that the exist- 
ence of true, coherent species among these fungi is a sufficient evidence 
of interbreeding, and hence of sexuality. There is in many groups a 
deficiency of specialized sexual organs, but these are rendered unneces- 
sary by abundant opportunities for direct conjugation among the 
mycelial filaments. 

That the cells of the more complex reproductive tissues of the higher 
fungi are known to have two nuclei, while in the younger mycelium 


TWO TYPES OF DOUBLE-CELLED STRUCTURES. 17 
there is only one, might also have been accepted as proving that con- 
jugation had taken place. This does not mean, of course, that cross- 
fertilization is indispensable for spore production among the fungi, 
but their habits of growth certainly give many opportunities for con- 
jugations between mycelia of different descent, by which the existence 
of compact and well-defined species can be maintained, although the 
peculiar structure of fungous tissues permits extreme variability of 
the size and external form of the fruiting bodies. 

In structural complexity, size, longevity, and other measures of 
organic efficiency the binucleate fungi have an intermediate position 
in the plant series. Their wide distribution and extensive differen- 
tiation into species, families, and orders are evidences of ample oppor- 
tunities in time and environment, so that it is not unfair to explain 
their evolutionary limitations by reference to their peculiar type of 
organic structure. 

Sexual reproduction is accomplished through conjugation or fusion 
of cells, a process which may be divided into three stages: (1) Plas- 
mapsis, the fusion of the cytoplasm or unspecialized protoplasm; 
(2) karyapsis,* the fusion of the nuclei or nuclear protoplasm; (3) 
synapsis, the fusion of the chromatin. The binucleate cells of the 
fungi may be said to have passed the stage of plasmapsis, but kary- 
apsis, or true fecundation, like that of the higher plants and animals, 
has not taken place. 

For the form of sexuality which produces the binucleate cell struc- 
tures of the higher fungi the name apaylogamy is proposed, in allusion 
to the fact that the two nuclei have not yet associated. The higher 
stage, where the nuclei fuse but the chromosomes remain apart, may 
be called paragamy, which implies that the union is still incomplete, 
but that a more intimate relation has been established. These two 
double-celled conditions may be further contrasted with haplogamy, 
the primitive method of undeferred combination of the sexual cells, 
nuclei, and chromatin. 

To the ‘‘asexual generation,” which is not asexual and not a gener- 
ation, the term paragamic phase may be applied among the higher 
plants and animals, the tissues of which are composed of cells with a 


@The etymology of these terms will be obvious to all students of biology, plasma 
and karyon being the familiar Greek renderings of protoplasm and nucleus, respec- 
tively. The other element, ais, signifies a binding or tying together and also a 
mesh or network, a meaning especially appropriate in view of the reticular struc- 
ture of living matter. 

The series might be completed more logically by using the distinctive word 
mitapsis as a substitute for synapsis, which in its etymology is scarcely more than a 
Greek equivalent for the general term conjugation. Mitapsis is derived from siros, 
a thread, and alludes to the threadlike condition assumed by the chromatin during 
the process of chromatic fusion. 


18 EVOLUTION OF CELLULAR STRUCTURES. 


double number of chromosomes. The binucleate structures of the 
fungi may be referred to as the apaylogamic phase. ‘The so-ealled 
‘sexual generation” may be called the Aaplogamic phase in both 
cases. These new terms might not be necessary if words were used 
for descriptive pur- 
. poses only, but in 
the present instance 
they have general 
implications too im- 
portant to be disre- 
garded. 
Haplogamice 
structures are built 
between synapsis 
and  plasmapsis, 
apaylogamic be - 
PLASMAPSIS tween plasmapsis 


Fic. 1.—Diagram showing the different types of cell structures and gnd karyapsis, par- 
their position in the life history of organisms. : 


PARAGAMIC PHASE 


KARYAPSIS SYNAPSIS 


HAPLOGAMIC PHASE 
APAYLOGAMIC PHASE. 


agamic between 
karyapsis and synapsis. Between the three critical points of cyto- 
logical activity there are three intervals, in which the organism can 
pause to gain additional size or numbers by vegetative division of 
cells. The relations between the cell structures and the nuclear 
processes are illustrated by the accompanying diagram (fig. 1). 

No organisms 


have, however, LOWER GROUPS Peete HIGHER FUNGI 4 MOSSES 
structures built in / 
all the three phases. 
The relative impor- 
tance of each phase Vv Y 
in the life histories sic Iain 
of the different pene FLOWERING PLANTS 
natural groups can ‘ ities P 
also be illustrated 
bysimplediagrams, 
as shown in figure 2. 
The relative im- | 
portance of the dif- Fig. Pe showing the eee importance of Be 
ferent phases inthe apaylogamic, and haplogamie phases in the life history of various 


life historv of the groups of organisms, 


various groups of organisms can be represented in another way, as 
is shown on Plate I. The diagrams on this plate show in addition 
a network of descent in its simplest form, composed of successive 
generations linked together at the first stage of conjugation. The 
generations themselves are seen to be composed of alternating 


TWO TYPES OF DOUBLE-CELLED STRUCTURES. 19 


simple and double celled phases in organisms of intermediate evolu- 
tionary rank, and finally the double-celled phase is shown to be an 
expansion of the fertilized egg, which constitutes an increasingly 
large part of the life history as organisms mount higher in the scale 
of evolutionary progress. 

It is thus easy to understand why the two types of double-celled 
structures have very unequal possibilities of organization. Two nuclei 
are evidently better than one, but their association is too slight, appar- 
ently, to gain much of the vital stimulus consequent upon the more 
effective method of conjugation followed by the higher plants, where 
the chromosomes of two fused nuclei lie in juxtaposition in the new 
nucleus. The higher organisms haye not merely double cells, but, 
what seems to be vastly more important, compound nuclei, a more 
advanced and energized stage of the sexual process, which enables them 
to maintain for exceedingly long periods of time the power of growth 
and subdivision.¢ 

The intercalation of the double-celled structure does not change the 
order of nuclear events in cross-fertilization, but it may be said to 
change fundamentally their chronological and physiological relations. 
The true historical sequence of conjugation is plasmapsis, karyapsis, 
and synapsis, but the apparent and practical sequence in the higher 
plants and animals becomes synapsis, plasmapsis, and karyapsis, the 
synapsis which ends one conjugation being followed closely by the 
plasmapsis which begins another. The suspension of nuclear changes 
for vegetative growth no longer occurs between synapsis and plas- 
mapsis, but between karyapsis and synapsis, the double-celled, para- 
gamic structure being built, as already stated, on a new and highly 
sexual plane, that is, out of cells in a state of prolonged sexual union. 

If, as may be supposed, the benefit of synapsis lies in the making of 
new associations of the ancestral chromatin elements, it is obvious that 
the bringing of two such newly energized nuclei together would pro- 
duce a condition which, for want of other words, might be called a 
multiple vital tension. The double-celled type of structure involves, 
therefore, not merely a transfer of emphasis to a new part of the life- 
eycle, but a new and improved sexual process, which raises the bio- 
logical equation to a higher power. From this standpoint it is obvi- 
ous that the morphological diversities of sex have a fundamentally 
important and truly physiological function in building up and main- 
taining the efficiency of the complex organization of the higher plants 
and animals. It is as illogical to ascribe the internal diversity of 


aAs noted before, some organisms, such as Caulerpa and Acetabularia, show a con- 
siderable degree of evolutionary progress, and have not as a matter of fact attained 
the cellular type of organization at all; they may, however, be found to have double 
nuclei and to be very striking examples of the expansion of the fertilized egg. 


20 EVOLUTION OF CELLULAR STRUCTURES. 


species to external environmental causes as to arbitrary mechanisms 
of heredity. 

The extent to which the static concept of a normally unchanging 
heredity has obscured evolutionary thought and investigation could not 
be better shown, perhaps, than by the fact that, notwithstanding the 
great multiplicity of terms which have been proposed for all the 
imagined kinds of variations, no name has been suggested for this nor- 
mal and necessary intraspecific diversity. The deficiency may be made 
good by the use of the word heterzsm for the whole group of phenomena, 
ranging from mere individual diversity to the highest specializations 
of heterism, exemplified by sexes, castes, and polymorphic conditions. 
It is true that the members of a species look alike when compared with 
those of other species, and there may be no harm in ascribing this 
likeness to heredity, but there is nothing to show that this heredity of 
general resemblance has anything to do with evolution except as an 
incidental result. Evolution does not take place between species, but 
inside of them; it is not an znterspecific phenomenon, but 7ntraspecific. 
Its principal factors are heterism and symbasis, not heredity and enyi- 
ronment, as believed by the selectionists, nor heredity and segrega- 
tion, as supposed by the mutationists. 


HEREDITY IN RETICULAR DESCENT. 


The greater efficiency of the double nuclei is, however, only one 
more evidence of the importance of sex as a means of diversity and of 
bringing diverse protoplasms together. The nuclear network of 
chromatin which controls the activities of the cell corresponds to the 
network of descent through which the cell has come into existence. 
Symbasis, or diversity of descent with normal interbreeding, is the 
foundation of the strength and vitality of the organism, because it 
increases the efficiency of the nuclei of the component cells. 

Inbreeding or defective fertilization, on the other hand, would cause 
nuclear deterioration, as so strikingly shown by Maupas in the so-called 
senile degeneration of ciliate Infusoria induced by keeping them too 
long without cross-fertilization. This phenomenon is, indeed, closely 
parallel to senile degeneration, but there is, nevertheless, an important 
difference. In true senile degeneration the vigor of the cells is declin- 
ing because of the absence or long postponement of conjugation. In 
monobasic degeneration, conjugation may take place, but is not effective 
because of insufficient diversity of descent. _Monobasis is the antithesis 
“of symbasis; it means descent without cross-fertilization, on single or 
very narrow lines. The inevitable result is degeneration, with a 
rapidity proportional to the closeness of the inbreeding and the com- 
plexity of the organisms. 

This intimate relation between organic descent and organic structure 
enables us to understand the phenomena of organic succession without 


HEREDITY IN RETICULAR DESCENT. 21 


resorting to abstract principles or to hypothetical mechanisms of hered- 
ity. The network of descent is, as it were, a map showing the alternative 
routes of the developing organism, and permitting normally any combi- 
nation of ancestral characters, as may well result from the endlessly 
varying arrangements into which the ancestral chromatin elements may 
fall at the time of synapsis or chromatic fusion. Twins developed from 
the same ovum would have the same arrangement of chromatin, which 
accords with their close similarity of form, but otherwise there is 
unlimited diversity, even among the simultaneous offspring of the 
same parents. It would seem, therefore, that instead of a mechanism 
of heredity inside the reproductive cells there is an automatic device 
for insuring diversity. The higher the organization the more com- 
plex the descent, and the greater the variety of nuclear configurations 
and the resulting individual diversity. 

Nevertheless, inheritance is not governed merely by chance, nor 
limited even to the infinity of nuclear networks to be made by the 
combinations possible among the ancestral chromatin elements. With 
the greater vitality of interbred organisms is associated also a stronger 
heredity or prepotency of the wild or more broadly symbasic types 
when such are crossed with inbred domesticated varieties. New varia- 
tions, too, appear to have the same effect as diversity of descent in 
lending greater vigor and prepotency. Even mutations, or degener- 
ative variations induced by inbreeding, are prepotent on their own 
plane of symbasis—that is, when crossed only with their own inbred 
relatives—though they are promptly obliterated or ‘* swamped” 
when brought into contact with the broadly symbasic wild type, the 
prepotency of the diverse descent being far greater than that attach- 
ing to the inbred variation. It is the prepotency of variations which 
renders evolution truly kinetic; for the methods of organic descent 
are such as to bring about a spontaneous change of type. The envi- 
ronment often influences the direction of this vital motion, but is in no 
proper sense an actuating cause. “ 

Cells are the units of organization, but species, as groups of inter- 
breeding individuals, are the units of evolution. The causes of evolu- 
tion are not revealed by hypothetical subdivisions of cells into char- 
acter units or determinate elements, but by ascertaining the methods 
of descent through which interbreeding maintains organic strength 
and evolutionary progress. Cells divide themselves, as we know, into 
other cells, and species into other species, but it is only as cells and as 
species that their vital, organic, evolutionary activities are accom- 
plished. /ndividuals vary and mutate, but only species evolve. To 
classify the various stages and functions of organisms under general 
and abstract terms may be desirable, but for evolutionary purposes it 


@Cook, O. F. Natural Selection in Kinetic Evolution, Science, n. s., 19:549. 1904. 


22 EVOLUTION OF CELLULAR STRUCTURES. 


is the network of descent which represents the concrete, significant 
fact, and it is this which can be resolved, if necessary, into its compo- 
nent lines, polygons, or nodes, to furnish units for the calculation of 
quantitative effects of inheritance, as in Galton’s Law of Ancestral 
Resemblances and Filial Regression, under conditions of normal sym- 
basic descent, or in Mendel’s Laws of Disjunction, in hybrids of 
abnormally inbred varieties. 

The recognition of the double character of the cells of the higher 
plants and animals permits many other phenomena of inheritance to be 
understood, though it seems to take us farther than ever from the hope 
of a merely mechanical explanation of the nature of heredity itself. 
If conjugation were concluded immediately, the well-known phenom- 
ena of sterile hybrids would be impossible, the sterility which puts an 
end to their existence being due, as now known, to the failure to 
perform synapsis or chromatin fusion. On the other hand, it may be 
that crosses between narrowly inbred varieties sometimes have the 
power of passing by the period of synapsis without a true fusion of 
the parental chromatin, perhaps in a manner corresponding to that in 
which Thalictrum produces seeds parthenogenetically, by avoiding 
chromosome reduction. The germ-cells might have a preponderance 
of chromatin from one parent or the other, or might even be quite 
unmixed, as claimed for Mendelian hybrids. It is obvious, however, 
that to explain Mendelism in this manner is to admit the essential 
abnormality of the phenomenon. 


SUMMARY. 


It has been held self-evident that there can be nothing in evolution 
except heredity and environment, and it was a simple deduction from 
such an aphorism that differences must all be due to environment, since 
‘heredity would, if nothing interfered, keep the descendants perfectly 
true to the physical characters of their progenitors.” Such heredity, 
however, is a pure figment of the scientific imagination; it is a 
hypothesis which lends us no aid in understanding the facts of 
organic succession. A stereotyped heredity could make nothing new; 
the interbreeding of diverse individuals and the prepotency of new 
variations are the constructive factors, not heredity and environment. 

Symbasis is the method, interbreeding the means, and sexuality the 
mechanism whereby organic evolution has been accomplished; these 
are the concrete and efticient causes of the vital motion of species. 
The association of organisms into species of similar individuals is not 
brought about by a predetermining hereditary mechanism, but by 
symbasic interbreeding. The highest organization has not been 
attained in ‘‘asexual generations,” but in structures completely and 
essentially sexual, built wholly of conjugating cells. There has been 


—-——-—- 


SUMMARY. 23 


no evolution away from sexuality. Long-continued violations of the 
law of symbasis bring only degeneration. 

This interpretation of evolutionary facts opens the way to an ade- 
quate physiological explanation of the significance of sex, and affords 
also a working theory of the chief cytological complications that have 
arisen as a consequence of sex—complications that have hitherto 
rendered obscure the nature of the cell-bodies of higher animals and 
plants. 

The external diversity of organic nature and the internal diversity 
of cells and of reproductive processes take on new and unexpected 
significance. Both are shown to be consequences of sexual specializa- 
tion, without which no evolutionary advance beyond simple-cell colo- 
nies has been possible. More than this, gradations in the perfections 
of the higher double-celled structure are correlated with definite stages 
of evolutionary progress, so that from the structure of an organism 
its kind of sexuality can be deduced. Evolution becomes, in the new 
view, a physiological rather than a morphological process, since the 
methods of descent affect the quality and efficiency of the organism 
even more promptly and fundamentally than they do its external 
form. 


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4) 
ry? EXPLANATION OF PLATE. 

The cireles (©), eights (8), and thetas (©) represent in each case the nucleus or 
nuclei belonging to a cell, and the succession of cell generations is shown by a 
string of nuclei either simple, in pairs (apaylogamic double cells), or fused (para- 
gamic double cells). The half circles (A ©) and quadrants (9 B ) represent the two 
cell generations formed during chromosome reduction. The brackets [ ] represent 
a cell at the period or periods when the organism is reduced to a unicellular condi- 
tion. All the signs for nuclei could be supposed to be inclosed by a cell wall, which 
has been omitted for the sake of clearness. For the same reason only the cell lineage 
leading directly up to the formation of the gametes has been shown, and no account 
has been taken of the enormous multiplication of cells which occurs not only to build 
up the bodies of animals and plants but also to form many gametes. Only a few of 
the numerous cell generations which make up the organisms in question are shown. 


EXPLANATION OF SIGNS. 


Plasmapsis—fusion of the cytoplasm, or unspecialized protoplasm. 
Karyapsis—fusion of the nuclei, or nuclear protoplasm. 


Synapsis—fusion of the chromatin elements. 


p 
K 
sho Heterotypic and homotypie divisions following synapsis. 


Nuclei of haplogamic phase—structures composed of simple cells haying 
O000 nuclei and chromatin elements completely fused. 


9999 Nuclei of apaylogamic phase—structures composed of double cells, each 
having two unfused nuclei. 


Nuclei of paragamic phase—structures composed of double cells having 
SITS) single nuclei containing unfused chromosomes. 


[ ] Cell, at periods where the organism is reduced to a single cell. 


£ d The expanded egg. 


EXPLANATION OF FIGURES. 


Fic. 1.—Lower organism, such as Sphzeroplea, having only simple-celled (haplo- 
gamic) tissues. The fertilized egg undergoes no development beyond merely splitting 
up into four spores when it germinates. 

Fia. 2.—Higher fungus, such as Agaricus or Puccinia, showing alternation of simple- 
celled and double-celled phases, the latter apaylogamic, i. e., with two unfused 
parental nuclei in each cell. The fertilized egg has expanded into a mass of apay- 
logamic tissue. 

Fic. 3.—Moss, showing alternation of a long simple-celled and a short double-celled 
phase, the latter paragamic, i. e., with parental nuclei fused but with their chro- 
mosomes still distinct and unfused. The fertilized egg has expanded slightly into a 
mass of paragamic tissue. - 

Fic. 4.—Fern, showing alternating phases as in moss (figure 3), but with a short 
simple-celled phase and a long double-celled phase, the paragamic phase haying 
developed at the expense of the haplogamic. The fertilized egg has expanded very 
much into a mass of paragamic tissue. 

Fic. 5.—Flowering plant (phanerogam), showing alternation of a very short simple- 
celled phase with a yery long double-celled phase, the paragamie phase having 
developed greatly at the expense of the haplogamic. The egg mother-cell develops 
only one cell (macrospore). The fertilized egg expands into a large mass of para- 
gamic tissue in which the greatly reduced haplogamic phase develops in a semipara- 
sitic manner, it having no free existence. 

Fic. 6.—Higher animal, having only double-celled tissues, the haplogamie phase 
having been completely suppressed by the greatly expanded paragamic phase. The 
egg mother-cell develops only one egg. The fertilized egg has expanded into a large 
mass of tissue. 

O 


26 


* 
a _—_—— 


PLATE I. 


fF Tey oF 
FIG.L— LOWER ORGANISM. 


ee|8) FIG.2. HIGHER FUNGUS. 


FIG.3.-MOSS. 
oP 
pioceocososesosoe(s 


75, 2 
apneeeso C2268 6289048 
‘oes 


“piscscccocscsccsoisfl i: 


nlocosqsacqsecoeocoosososocsoe(s}— 
20, 
‘jpjocooosoosecocosoocsocosossorisie 


0, 


oa 


ISESEESESSSSESESESSSSEESCEOSOI|S 


aa “x 
FIG. 5-FLOWERING PLANT. 


a 


*» 
prloseeeoeesoeoeososesesssesesess 
oe plococecececosesococcosssscssoce[sie 
~ ‘pulecoocoosesoceeeseeecessesossse Px 
oe neseeoseceeoooeoceseoessscesooe om ( 
“preceocescoooosssooesosoeseseee9 SF 5 Oe ae Se es 
oh, FIG.6-HIGHER ANIMAL. 


DIAGRAM ILLUSTRATING THE NETWORK OF DESCENT, SUCCESSION OF GENERATIONS, 
ALTERNATING PHASES, AND EXPANSION OF THE FERTILIZED EGG. 


£- 


‘ 
a 
* 
“a . 
— a . 
‘ a = 

ye 3 : 
. es 
a + a awed GaoaA™. 


~ ae oe : 


PLATE I. 


Bureau of Plant Industry, U. S. Dept. of Agriculture 


5 
“) 


.8 


Bu 


‘VUSVI1V 40 dv 


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gy 
Le YOM INS 
° 


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. f Al SEL 4 
oll 02l o82l oct o9EI oS). 091 o¥91 =. 91 


U. S. DEPARTMENT OF AGRICULTURE. 
BUREAU OF PLANT INDUSTRY—BULLETIN NO. 82. 


B. T. GALLOWAY, Chief of Bureau. 


GRASS LANDS. OP THE SOUTIE ALASKA COAST 


BY 


C. V. PIPER, 


AGROSTOLOGIST IN CHARGE OF FoRAGE PLANT INTRODUCTION. 


GRASS AND FORAGE PLANT INVESTIGATIONS. 


IssuED AuGusr 22, 1905. 


WASHINGTON: 
GOVERNMENT PRINTING OFFICE. 
£5.05. 


BUREAU OF PLANT INDUSTRY. 


B. T. GALLOWAY, 
Pathologist and Physiologist, and Chief of Bureau. 


VEGETABLE PATHOLOGICAL AND PHYSIOLOGICAL INVESTIGATIONS. 
ALBERT T°. Woops, Pathologist and Physiologist in Charge. 
Acting Chief of Bureau in Absence of Chief. 
BOTANICAL INVESTIGATIONS AND EXPERIMENTS. 
FREDERICK VY. CoviLLe, Botanist in Charge. 

GRASS AND FORAGE PLANT INVESTIGATIONS. 

W. J. SPILLMAN, Agriculturist in Charge. 
POMOLOGICAL INVESTIGATIONS. 

G. B. Brackett, Pomologist in Charge. 

SEED AND PLANT INTRODUCTION AND DISTRIBUTION. 
A. J. Prerers, Botanist in Charge. 
ARLINGTON EXPERIMENTAL FARM. 

L. ©. Corpetr, Horticulturist in Charge. 
EXPERIMENTAL GARDENS AND GROUNDS. 

BE. M. Byrnes, Superintendent. 


J. E. RocKWELL, Editor. 
JAMES E. JONES, Chief Clerk. 


GRASS AND FORAGE PLANT INVESTIGATIONS. 
SCIENTIFIC STAFF, 
W. J. SPILLMAN, Agriculturist in Charge. 


A. 8S. Hirescock, Systematic Agrostologist in Charge of Herbarium. 

C. V. Preer, Agrostologist in Charge of Forage Plant Introduction. 

DAviIp GRIFFITHS, Assistant Agrostologist in Charge of Range and Cactus 
Investigations. 

C. R. BALL, Assistant Agrostologist in Charge of Work on Arlington Farm. 

S. M. Tracy, Special Agent in Charge of Gulf Coast Investigations. 

ID. A. Bropig, Assistant Agrostologist in Charge of Cooperative Work. 

HARMON BENTON, Assistant Agriculturist. 

I’. L. Ricker, Assistant in Herbariwn. 

J. M. WestGate, Assistant in Charge of Alfalfa and Clover Investigations. 

Byron Hunter, Assistant in Charge of Pacific Coast Investigations, 

R. A. Oakey, Assistant in Domestication of Wild Grasses. 

C. W. WARBURTON, Assistant in Fodder Plant and Millet Investigations. 

M. A. Crossy, Assistant in Southern Forage Plant Investigations, 

J. S. Corron, Assistant in Range Investigations. 

HaRoLp T. NIELSEN, Epwarp J..Troy, LyMAN E, Carrier, Leroy C. WILson, 
LAWRENCE G. Dopar, Assistants in Agronomy. 

AGNES CHASE, Agrostological Artist. 


2 


‘oN 


LETTER OF TRANSMITTAL. 


U. S. DeparTMENT or AGRICULTURE, 
Bureau or Piant INpustry, 
OFFICE OF THE CHIEF, 
Washington, D. C., May 19, 1905. 

Sir: I have the honor to transmit herewith a paper entitled * Grass 
Lands of the South Alaska Coast,” and to recommend that it be pub- 
lished as Bulletin No. 82 of the series of this Bureau. 

This paper was prepared by Mr. C. V. Piper, Agrostologist in 
Charge of Introduction of Grasses and Forage Plants, Grass and 
Forage Plant Investigations, and has been submitted by the Agrostol- 
ogist with a view to publication. ; 

The four plates accompanying the paper are necessary to a proper 
understanding of the text. 

Respectfully, B. T. GatLoway, 
Chief of Bureau. 
Hon. James Wiison, 
Secretary of Agriculture. 


PREP AGE. 


Since the discovery of gold in Alaska in 1897 continuous calls for 
information concerning the agricultural possibilities of the Alaska 
Peninsula have come to the Department of Agriculture. Much valu- 
able information on this topic has been secured by the Office of Ex- 
periment Stations largely through the Alaska experiment stations 
at Sitka, Kenai, and Copper Center in charge of Prof. C. C. George- 
son, but as the work of these experiment stations was necessarily 
largely local in character, and as it was highly desirable to study con- 
ditions in sections remote from the stations, the Office of Experiment 
Stations requested the Bureau of Plant Industry to send some one to 
explore as large an area of the Alaskan country as might be feasible. 
Accordingly Prof. C. V. Piper, of the Office of Grass and Forage 
Plant Investigations, was detailed to make this exploration under 
the joint auspices of the Office of Experiment Stations and the 
Bureau of Plant Industry. The summer of 1904 was spent in this 
work. The area explored is shown in black on the map constituting 
Plate I. Many interesting facts relating to agricultural possibilities 
in the region covered were recorded, and Professor Piper discusses 
them in the following pages in detail. 

For further information concerning the results of this exploration 
the reader is referred to the Annual Report of the Office of Experi- 
ment Stations for the year 1904. 

W. J. Spruuman, Agrostologist. 

Orrice or Grass AND Forace PLANT INVESTIGATIONS, 

Washington, D. C., April 14, 1905. 


5 


“\ 
CONTENTS. 
eRe erreas ie Ge mes ae Le. SP coe So ee a 
upmumneneron the prass lands. <=. 2 Po a ee oe 
eI Fira ie en Sosa ee. Se ee ene 
Alaska Peninsula and adjacent islands_______-_ Pull ic ees ee eke eee 
Unalaska and the neighboring islands ___._______-_____________._.__- 
8 UB GEL DSIS TTC ERS 2 he NS a ence oe Ns co 
The Yakutat plains - Se SE re he SIN ge OR 
Important factors relating to the agricultural value of the grass Jands. ao 
The abundance and permanence of native fodder plants. =» = -- _§_- 
Seererepperrtst Sian. oe fehl fo 8 EE a at oe i Sea 
TEES Chl Tins CEE ee ce Pa Rae peer ie Re eS A Ee Rene 
IBinomrass. jo 822. tee BS LEY ek Reise sb ova tne men os Coen 
Paver topo. = 22s Soe: Bot ne Oa ee sap A rage ha oR Nea LS 
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Rene naar ey ie Rae Ses Se a ee te ee 
0 OUST Erne Re Soe a SES ar eee ee ane ee ee a 
OTE EET ot Pa ey ae: a A ee 
Hood. value of native Alaskan grasses... 2 2.2 eek 
SirlnncaleuLOn al CTODS ee a2 tee oe ae oe Es ee 
palace alone as a ration for milch cows __-..-.--..-----.-.-- -.2-.-2: : 
masks Oxperionce in stock raising... 2. =.2.-_---. --4..--2-) ese 
(BIG S: 2 SS Se Pa eigen, Sart eke eed, 5 St” Cg em Re ee 
SAE, co ks Bete epee Se ae ene Say oe See ee ee ees 
MHOC PMS UATOTY. 8r-f. hee Wu AG Re eae Ry ge ee RN ees ee 
DRE ne oy Shae. eS =e OT SCD ES Be eine NEL SY Oe ea 
Populawou and available markets (2. <--22-- 252-4. -. 52+: -4--2--2-5-- 
Bre oiicranostransporiatlon 222 9222 )aa22 7S pepe oS) ee SL 
Desirability. of south.Alaska asa home =... 2-22 = 2) 22 2 
OTS AIRE es Se 2 ie OR aie > ee ee See ee 
SRO NST ALT OPO Ni Cops he 2 SE eas a i PA At ee Re ee eee 
LS) Sasa ee Se PEE 6 EIN agi Sag eS Dr Ae 
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Land laws applying to Blasi ee RS Res 2k eee ery, Sean Lalieig han em 
Homesteads - __ _- Rt ea eet eee Rete Sat, Sure AS ty et a SE 
Application for a homestead for surveyed land-__-_-__------------ 
Inceptive rights of homestead settlers _-_--__--------------------- 
Homestead settlers on unsurveyed lands ____ ___---_--- ---------- 
Smithson Wi Prazinis CIStricts= 4°25. 25 eel Shae oe oe ps st 
Homestead claims not liable for debt and not salable_-_._------ 
Soldiers and sailors’ homestead rights -----.-...------------------ 
Soldrers*additional homestead entry. =.= --=-2=----= ----.---=2= t= 


UTS Ug SPSS aa Se eee See Ser eee ale 


99 
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ILLUSTRATIONS. 


Page. 
Piaté I. Mapof Alaskass-. 22.52.20 22-2 ee eee Frontispiece. 
II. Fig. 1.—A view of the flat lands lying at the head of Woman’s 
Bay, Kadiak Island, Alaska. Fig. 2.—Mowing beach rye on 
Kadiak Island, Alaska _....22--=-_. 2.2) 2 38 
III. Bluetop (Calamagrostis langsdorfii), 6 feet ae on Kadiak 
Island, Alaska, July, 1904 _._._...-__...-_32) 2 38 
IV. Fig. 1.—A view of Kadiak, Alaska, Noveniber ies 1903. Fig. 2.— 
A different view of Kadiak, March 26, 1904 _.......-----.-.--- 38 


8 


B. P. I.—164. G. F. P. I.—113. 


GRASS LANDS OF THE SOUTH ALASKA COAST. 


INTRODUCTION. 


A glance at the accompanying map of Alaska (Pl. I) will show . 
that the coast line beginning at Dixon Entrance, in longitude 132°, 
latitude 54° 30’, and extending to Unalaska, in longitude 166° and 
latitude 54°, is nearly in the form of a semicircle, or, rather, of a half 
ellipse, the east and west diameter of which would be about 2,000 
miles and the north and south diameter about half this distance. 
Near the northernmost part of this coast line are two large inlets, the 
eastern one Prince William Sound, the western one Cook Inlet. It 
will be further noticed that islands are very numerous on the coast 
and that the coast line is much indented by narrow inlets or fiords, a 
fact better shown on larger maps. The principal places mentioned in 
this paper are likewise indicated on the map. Officially, the region 
from Mount Saint Elias eastward is known as southeastern Alaska, 
that west of this peak as southwestern Alaska. From an agricultural 
standpoint, however, there is a much better and very marked divid- 
ing line. From Cook Inlet eastward practically all of the lands lying 
near the coast are densely timbered up to an altitude of 2,000 to 3,000 
feet. From Cook Inlet westward, excepting Afognak Island and a 
small portion of Kadiak Island, the lands are devoid of timber, and 
are for the most part grass covered. 

The total area of the coast grass lands is about 10,000 square miles, 
nearly all of which lies between Cook Inlet and Unalaska, a distance 
of about 700 miles. At least one-half of this land would seem capa- 
ble, in time at least, of profitable utilization. From various causes it 
has remained until now practically unused. 

South Alaska is a mountainous country, a great range of snow- 
capped peaks on the mainland paralleling the entire coast. Eastward 
from Cook Inlet great numbers of glaciers arise in the higher moun- 
tains, and many of these rivers of ice extend downward to the sea. 
Westward from Cook Inlet no glaciers reach the sea, although many 
of the mountain peaks are from 5,000 to 8,000 feet high. This strik- 
ing difference apparently depends on a much smaller annual rainfall 


and snowfall. 


9 
29975—No. 82—05 m——2 


10 GRASS LANDS OF THE SOUTH ALASKA COAST. 


In general the lands are hilly, sometimes rising abruptly from the 
seashore, but seldom too steep to afford a luxuriant grass covering. 
More often, however, the hills near the coast are low and rounded, 
with intervening valleys. In places there are wide areas contiguous 
to the coast of from 100 to 1,000 feet elevation and comparatively 
level. Most of the smaller islands, too, have comparatively gentle 
slopes, and either are under 1,000 feet elevation or have but few hills 
reaching above that height. The coast line everywhere is indented 
by numerous bays or inlets, into many of which rivers flow. At the 
heads of these bays there are, as a rule, considerable areas of flat or 
nearly flat lands. Such locations naturally afford the most advanta- 
geous sites for agricultural settlements, especially as these flat lands 
are exceedingly well grassed, and with little preliminary labor can be 
prepared for mowing. 

Where the land is level it is very likely to be wet and covered with 
2 growth of peat moss. Under such circumstances it supports but a 
scanty vegetation. Even on the hillsides this peat moss may become 
established, and where it does so the grasses quickly become less lux- 
uriant. The decay of this moss and of other vegetation results in the 
formation of a humous soil, very retentive of moisture. So deep 
does this humus become that the real soil is often entirely concealed. 
Where it is possible to destroy this moss by burning, the result is 
always a heavy crop of grass or other plants. Most of the land that 
lies at less than 1,000 feet elevation is covered by a most luxuriant 
growth of native grasses. Over large areas these grasses are fre- 
quently 6 feet high, thus furnishing a large quantity of fodder. “On 
the remaining areas, lying at higher elevations or on exposed slopes, 
the grasses are too short to cut for hay, but furnish splendid grazing. 

That grass in Alaska is exceedingly abundant and fairly nutritious 
and that cattle will thrive upon it are facts beyond question. But 
these facts in themselves are not sufficient to enable a prospective 
settler thinking of engaging in stock raising to determine whether 
or not such a venture would be likely to prove profitable. The mere 
abundance of grass of fair quality is not sufficient to insure success 
in stock raising in an isolated region like that under consideration. 

The following statements regarding the Alaska grass lands and 
the factors that have a bearing on their profitable utilization are 
based on as complete a survey as one season’s work would permit, 
together with the facts previously recorded by reliable authorities. 
A detailed report of the conditions actually observed will appear in 
the Annual Report of the Office of Experiment Stations for 1904. 
The present bulletin designs rather to cite these facts in their bearing 
pon the south Alaska grass lands as a desirable field for stock 
raising, 


LOCATION OF THE GRASS LANDS. ah 


THE LOCATION OF THE GRASS LANDS. 


The accompanying map (PI. I) indicates the general location of 
the southern Alaska areas which are covered with grasses. These 
areas differ considerably in detail and are here discussed separately. 


KADIAK ISLAND. 


Kadiak Island, which lies off the mouth of Cook Inlet, is about 100 
miles long by 50 miles wide. It is mountainous in character, the hills 
rising more or less gently from near the seashore to heights of 1,000 
to 3,000 feet. At the end of July, 1904, there was still considerable 
snow at 2,000 feet (Pl. IV), but this is said to be quite unusual. This 
island, like most of the Alaska coast, is much cut into by long, narrow 
bays, into most of which flow streams. The flat lands lying at the 
deltas of these streams are, as a rule, very heavily covered with 
grasses (Pl. II). The slopes also, up to an altitude of 1,500 feet, 
are well grassed, except where there are thickets of alder or willow; 
but these slopes are usually too steep to utilize otherwise than by 
grazing. The total area of these hillside lands is much greater than 
that of the approximately level stretches, in the proportion of at least 
20 to 1. 

On the hillsides the principal grass is bluetop (Calamagrostis 
langsdorfii), which often covers large areas in a pure growth. This 
was exceedingly fine on hillsides burned over in March, by which 
means the old straw and moss were destroyed, thus permitting bet- 
ter drainage and making the soil warmer. In such places this grass 
is often 6 feet high. On the contrary, if the hills are burned over 
in June the fire is likely to kill the grass roots as well as the moss, 
with the result that fireweed usually takes possession of the ground. 

Other grasses than bluetop on the hillsides are relatively unimpor- 
tant, though sometimes considerable areas of Siberian fescue occur, 
and on the higher slopes are a number of low grasses of forage value. 

On the flat lands before mentioned the tall beach sedge (Carex 
eryptocarpa) forms a broad fringe along the shores of the bays and 
sloughs, especially on lands which are occasionally covered by tide 
water. Back of this sedge, beach rye (Elymus mollis) forms a more 
or less broad zone, often mixed with patches of a coarse bluegrass 
(Poa glumaris). In the still drier portions bluetop occupies the 
ground almost exclusively. The three plants mentioned furnish the 
great bulk of forage on Kadiak Island, and indeed on most parts of 
the Alaskan coast, but the bluetop is more abundant than all of the 
other grasses combined. 

Bluetop has slender stems and thin leaves, thus curing very readily 
and making a sweet and palatable hay. Beach rye, on the contrary, 


12 GRASS LANDS OF THE SOUTH ALASKA COAST. 


has thick stems and thick leaves, in consequence of which it cures 
slowly. Beach sedge has a three-sided, solid, pithy stem, and is 
therefore very difficult to dry. All three of these plants grow so 
luxuriantly that they often yield 3 tons of hay or more per acre. 

Of forage plants other than grasses the lupine and fireweed, here- 
after described, are both abundant. In a green state they are readily 
eaten by sheep, but cattle prefer the grasses. 

In portions of the island which have been more or less closely 
grazed for some years it was noticeable that the taller wild grasses 
had largely disappeared, being replaced principally by bluegrass 
(Poa pratensis) and wild barley (Hordeum boreale). Cattle seem 
to be much more fond of the former than of the latter grass, although 
in parts of northern Europe the wild barley is considered a most 
excellent grass. 

All of Kadiak Island, except a small portion in the extreme north- 
east, is practically timberless, as are most of the adjacent islands. 
In the valleys, however, there is usually a small number of cotton- 
woods and willows, and on wet slopes scrub willows and alders form 
dense thickets. Afognak Island, however, which les northeast of 
Kadiak, is quite densely covered with spruce. 


ALASKA PENINSULA AND ADJACENT ISLANDS. 


The whole region to the west of Kadiak Island might briefly be 
described as similar to that island, but entirely deviod of timber, the 
shrubs being more scrubby and the grasses less luxuriant. The 
peninsula itself is very mountainous, and for considerable stretches 
along the coast the hills rise abruptly from the water’s edge. In the 
bays and inlets, however, there are frequently considerable areas of 
comparatively level lands well grassed, though seldom as luxuriantly 
covered as those before mentioned. The islands lying off the coast 
are comparatively low, and some of them are said to be exceedingly 
well adapted to stock raising. Such areas as were examined indicate 
that in general there is a greater variety of forage grasses than to 
the eastward, but most of them are smaller in size. 

At the present time there is a mail steamer plying once a month 
between Valdez and Unalaska. This boat carries the mail, and stops 
at such points as business demands. The population of this entire 
region is exceedingly sparse, and many of the outlying islands would 
probably have to be reached by means of sailing craft. 


UNALASKA AND THE NEIGHBORING ISLANDS. 


Unalaska and the neighboring islands differ on the whole compara- 
tively little from Kadiak Island, though the vegetation as a rule is 


LOCATION OF THE GRASS LANDS. 13 


decidedly less luxuriant. The grasses are much the same in kind, 
although differing in their relative abundance. Some difficulty 
would be experienced on these islands in finding sufficient tall grass 
to furnish winter fodder in case large quantities were necessary, 
though in some of the more sheltered valleys small areas were ob- 
served where the grasses were very tall. There is quite a herd of 
cattle at Unalaska which, according to local reports, receive but very 
slight attention during the winter, only a small quantity of feed 
being cut for them. The principal advantage of Unalaska and the 
neighboring islands would seem to lie in the fact that they are on 
the line of travel of the vessels going to the Yukon and to Nome. If 
sufficient numbers of cattle were raised on these islands, doubtless 
little difficulty would be experienced in finding a market for them at 
the above-mentioned points. Indeed, a Seattle company, which pur- 
poses, among other things, to engage in cattle raising primarily for 
these northern markets, has already begun operations on Akun 
Island. 


KENAI PENINSULA. 


Kenai is the name given to the large peninsula lying between Cook 
Inlet and the Gulf of Alaska. That portion of it on the east side of 
Cook Inlet and north of Kachemak Bay, comprising an area 100 
miles long by 20 to 30 miles wide, is an extensive plateau. Its south- 
ern portion, on Kachemak Bay, lies 500 to 1,000 feet or more above 
the sea level. It slopes mainly to the westward, so that that part 
from Anchor Point northward is but 100 to 200 feet above the sea 
level. Most of this land is timbered with spruce, but there are con- 
siderable areas of grass near Anchor Point, near Homer, and on the 
north side of Kachemak Bay. 

At Homer there is an extensive sand spit, about 4 miles in length 
and from one-fourth to 1 mile across, which supports a good growth 
of several grasses and sedges. Beach rye is the most important and 
most abundant, but red fescue, bluegrass, and seashore grass furnish 
considerable grazing. At the base of the spit the land rises grad- 
ually to the high plateau above, the scattered timber giving the ap- 
pearance of mountain parks.* The open portions of this land support 
a luxuriant growth of bluetop, often 6 feet tall. At a rough estimate 
the open grass lands in this vicinity comprise about 2,000 acres. 

The site of a proposed Finnish colony is on the north side of Kache- 
mak Bay, not far from its head. From the colony site to the head of 
the bay are extensive tide flats, which are mainly covered with 
sedges about 2 feet high. The marshy nature of these lands, together 
with the coarse nature of the forage, makes them of but limited value. 
Undoubtedly they can be much improved by diking. 


14 GRASS LANDS OF THE SOUTH ALASKA COAST. 


The grass lands of the colony site proper consist of about 500 acres 
of excellent land, covered with a luxuriant growth of bluetop. 
These lands lie close to the seashore and less than 100 feet above it. 
Back of these lands are hills 500 to 1,500 feet high, the plateau on the 
top of which consists in part of extensive grass areas. Much of this 
grass is bluetop, often 6 feet high. Other areas are pure growths 
of Siberian fescue. Interspersed with these are several other 
good grasses, but none of them in great quantity. These plateau 
grass lands are apparently very extensive. To render them accessible 
will, however, require the building of roads or trails up to the easiest 
slopes. At Anchor Point there is but little grass land near the sea- 
shore, but on the plateau behind are considerable areas much like 
those just described. The plateau at this point is, however, much 
lower. 

An important circumstance in relation to all of the grass lands of 
this region lies in the fact that they are underlaid with coal, which is 
exposed for miles in the bluffs along the coast. In view of this fact 
it is doubtful if title to the land can be gained by homesteading it. 

At Kenai there are no naturally grassed lands, except the sand 
dunes along the beach and the marshes lying inside of them. The 
dunes are covered principally with beach rye and bighead sedge 
(Carex macrocephala). In the brackish marshes red fescue and sea- 
shore grass are plentiful. Here also is found poison parsnip (Cicuta 
douglasii) 11 small marshes, and there is a record of some native cows 
having been killed by it several years ago. : 


THE YAKUTAT PLAINS. 


The only extensive areas of grass lands known in southeastern 
Alaska are those lying in the river valleys near the coast south of 
Yakutat. Inasmuch as these lands have been several times referred 
to in reports, and as they are now in part accessible owing to the 
building of the Yakutat and Southern Railway, a careful examina- 
tion was made of them. The above-mentioned railway has been built 
primarily to reach the several rich salmon streams flowing into the 
ocean south of Yakutat, it being impracticable to fish them by ap- 
proach from the ocean. This railway is projected to be built to the 
Alsek River, a distance of 45 miles. At present it is built only to 
the Setuck River, 10 miles from Yakutat. 

Practically the whole of this region is an old glacial moraine, com- 
posed of fine gravel, which slopes very gently to the seashore. The 
land close to the seashore is somewhat higher than that lying behind, 
and is heavily timbered. Owing to this strip of higher land most 
of the streams flow parallel to the coast for some distance near their 
debouchments. It is along the valleys of these streams that the grass 


LOCATION OF THE GRASS LANDS. 15 


lands lie, but owing to the flatness of the land and the slight eleva- 
tion above the sea level they are very ill-drained, notwithstanding 
the gravelly nature of the soil. 

i Traveling along these rivers in a canoe one receives the impression 
that the grass is tall and rank on these flat lands. This, in fact, is 
the case on a very narrow strip just along the river banks, where there 
is a fine growth of bluetop (Calamagrostis langsdorfii) and sedge 
(Carex sitchensis Presc.). This strip of tall grass is, however, 
nearly always confined to the immediate banks of the rivers. The 
great mass of the land is covered with a thin layer of bog moss, which 
supports but a scant vegetation of grass and sedges less than a foot 
high. 

' It is a conservative statement to say that fully 80 per cent of these 
Yakutat grass lands are thus scantily grassed. Apart from this 
scant amount of grass, which practically precludes the cutting of 
winter forage, another serious difficulty presents itself in the fact 
that poison parsnip (Cicuta douglasii) occurs quite plentifully over 
all the land that is the least boggy, which, as before stated, is 80 per 
cent of the area. Thus, even if these meadows were used only for 
grazing, great care would need to be exercised in the spring, when 
grass is scanty and the sweet but very poisonous tubers of this plant 
are frequently forced to the surface by the frost. 

While the above statements are true concerning the Yakutat 
meadows as a whole, there are small areas which are exceptional. 
For example, along the lower Ankow River occurs a narrow strip 
of several hundred acres well grassed with silver-top (Deschampsia 
cespitosa) and beach rye (Hlymus mollis) and free from Cicuta. 
Care would need to be exercised in utilizing even this, as the sur- 
rounding boggy lands bear an abundance of poison parsnip. 

Again, the strip of land lying just within the ocean dunes is often 
well grassed with beach rye and red-fescue (Festuca rubra). 

A particularly good area of arable land hes along the railway 
where it reaches the Setuck River. This consists of 3 or 4 square 
miles of gravelly, well-drained, level land, at present looking much 
like a worn-out meadow. It is apparently very well adapted to such 
cultivated grasses as smooth brome-grass and tall meadow oat-grass. 
It will undoubtedly grow all sorts of hardy vegetables. The present 
grass covering is rather scanty, but it is probable that this can be 
greatly increased by cultivation. This particular piece of land is 
well worthy of the attention of homesteaders. 

It is within the bounds of possibility that the larger part of the 
Yakutat plain can be drained and made into fine meadow lands. In 
its present state, however, this land is not adapted to stock raising, 
with the exception of such small areas as above noted. 


16 GRASS LANDS OF THE SOUTH ALASKA COAST. 


IMPORTANT FACTORS RELATING TO THE AGRICULTURAL 
VALUE OF THE GRASS LANDS. 


In determining whether or not the grass lands previously described 
offer a desirable field for settlement, a number of factors that bear 
more or less directly upon the problem need consideration. These 
factors may be discussed in the following sequence: 


(1) The abundance and permanence of the feeds available. 
(2) The possibility of raising forage on cultivated lands. 
(3) The known facts in regard to live-stock raising. 

(4) The available markets. 

(5) Transportation facilities and freight rates. 

(6) The desirability of south Alaska as a home. 

(7) The choice of a location 


THE ABUNDANCE AND PERMANENCE OF NATIVE FODDER PLANTS. 


Live-stock husbandry in Alaska will have to depend primarily 
upon the native plants, supplemented in time, perhaps, by such 
additional ones as experiments shall indicate may compete with the 
native plants, or which upon cultivated land will yield heavily 
enough to be profitable. The most important and abundant of the 
native forage plants are as follows: 

Bluetop.—Bluetop (Calamagrostis langsdorfii) is by far the most 
plentiful tall grass in Alaska, growing along the whole coast. On 
Kadiak Island and the Kenai Peninsula it is especially abundant, 
often being 6 feet high and very dense (Pl. III). It grows with 
special luxuriance on hillsides that have been burned over early in 
the spring. This burning destroys the moss, and thus makes the soil 
better drained and warmer. Bluetop also flourishes on the level 
boggy lands, but prefers a well-drained soil. Owing to its thin 
stems and leaves it cures very readily, and is therefore the usual hay 
grass of Alaska. It is often called redtop, but this name should be 
restricted to the true redtop, a very different grass. 

There are no accurate data bearing on the point as to how well 
this grass will withstand continued cutting, but the general belief is 
that it rapidly becomes thinner in stand. It is noticeable about 
villages where cows are kept that the bluetop is scarce, being replaced 
by other grasses, especially bluegrass and wild barley. The area of 
bluetop is so great, however, that in many places it would be quite 
practicable to manage so as not to cut the same plats two years in 
suecession, which practice would probably maintain the density of 
the stand. 


Beach rye—Along all the quiet shores and inlets of Alaska, 
wherever there is low land near the beach, there is a strip of beach 


rye (Llymus mollis) occurring just above high-tide level. Some- 


NATIVE FODDER PLANTS. 17 


times this strip is only a few feet wide, but on the low level lands 
near the heads of fiords there are often large areas of it 3 to 5 feet 
high (Pl. II, fig. 2). One patch of it examined had been cut the 
year previous, and on this the stand was scarcely half as dense as on 
neighboring pieces which had not been cut. This observation ac- 
cords with the experience of others. 

Where sand dunes occur on the coast, as at Kenai and near Yakutat, 
beach rye is an important sand binder. In such locations it is often 
very different in appearance from that found in other situations, 
the heads being short and thick. This is the result of infestation by 
a parasitic worm. 

Bluegrass.—The true Kentucky bluegrass (Poa pratensis) 1s com- 
mon all along the Alaska coast, where if thrives to perfection. It 
shows a tendency to occupy the ground where closely grazed, and 
cattle exhibit a marked preference for it. Several closely allied 
species also occur, and it is an important fact that they persist and 
increase where other grasses disappear, which seems to insure the 
permanence of pasturage of a high quality. 

Silver-top.—The very nutritious grasses known as_ silver-top 
(Deschampsia cespitosa and PD. bottnica) occur in some abundance, 
especially in gravelly soils, whether on the hillsides or near the sea- 
shore. Owing to their stems being nearly leafless they yield but 
little hay, but the numerous fine basal leaves furnish most excellent 
forage. 

Siberian fescue.—Siberian fescue (Festuca altaica) makes large 
tussocks, especially in gravelly soil and in open timber up to 1,000 
feet elevation. In such locations it often makes a nearly pure growth 
It seems to be fully as nutritious as the well-known sheep fescue, 
but is a much larger grass. 

Sedges.—Two tall species of sedge, Carex cryptocarpa and C. 
sitchensis, in places make dense stands 3 feet high or more, especially 
in wet soil; in the case of the former, more especially in tidal 
marshes. Considerable quantities of this sedge were cut for hay 
near Kadiak, and it is said to furnish excellent feed. These sedges 
are both quite smooth and soft, unlike most others. 

Alaska lupine.—The blue-flowered plant known as Alaska lupine 
(Lupinus unalaschensis) is quite tall, often 3 feet high, and some- 
times occupies large areas almost to the exclusion of other plants. 
It is thick leaved and rather fleshy, and is the only leguminous plant 
that is really abundant in Alaska. Sheep eat it readily. Should 
it prove palatable as well as nutritious to cattle the problem of a good 
winter ration for milch cows would be considerably simplified. Ex- 
periments with it as silage, both pure and mixed with grass, are much 
to be desired. 


29975—No. 82—05 


3 


18 GRASS LANDS OF THE SOUTH ALASKA COAST. 


With the exception of this plant the only legumes of forage value 
in the grass regions are two species of wild pea, both of which, 
unfortunately, are rather scarce. 

Fireweed.—The well-known plant called fireweed (E'pilobium an- 
gustifolium) often occupies the ground to the exclusion of others, 
especially where the land has been burned over in summer and the 
grass roots thus destroyed. Sheep seem fond of it. It is possible 
that this plant may prove profitable as silage, at least when mixed 
with grasses, but no tests with this end in view seem to have been 
made. Its great abundance at times makes such a test desirable. 


There are three possible ways of preserving the above-mentioned 
plants for winter feed. The more easily dried—as bluetop and blue- 
grass—may be made into hay. Continued sunshiny weather on the 
Alaska coast is not to be depended upon, so that haymaking is 
accomplished only with much uncertainty. Where one needs but a 
small amount of fodder, little difficulty is experienced in select- 
ing the few necessary sunshiny days. Where, on the contrary, one 
needs great quantities of winter feed, haymaking is impracticable. 
Resort in such cases must be had either to brown hay or to silage. 
Brown hay is simply half-cured hay, made by stacking the grass 
green or half dry—really a compromise between hay and silage. 
Sometimes salt is scattered over the layers while it is being stacked. 
It is more or less used in all countries where haymaking is difficult. 
While analyses show it to contain practically as much nutriment as 
hay or silage, cattle are not eager for it, and it can be considered 
only an emergency feed. 

Unquestionably when large quantities of winter forage are needed 
for stock, silage must be depended upon, and undoubtedly, all things 
considered, it will be the most satisfactory feed. Practically the only 
Alaska forage plant thus far used as silage is beach rye, and the 
experiences with this plant of Prof. C. C. Georgeson, special agent 
in charge of the Alaska Agricultural Experiment Stations, and of 
others who have grown it, show it to be both palatable and nutri- 
tious. In all probability other Alaskan grasses, and perhaps other 
plants, will be found to be quite as satisfactory. 

Where timber is available silos may be constructed of logs, like 
the one at the Sitka Experiment Station. This silo has the advantage 
of enabling a man to utilize his own labor. On the other hand, the 
material for stave silos can be secured at very reasonable prices, and 
this doubtless is the best silo to use in the timberless regions. 


FOOD VALUE OF NATIVE ALASKAN GRASSES. 


Chemical analyses have been made of the principal Alaskan grasses, 
and while these can be properly interpreted only in connection with 


CULTIVABLE FORAGE CROPS. 19 


digestion experiments, their comparison with the analyses of stand- 
ard grasses furnishes some measure of their value. 


Analyses of Alaskan grasses (air-dried samples taken when in flower). 


F | : Nitro- | Crude 
Species. Water. | Protein. Fat. | gen-free fiber Ash. 
| } extract. 


| | 
Per cent. Per cent.| Per cent.| Fercent. Per cent.| Per cent. 


Calamagrostis eengedon ee ae 7.18 4.58 | 1.03 | 40. 37 2.94 3.90 
Carex cryptocar. Z (Sete) Ses: 3s. 5. 85 10. 32 | 2.12 45.34 25. 72 10. 65 
Elymus mollis (Beach rye) ------------- | 11. 92 | 12.71 2.26 35. 29 | 30. 31 7.51 
Phleuwm pratense (Timothy) ----..------ 8.59 | 8.94 2.14 45.69 30.06 4.58 
Poa pratensis (Bluegrass) -----.-------- 8.11 | 8.94 | 2.04 | 41.45 | 34. 24 5. 22 
Deschampsia bottnica (Silver-top) - ---- 8.75 7.44 2.07 | 47.05 31.54 4.15 
Calamagrostis aleutica ____--.---------- 8.33 10.00 1.37 37.89 38.89 4.52 


| 


Analyses of standard grasses for comparison. 


Nitro- | | 
Species. Water. Protein. Fat. gen-free Candé Ash. 


extract. 


| Percent. Percent. Percent. Per cent. Per cent.| Per cent. 


Poa pratensis (Bluegrass) ........------ 17.44 10. 80 3.45 46. 10 22.09 7.35 

Agrostis alba (Redtop) .......--.....-..| 14.30 Sa BBO aT er | 5.90 

Phleum pratense (Timothy) .---..-.--_- 15. OL 6.01 3.01} 41.90 29.59 | 4.48 

Dactylis glomerata (Orchard grass) __- 14. 30 7.34 2.28 47.08 23.58 | 5.42 

Deschampsia ceespitosa (Silver-top) -- 14. 30 . O4 1.06 37.20 29.03 9. a 
Oo. 


Calamagrostis canadensis (Bluejoint) - 6. 87 11.19 3.45 35. 82 37.18 


The analyses of the Alaskan grasses were all made by the Bureau 
of Chemistry of the Department of Argiculture, and with the excep- 
tion of the first three, from material collected in 1904, were originally 
published in Bulletin No. 48, Office of Experiment Stations. The 
other analyses have been compiled from various authorities. 


CULTIVABLE FORAGE CROPS. 


The experiences of a number of individual investigators, as well as 
the tests made at the Sitka and Kenai experiment stations, throw a 
good deal of light on the possibility of growing fodder plants and 
forage crops on cultivated land. Much more testing is necessary, 
however, before some of the conclusions which at present seem prob- 
able can be considered demonstrated. 

In the way of grasses the tests made at Sitka by Professor George- 
son on muck soils showed tall meadow oat-grass to be the most prom- 
ising. Tall fescue, bluegrass, meadow foxtail, and redtop did fairly 
well, while orchard grass, timothy, and Italian rye-grass were not 
promising. From observations on a number of these-and other 
grasses introduced by chance, some rather definite conclusions may 
be drawn. Timothy is more or less abundantly introduced at various 
places on the coast, but does not as a rule thrive very well, being often 
inferior in size to the native mountain timothy, It is altogether 


20 GRASS LANDS OF THE SOUTH ALASKA COAST. 


probable, however, that a variety of timothy suited to the conditions 
might readily be secured by selection, as chance specimens of the 
plant seen were very fine. The success of such a selection, however, 
will largely depend on the possibility of growing seed in Alaska. 

Among other useful grasses that have become accidentally intro- 
duced and show marked adaptability to the conditions are redtop, 
rough-stalk meadow grass, bluegrass, and fowl meadow grass. 

White clover thrives everywhere along the coast and is an aggres- 
sive plant. Red clover and alsike are not promising and alfalfa 
does not thrive. 

In the way of cereals, the earliest varieties of oats and barley will 
mature for two or possibly three out of five seasons. Of course, such 
a crop is not entirely lost if the grain fails to mature, as it can be 
utilized as hay or silage. On this account it will probably be wisest 
to grow the crop mixed with field peas, as such a mixture will make 
excellent silage, whereas oats alone could only be preserved as hay, a 
difficult thing to do so late in the season. It is to be clearly under- 
stood that under present conditions it is unnecessary to plant any 
cultivated ground in such crops as grass, or perhaps even legumes. 
The above facts are of value simply as indicating what well-known 
forage plants will thrive, thus to some extent showing the future 
agricultural possibilities of Alaska. 


SILAGE ALONE AS A RATION FOR MILCH COWS. 


The writer has been unable to find any published data on results 
obtained by feeding milch cows nothing but grass silage. Presuma- 
bly the best of results would not thus be obtained. 

In order to obtain some light on the subject, Dr. James Withy- 
combe, director of the Oregon Experiment Station, was requested to 
conduct such a test. The results of his experiment are reported as 
follows: 

The silage test was made on a nonbreeding Jersey cow which freshened in 
February, 1902. In January, 1904, this cow was fed largely on silage, with a 
moderate amount of mill feed and light ration of hay as a preliminary prepara- 
tion. From February 1 to April 30 she was fed wholly on corn silage and a light 
ration of ground oats daily. She consumed during the ninety days’ feeding 
3,785 pounds of corn silage and 270 pounds of the oat chop. The following table 
shows variation in weight and her production : 


| Aver- 
Date. Weight. Milk. | age Fat. Date. Weight.) Milk. | age | Fat. 
| test. | test. | 
} eee Se 
Lbs. Lbs. | P.ct.| Lbs. | Lbs. Lbs. | P.ct. | Lbs. 
December 1... 955 | 196| 5.8] 11.86 || March1 ...... 925 195 h.8| 11.31 
January 1----- 945 | 199 | BO)" AL, 41) April ba 890 221 5.5 12.15 
February 1 -__. 905 | 178} 5.7} 10.15 || April 30....... 880 | conch nulsads sane eae 


| 


ALASKAN EXPERIENCE IN STOCK RAISING. 21 


The cow was in good condition at the close of the experiment, which indicates 
- that silage may with safety constitute a large portion of the ration of a dairy 
cow. | ‘ 

This experiment was undertaken at the suggestion of the Government agros- 
tologist to determine in a measure if it were practicable to winter cattle in 
Alaska on grass silage. 

The 3 pounds of ground oats were fed daily for the purpose of bringing the 
corn silage up to a protein standard equaling that of mixed-grass silage. 


Protein percentage of feed consumed. 


SSIS DUPER.) (ON EE ee 2 Se ee eee = 2 ee ee 11. 56 
TNE ADL «SAG Tg eth ss a ek as ee ee 2 a ee ee ee ee 1.58 
Protein percentage of grass silage_________ (approximately) 2. 72 


Average amount of total protein consumed daily in 42 pounds of corn silage 
and 3 pounds of ground oats, 1.01 pounds. Approximate amount of protein con- 
tained in 40 pounds of grass silage, 1.08 pounds. 

It will thus be seen that this test indicates that cattle can be successfully 
wintered on grass silage and that dairy cows may be expected to yield a reason- 
able amount of milk when fed exclusively on this feed. 


ALASKAN EXPERIENCE IN STOCK RAISING. 


Hogs.—A few hogs were seen at various Alaska villages. They 
are fed refuse, and graze on various succulent plants when obtain- 
able. They are very fond of wild rice, the bulb of a lilylike plant 
(Fritillaria kamtschatica), which, however, is not very abundant. 
Unfortunately these animals are prone to feed on fish offal and other 
sea refuse, and as a consequence their flesh has a disagreeable flavor. 
Unquestionably there is too little feed adapted to hogs to make their 
raising profitable in Alaska. 

Goats.—Angora goats have been tested by the Alaska Commercial 
Company at Kadiak and by Rey. C. P. Coe at Wood Island. 
According to Mr. Washburn, formerly resident superintendent of the 
Alaska Commercial Company at Kadiak, the company had a few 
years ago about 50 head of these animals on Ukamak Island, near 
Kadiak, which were entirely self-sustaining, increasing about 60 per 
cent each year. The mohair is said to have been good, both in quan- 
tity and quality. 

Rey. C. P. Coe, of Wood Island, has several head of Angora goats 
which have passed the last two winters with but little care. This 
year his herd has shown very satisfactory increase, and no difficulty is 
anticipated in wintering the kids. A large part of their feed is 
derived from willows and other browse, and where this is abundant 
the animals need but little feed in winter. Owing to their tracta- 
bility and the ease with which they are kept, especially where browse 
is abundant, Angora goats should prove most useful animals both 
for the natives and for whites. 


22 GRASS LANDS OF THE SOUTH ALASKA COAST. 


Sheep husbandry.—Two definite attempts have been made to 
establish sheep raising in south Alaska, though small numbers have 
been kept at various points for short periods. The first attempt was 
made by the Alaska Commercial Company, which in 1883 imported 
a band of about 300 sheep from California. Unfortunately no accu- 
rate record of this experiment is available, and the accounts of 
various persons differ considerably. Many of the sheep died the 
first winter, according to some reports from lack of shelter, according 
to others from scab. The remainder were kept on a small island 
near Kadiak, where the only shelter was a small grove of spruce, but 
in winter they were usually transferred to new grazing grounds 
where they could feed on the tall, dry grass. In very severe weather 
they were sometimes sheltered and fed hay. These sheep are said 
to have yielded about 5 pounds of excellent wool per head each year, 
and the annual increase is reported to have been about 60 per cent of 
the adult animals. No particular care was given them, and the last 
were slaughtered about six years ago. The venture, even excluding 
the loss of the first winter, seems not to have been profitable. 

The only sheep now in Alaska are on the ranch of the Frye-Bruhn 
Company, near Kadiak, who have about 80 head. These sheep are the 
remnant of 9,000 which were shipped in from Oregon in 1902 and 
1903, the remainder having perished. At first sight it would seem 
that this appalling loss of more than 98 per cent was conclusive eyi- 
dence that sheep raising in Alaska is not lkely to prove profitable. 
Inquiry into the causes of the mortality do not bear out this conelu- 
sion necessarily. About 500 of the sheep were drowned in March, 
1903, by being caught at the head of a narrow cove by the incoming 
tide. One hundred and fifty head were lost by becoming frightened 
and jumping over cliffs. The*rest of those that died succumbed to 
scab, which broke out in January, 1903. Owing to lack of shelter it 
was then impossible to treat them by dipping, as that would practi- 
cally have been equivalent to killing them. The result was that all 
but 80 died of the disease. Thus all the mortality was due to causes 
entirely preventable. It was interesting to learn that several head of 
these sheep which ran wild survived the winter without care, and the 
writer was informed by trustworthy witnesses of other cases of this 
kind. In the light of present knowledge it is difficult to say whether 
sheep can be profitably raised in southwestern Alaska. 

In regard to the two attempts which have been made, it is 
noteworthy that in both instances the animals were shipped from a 
comparatively warm and dry climate to one cool and notably wet; 
furthermore, that none of them perished from any cause directly 
connected with the Alaska conditions. 

There are, however, some further difficulties in connection with 


ALASKAN EXPERIENCE IN STOCK RAISING. 23 


sheep raising in Alaska which need careful consideration. It is the 
general opinion in Kadiak that in an ordinary winter sheep can not 
safely be left without care after the beginning of January. Indeed, 
many would place the time a month or six weeks earlier. New grass 
never appears before May 15, and often not until June 1. Therefore, 
under the best of conditions, sheep will need four and a half months 
of feeding and shelter. The superintendent of the Frye-Bruhn ranch, 
after one winter’s experience, thinks that feed and shelter should be 
given for a longer period than that mentioned. 

Another serious difficulty hes in the lateness of the lambing season. 
It is generally agreed that lambing should not take place before 
June 1. The lambs will need shelter and feed by December 1 or 
earlier, unless one takes serious chances of losing many. 

Whether sheep raising could be made profitable at present under 
such conditions remains to be demonstrated. The mere fact that 
sheep in small numbers have wintered without care is no proof that 
successful sheep husbandry can thus be carried on, nor even that one 
or two months’ feeding will suffice. The risks involved in such a 
procedure are too great to warrant a careful stock raiser in taking 
any chances. 

Destructive wild animals are no menace to sheep raising on the 
islands. Eagles may destroy a few lambs, but these birds are easily 
exterminated. Kadiak bears are too scarce and too easily destroyed 
to merit consideration. On the mainland, however, both wolves and 
brown bears may prove troublesome. 

In the hght of present knowledge one is safe in saying that sheep 
can be raised on the Alaska coast 1f adults are given five months’ feed 
and shelter and the lambs a month more—this with the ordinary 
sheep of the western ranges. With more hardy breeds better adapted 
to the conditions the outlook for success would be better. It need 
hardly be said that extreme caution should be taken to import only 
perfectly healthy animals. The great mortality caused by scab and 
the great danger of such a disease as foot-rot in a damp climate de- 
mand that extreme care be taken not to introduce these diseases. 


Cattle-—Cattle have been raised at nearly all the Alaskan coast 
settlements ever since the Russian occupation. Some of the original 
stock, according to local tradition, is still represented in the band of 
cattle at Nannilchuck. These are small animals, but said to be very 
hardy. Nearly all of the cattle kept near the villages are milch cows, 
mostly grades, but a number of Holsteins and Jerseys were seen. 
When owned by whites the animals are given shelter and feed for 
about five months. When they belong to the natives they are forced 
to exist through the winter with little or no care, eking out an exist- 
ence by feeding on browse and seaweeds. No accurate data could be 


24 GRASS LANDS OF THE SOUTH ALASKA COAST. 


gathered concerning the amount and character of the milk yield, but 
it was universally said that the milk is most excellent in summer, and 
good in winter when the animals are properly fed. It is unfortunate 
that no accurate records could be obtained as to the winter yield of 
cows fed only on native hay or silage. 

Several herds of beef cattle have been successfully maintained in 
the neighborhood of Kadiak. The experience of the Alaska Com- 
mercial Company is thus summarized by Mr. Washburn, the former 
superintendent at Kadiak: 

We have bred stock on the islands.of Kadiak, Ukamak, and on Long Island. 
On Long Island we have about 40 head of cattle. These cattle are fed from two 
to six weeks each winter. The remainder of the time they have been able to 
get their own subsistence. During occasional winters we have carried our stock 
through with no feeding. We have had very good increase from them, and 
should say that the percentage of calves raised from the breeding cows is about 


75. The cattle on this island have not been housed except during the short 
period when we were obliged to feed them. 

On Ukamak Island we have a herd of about 20 head, which are entirely self- 
sustaining. We have not found it necessary either to feed or shelter these cat- 
tle during the winter season, and the increase has been fully as good as that of 
the herd on Long Island. 

On Kadiak Island we have not kept any stock cattle, but only a herd of dairy 
cows and some working horses. These we have, of course, fed regularly during 
the winter season for about five months. We are able to cure sufficient hay on a 
lot we have leveled, and we have used the only mowing machine in western 
Alaska. We have obtained very good results from feeding the Alaska hay to 
both cows and horses, and find that they require no more grain when fed this 
hay than when we feed hay imported from California. 

The Frye-Bruhn Company, of Seattle, began operations near 
Kadiak in July, 1903, importing about 200 head of beef cattle, mostly 
Herefords. Owing to unpreparedness and inexperience, about 140 
head of this number were lost during the first year. Most of these 
were killed by falling over cliffs. Owing to the fact that the earliest 
grass appears on the steep southerly slopes, the cattle crowded in such 
places; in some instances the sod, loosened by the frost, gave way and 
precipitated them over the cliffs. In other cases the cattle used their 
horns when crowded, the wounded ones losing their foothold in’ 
endeavoring to escape. As precautions, more care is taken in select- 
ing the early feeding grounds and the cattle have been dehorned. 

The common experience of cattle owners in Alaska has been that 
the animals fatten readily on the grass in the spring, and remain 
in good condition without care until late in the autumn. Some 
Herefords slaughtered at Kadiak in July furnished beef of remark- 
ably fine quality. 

From the experience had at the Kenai Experiment Station, oxen 
keep in good working condition all winter on no other feed than 
native grass hay and silage, and the limited experience of others 


POPULATION AND AVAILABLE MARKETS. 95 


has given similar results. It is not probable, however, that animals 
will remain fat on such feeds alone. 

Nothing has been done up to the present time in the way of intro- 
ducing breeds that are likely to be especially adapted to the peculiar 
conditions. It is highly probable, as has been pointed out by Pro- 
fessor Georgeson, that long-haired hardy breeds like the Galloway 
or the West Highland cattle will prove much more successful than 
breeds adapted primarily to a drier and warmer climate. 


POPULATION AND AVAILABLE MARKETS. 


No very accurate data are available as to the present population of 
the Alaska coast towns and villages, which furnish the only markets 
close to the grass lands. The population of the principal towns 
along the coast is approximately as follows: Sitka, 1,500; Valdez, 
1,000; Seward, 500; Kadiak, 50; Unalaska and Dutch Harbor, 
600. The total population from Valdez to Unalaska, inclusive, is 
about 8,000, of whom less than one-half are whites. From Valdez 
to Sitka, excluding the former, the population is perhaps 4,000, about 
half of them white. Thus the coast of Alaska from Sitka to Una- 
laska provides a market population at present of not more than 
6,000 people, as no market for meat or dairy products can be expected 
so far as the natives are concerned. 

No account is here taken of the towns lying along the interior 
channels in southeastern Alaska, whose populations aggregate per- 
haps 8,000 whites, though a portion of this market could perhaps be 
reached. 

Skagway and Valdez are the principal south Alaskan points which 
supply the interior, and consequently are of especial importance in 
considering markets. 

A considerable market for beef and dairy products could perhaps 
be established by shipping from Unalaska to the population of the 
Nome district and the lower Yukon. Unalaska is on the line of 
transportation from Puget Sound to Nome and the Yukon River, 
though at present few of the vessels stop there. 

Thus the present available markets in Alaska for live-stock 
products are very limited. The supply for these markets at the 
present time is shipped from Puget Sound. 

It is evident, however, that it is possible to raise in Alaska far more 
produce of this kind than the local markets can consume. The only 
other-markets that can possibly be reached are those furnished by the 
cities of British Columbia and of the State of Washington. Freight 
rates are at present, and perhaps will be for some time to come, such 
that dairy products and wool are the only articles that could profit- 
ably be shipped to such distant ports. 


26 GRASS LANDS OF THE SOUTH ALASKA COAST. 


No predictions can here be ventured concerning the future devel- 
opment of south Alaska. The present resources are mainly furs, 
fisheries, and mines. The fur industry is becoming less and less 
important. The fisheries are already highly developed, but are 
‘rapable of considerable increase. The mines undoubtedly will be- 
come more and more important. It is probable, too, that the exten- 
sive explorations now carried on in prospecting for oil will result 
in the development of another important industry. 


FREIGHTS AND TRANSPORTATION. 


At the present time both freight and passenger rates to and be- 
tween Alaskan ports may be considered moderate. The great bulk 
of the freight traffic is northward, a condition that is not unlikely 
to continue. Any permanent increase in the traffic to and from 
Alaskan ports will naturally be accompanied by a corresponding 
lowering of rates. The transportation companies doing business im 
south Alaska seem to be quite as liberal as conditions will permit, 
and so far as expressed sentiment goes their general policy will be 
the wise one of encouraging as far as possible any industry that 
promises to add to the sum total of the traffic. 


DESIRABILITY OF SOUTH ALASKA AS A HOME. 


Climate—The south Alaska coast les in the same latitude as 
northern Labrador, the north of Scotland, and the south of Sweden, 
but none of these regions is very similar to it. In fact, south Alaska 
has several peculiarities which render close comparison with any 
other region difficult. In general, the climate is a moist one, accom- 
panied by no great extremes in temperature. The thermometer very 
seldom reaches zero in winter, nor does it exceed 75° F. in summer. 

The following tables give the more important meteorological data 
as compiled from various published reports, localities in Sweden, 
Canada, and the State of Washington being included for comparison : 


Monthly and annual mean temperatures at points in Alaska and eslewhere. 


Station. Jan. »Feb. | Mar.| Apr. | May.|June. July, Aug.|Sept.} Oct. | Nov.| Dec. ey 
|) SER seen erie [or ete Woe Ut eal ake | © P| oF. | 89, (eae eee 
SN Or a i choy 31.4 | 32.9 | 85.6 | 40.8 | 47.0 | 52.4 | 55.4 | 55.9 | 51.5 | 44.9 | 88.1/ 83.3] 43.3 
Dithkeo 2.2... 34.2 | 33.0 | 37.2 | 41.9 | 46.9 | 51.6 | 54.4 | 56.6 | 52.3 | 45.7 | 39.8 | 36.0) 44.5 
Kadiak_. .....-| 30.0 | 28.2 | 82.6 | 36.3 | 43.2 | 49.5 | 54.7 55.2 | 50.0 | 42.3 | 34.7 | 80.5] 40.6 
Unalaska@._.......) 30.0 | 31.9 | 30.4 | 35.6 | 40.9 | 46.3 | 50.6 | 51.9 | 45.5 | 37.6 | 33.6 | 30.1) 38.7 
Unalaska?____. 33.5 | 80.5 | 32.6 | 85.2 | 40.4 | 45.9 | 49.6 | 50.3 | 46.0 | 40.4 | 34.6 | 82.8 39. ¢ 
Port Angeles, | 

Wash ............| 34.7 | 36.7 | 41.7 | 45.6 | 50.6 | 54.0 | 56.6 | 56.8 | 52 7 | 47.7 | 42.4 | 88.2] 46.1 
Ottawa, Canada 11.9 | 12.2 | 17.6 | 41.5 | 63.6 | 66.9 | 70.4 | 68.7 | 57.7 | 48.1 | 84.5 | 17.8] 42.1 

Stockholm, Swe- | | | 
den. ..........-...| 33.5 | 29.5 | 33.8 | 39.5 | 52.5 | 57.0 | 59.1 | 59.3 | 53.6 | 40.6 | 35.6 | 27.3) 43.4 

| / | | ; 


aFrom records kept By the Russian Government. e 
>From records of the United States Signal Service 


aos 


DESIRABILITY OF SOUTH ALASKA AS A HOME. 27 


Average precipitation at points in Alaska. 


| “iy Bs 
eS | @ ee ae oes 
Station. par pos 7 sg Ds Zeek age 
I 5 | 4 — | . ; z A 2 a A ee Pee 
ted ee ye Pe | ae ee fe £ 2 2 £ |\Se0 
ajo Se ee Sl os les Ss oe 2 2 B) 5S Ss 
eH |e | & / j/AlH}R] 4] a OMe A Se 8 
: In. | In. | In. | In. | In. | In. | In. | In. | In. | In. | In. | In. | In. | In. 
Te eee | 7.95) 8.02] 7.78) 5.03) 3.89 3.87) 4.14) 6.67 10.94) 12.96) 10.77) 8.52 90.54 29.51 
Mendinke | 6.56) 3.70) 4.86) 4.01) 5.92 4.91) 3.38) 4.97| 7.26) 8.09 6.56 7.94 68.16 26.44 
UnalasKa __--__----- 13. 81| 7.68) 6.48) 7.51] 4.49 4.26) 2.78, 3.40 8.64) 11.98 9.30 11.81 92.14 23.57 


In comparing the data for Sitka, Kadiak, and Unalaska it will be 
noted that the average mean temperature of Sitka is a little higher 
than that of Kadiak, which in turn is higher than that of Unalaska. 
It will also be noted that Sitka and Unalaska have about the same 
rainfall—considerably greater than that of Kadiak. 

A matter of more practical consequence than either the copious 
rainfall or the comparatively high mean temperature is the rather 
low total of effective temperatures during the months from May to 
September, inclusive. By effective temperature is meant that above 
43° F., at which plant growth practically begins. These totals, as 
compiled by Evans,? are as follows: 


“SO Lea gS airs sae ter eee oes eee 1, 479. 1 
LSC TS Sa ia a ale es Se Sa Set es ees i 152-4 
ple k OS) 2p ee Te oa i ot es ee ee 624. 5 
erie A ieoles:. Waly Soi!) 3 oi vey ores la foe eet ee 1, 671. 0 
UTE TE CETTE ih 2 Se Se ee ee 5, 424. 7 
SYED US LT ee a eee es See ES ae eee ee ae oy ee f, 69227 
Saas eRIOTE ee Sy OCOD sss Shee a id se ee ZOOL D 


The difference in totals between Sitka and Kadiak is very striking, 
but not so much as that between Kadiak and Unalaska. Undoubt- 
edly this effective temperature factor is the principal cause of the 
sharp demarcation between the timbered and the timberless regions 
on the Alaska coast. 

Garden products.—This same factor—the low total of effective tem- 
peratures—limits also the variety of garden products that can be 
grown, but along the whole coast a considerable variety of vegeta- 
bles is successfully raised, such as potatoes, turnips, cabbage, cauli- 
flower, Brussels sprouts, kale, lettuce, peas, radishes, and rhubarb. 
Red currants and red raspberries grow wild on Cook Inlet, and these 
hardy varieties will thrive at most places along the coast—at Sitka 
even the ordinary garden varieties ripening. In southeastern Alaska 
salmon berries, cranberries, and huckleberries grow wild in abun- 
dance. 


4 Bulletin No. 48, Office of Experiment Stations, U. S. Department of Agri- 
culture. 


28 GRASS LANDS OF THE SOUTH ALASKA COAST. 


Fuel.—In the timbered region a supply of fuel is easily obtainable, 
while in the timberless country a rather scant quantity is secured 
from scrubby willows and alders and from beach drift. Coal of an 
inferior quality, but still fairly satisfactory for domestic use, is 
abundant along Cook Inlet. At present none of this is mined, but 
considerable quantities are gathered from exposed ledges, or from 
drift on the beaches. Most of the coal used along the Alaska Penin- 
sula, however, is at present shipped from Puget Sound. In some 
localities the paraffin residue from oil seepage is utilized as fuel. 


CHOICE OF A LOCATION. 


In general, Kadiak and the neighobring islands and the Cook Inlet 
country are the most favorable places for live-stock raising on 
account of a great luxuriance of grasses and contiguity to timber. 
The Cook Inlet region enjoys the reputation of being the garden spot 
of the Alaska coast, apparently producing finer vegetables than else- 
where, though lying farther north than the Alaska Peninsula and 
most of the territory described in these pages. The accessible grass 
lands here are, however, comparatively limited. 

On the other hand, Unalaska and the neighboring islands, while 
possessing less abundant grass and perhaps a less favorable climate, 
can perhaps reach markets in the Nome region and on the lower 
Yukon. At Yakutat, while the grass is not overabundant, the loca- 
tion is more favorable for shipments southward. 

The prime requisite of any Alaska location is a sufficiently large 
available supply of winter forage. Of summer range there is an 
abundance nearly everywhere, but the utilization of this is definitely 
limited by the number of cattle one can safely winter. The all- 
important point is therefore to have a sufficient acreage of land from 
which hay or silage can be secured. By selecting locations on the 
flat lands that so commonly occur at the heads of the narrow fiords 
one can easily control for all practical purposes great areas of grazing 
lands. 

The writer can not refrain from quoting here the following opinion 
of a widely traveled man from California, who for three years has 
been engaged in placer mining on the beach on the west side of Kadiak 
Island and who is seriously considering taking up a homestead and 
bringing his family to Alaska : 

In all my travels I have never found a place where one can live so well or 
so cheaply as I have done for the past three years. I can raise all sorts of 
hardy vegetables and berries, besides the wild ones, and have unlimited grass 
to keep cattle and sheep. Fish of the choicest sorts—salmon, halibut, cod, and 
many others—are very abundant, and the stream flowing by my cabin door 
swarms with trout. In the way of big game there are bears, Of-small game 


LAND LAWS APPLYING TO ALASKA. 29 


ducks and geese are plentiful in the spring and fall, and fresh gull eggs may be 
had for the gathering. To add to all this, if ready money is not available, I 
ean always make good wages at least by washing out gold on the beach. 


Surely there is here a combination of resources that makes failure 
well-nigh impossible. 


LAND LAWS APPLYING TO ALASKA. 


The following report regarding the methods by which title may be 
securea to agricultural lands in Alaska was prepared. in the office 
of the Commissioner of the General Land Office, through the cour- 
tesy of the Secretary of the Interior. It refers solely to acquiring 
title to agricultural lands and not to the town-site or mineral laws, or 
to mission claims under section 27 of the act of June 6, 1900 (31 Stat. 
L., 330) : 


Section 1 of the act of Congress approved May 14, 1898 (30 Stat. L., 409), 
extending the homestead laws to Alaska, may be summarized as follows: 

First. Extending the homestead laws and the rights incident thereto to the 
district of Alaska. 

Second. Extending to such district the right to enter surveyed lands under 
provisions of law relating to the acquisition of title through soldiers’ addi- 
tional homestead rights. 

Third. Granting the right to enter unsurveyed lands in said district under 
provisions of law relating to the acquisition of title through soldiers’ additional 
homestead rights. ; 

Fourth. Prohibiting the location in said district of any indemnity, deficiency, 
or lieu lands pertaining to any land grant whatsoever orginating outside of said 
district. 

Fifth. Limiting each entry under this section to 80 rods along the shore of 
any navigable water, and reserving along such shore a space at least 80 rods 
between all such claims, and prohibiting the entry or disposal of the shore 
(meaning land lying between high and low water mark) of any navigable waters 
within said district. 

Sirth. Limiting each homestead in said district, whether soldiers’ additional 
or otherwise, to 80 acres in extent. 

‘This section was amended by the act of March 38, 1903 Soe Stat. L., 1028), the 
provisions of which may be stated as follows: 

The amendatory act does not specifically reenact that portion of the act of 
1898 which granted the right to enter wisurveyed lands in the district of Alaska 
under the provisions of law relating to the acquisition of title through soldiers’ 
additional rights, but it is provided thereby “that no more than one hundred 
and sixty acres shall be entered in any single body by such scrip, lieu selection, 
or soldiers’ additional homestead right,’ which seems to negative any intention 
to modify or repeal the existing law with regard to the exercise of such rights 
in the district of Alaska further than to limit the amount which may be entered 
in a single body to 160 acres. Further, that portion of the amendatory act 
which provides that “ no indemnity, deficiency, or lieu-land selections pertaining 
to any land grant outside of the district of Alaska shall be made, and no land 
serip or land warrant of any kind whatsoever shall be located within or exer- 
cised upon any lands in said district, except as now provided by law,” seems to 


30 GRASS LANDS OF THE SOUTH ALASKA COAST. 


recognize that there are such outstanding rights; but, unless so'diers’ additional | 
homestead rights are thereby considered as scrip rights, this Department is not 
advised as to any other law permitting the exercise of any such rights in the 
district of Alaska. Soldiers’ additional homestead applications, under sections 
2306 and 2307, Revised Statutes, are received as heretofore, but not more than 
160 acres can be taken in a single body. 

The act of 1898 is amended so as to increase the amount of land which may 
be entered as a homestead in the district of Alaska to 320 acres, and in providing 
therefor grants such rights to “any person who is qualified under existing laws 
to make homestead entry of the public lands of the United States who has 
settled upon, or who shall hereafter settle upon, any of the public lands of the 
United States situated in the district of Alaska, whether surveyed or unsur- 
yeyed.” If a person be qualified, therefore, to make homestead entry under 
existing laws, he may enter not to exceed 320 acres, upon which he may have 
settled, in the district of Alaska, and without regard to the amount he might be 
authorized to make homestead entry of elsewhere; but the right to locate a 
soldier’s additional homestead right in the district of Alaska, without settle- 
ment, is not thereby changed. Only 160 acres or less may be commuted. 

No entry of any kind in the district of Alaska can, however, be allowed for 
land extending more than 160 rods along the shore of any navigable water, 
which is twice the extent originally permitted by the act of 1898, and along 
such shore a space of at least 80 rods is reserved between all claims, being the 
same as originally provided in the act of 1898. 


HOMESTEADS. 


The homestead laws secure to qualified persons the right to settle upon, 
enter, and acquire title to not exceeding 320 acres of public land, by establishing 
and maintaining residence thereon and improving and cultivating the land for 
the continuous period of five years. 

A homestead entryman must be the head of a family or a person who has 
arrived at the age of 21 years, and a citizen of the United States, or one who 
has filed his declaration of intention to become such, as required by the nat- 
uralization laws, to which section 5 of the act of March 3, 1891 (26 Stat. L., 
1095), attaches the conditions that he must not be the proprietor of more than 
160 acres of land in any State or Territory, and that since August 30, 1890, 
he has not acquired title to, nor is now claiming under any of the agricultural 
public-land laws, an amount of land which, together with the land now applied 
for, will exceed in the aggregate 320 acres. 

Where a wife has been divorced from her husband or deserted, so that she 
is dependent upon her own resources for support, she can make homestead 
entry as the head of a family or as a femme sole. 

Where an unmarried woman settles upon a tract of public land, improves 
the same, establishes and maintains a bona fide residence thereon with the 
intention of appropriating the same for a home under the homestead law, and 
thereafter marries before making entry of said land, or before making appli- 
cation to enter said land, she does not, on account of her marriage, forfeit her 
right to make entry and receive patent for the land: Provided, That she does 
not abandon her residence on said land and is otherwise qualified to make 
homestead entry: And provided further, That the man whom she marries is 
not, at the time of their marriage, claiming a separate tract of land under the 
homestead law. (Act June 6, 1900, 31 Stat. L., 683,) 


_---- 


LAND LAWS APPLYING TO ALASKA. OL 


APPLICATION FOR A HOMESTEAD FOR SURVEYED LAND. 


To obtain a homestead the party should select and personally examine the 
land and be satisfied of its character and true description. 

He must file an application, stating his true name, residence, and post-office 
nddress, and describing the land he desires to enter, and make affidavit that he 
is not the proprietor of more than 160 acres of land in any State or Territory ; 
that he is a citizen of the United States, or that he has filed his declaration of 
intention to become such, and that he is the head of a family, or over 21 years 
of age, as the case may be; that his application is honestly and in good faith 
made for the purpose of actual settlement and cultivation, and not for the bene- 
fit of any other person, persons, or corporation, and that he will faithfully and 
honestly endeavor to comply with all the requirements of law as to settlement, 
residence, and cultivation necessary to acquire title to the land applied for; 
that_he is not acting as agent of any pevson, corporation, or syndicate in making 
such entry, nor in collusion with any person, corporation, or syndicate to give 
them the benefit of the land entered, or any part thereof, or the timber thereon; 
that he does not apply to enter the same for the purpose of speculation, but in 
good faith to obtain a home for himself, and that he has not, directly or in- 
directly, made, and will not make, any agreement or contract in any manner 
with any person or persons, corporation, or syndicate whatsoever, by which 
the title which he might acquire from the Government of the United States 
should inure, in whole or in part, to the benefit of any person except himself ; 
and, further, that since August 30, 1890, he has not acquired title to nor is he 
claiming under any of the agricultural public-land laws an amount of land 
which, together with the land he is seeking to enter, will exceed in the aggre- 
gate 320 acres, and that he has not theretofore had the benefit of the home- 
stead laws, and must pay the legal fee and that part of the commissions which 
is payable when entry is made, and furnish the usual nonmineral affidavit. 

On compliance by the party with the foregoing requirements the receiver will 
issue his receipt for the fee and that part of the commissions paid, a duplicate 
of which he will deliver to the party. The matter will then be entered in the 
records of the district office and reported to the General Land Office. 

The applicant must in every case state in his application his place of actual 
residence and his post-office address, in order that notices of proceedings rela- 
tive to his entry may be sent him. The register and receiver will note the post- 
office address on their tract books. 


INCEPTIVE RIGHTS OF HOMESTEAD SETTLERS. 


An inceptive right is vested in the settler by the proceedings hereinbefore 
described. He must, within six months after making his entry, establish his 
actual residence in a house upon the land, and must reside upon and cultivate 
the land continuously in accordance with law for the term of five years. Occa- 
sional visits to the land once in six months or oftener do not constitute resi- 
dence. The homestead party must actually inbabit the land and make it the 
home of himself and family, as well as improve and cultivate it. 

At the expiration of five years, or within two years thereafter, he may make 
proof of his compliance with law by residence, improvement, and cultivation for 
the full period required, and must show that the land has not been alienated 
except as provided in section 2288, Revised Statutes (sec. 2291, Rey. Stat.), as 
amended by section 3 of the act of March 3, 1891 (26 Stat, L., 1095), 


32 GRASS LANDS OF THE SOUTH ALASKA COAST. 


The period of continuous residence and cultivation begins to run at the date 
of actual settlement in case the entry at the district land office is made within 
the prescribed period (three months) thereafter or before the intervention of a 
valid adverse claim. If the settlement is on unsurveyed land, the latter period 
runs from the filing of plat in the district land office. (Act May 14, 1880, 21 
Stat. L., 140.) 


HOMESTEAD SETTLERS ON UNSURVEYED LANDS. 


A homestead settler on unsurveyed public land not yet open to entry must 
make entry within three months after the filing of the township plat of survey 
in the district land office. (Act May 14, 1880, 21 Stat. L., 140.) 


CULTIVATION IN GRAZING DISTRICTS. 


In grazing districts stock raising and dairy production are so nearly akin to 
agricultural pursuits as to justify the issue of patent upon proof of permanent 
settlement and the use of the land for such purposes. 

Proofs can only be made by the homestead claimant in person, and can not be 
made by an agent, attorney, assignee, or other person, except that in case of the 
death of the entryman proof can be made by the statutory successor to the nae 
stead right in the manner provided by law. 

Sections 2291 and 2292, Revised Statutes, provide for obtaining title to lands 
entered by a homestead settler by his heirs. The act of June 8, 1880 (21 Stat. 
L., 166), provides for homestead claimants who become insane. 


HOMESTEAD CLAIMS NOT LIABLE FOR DEBT AND NOT SALABLE. 


No lands acquired under the provisions of the homestead laws are liable for 
the satisfaction of any debt contracted prior to the issue of the patent. (Sec. 
2296, Rev. Stat.) 

The sale of a homestead claim by the settler to another party before becoming 
entitled to a patent vests no title or equities in the purchaser as against the 
United States. In making final proof the settler is by law required to swear that 
no part of the land has been alienated except for church, cemetery, or school 
purposes or the right of way for railroads, canals, or ditches for irrigation or 
drainage across it. (Sec. 2288, Rev. Stat., as amended by sec. 3 of the act of 
March 3, 1891, 26 Stat. L., 1095.) 


SOLDIERS AND SAILORS’ HOMESTEAD RIGHTS. 


Any officer, soldier, seaman, or marine who served for not less than ninety 
days in the Army or Navy of the United States during the rebellion, and who 
was honorably discharged and has remained loyal to the Government, and who 
makes a homestead entry of 520 acres or less on any land subject to such entry, 
is entitled under section 2305 of the Revised Statutes to have the term of his 
service in the Army or Navy, not exceeding four years, deducted from the period 
of five years’ residence required under the homestead laws. 

If the party was discharged from service on account of wounds or disabilities 
incurred in the line of duty the whole term of enlistment, not exceeding four 
years, is to be deducted from the homestead period of five years; but no patent 
can issue to any homestead settler who has not resided upon, improved, and cul- 
tivated his homestead for a period of at least one year after he commenced his 
improvements. (Sec. 2805, Rev. Stat.) 


LAND LAWS APPLYING TO ALASKA. 33 


Similar provisions are made in the acts of June 16, 1898 (30 Stat. L., 473), and 
March 1, 1901 (81 Stat. L., 847), for the benefit of like persons who served in 
the late war with Spain or during the suppression of the insurrection in the 
Philippines. 

A party applying to make entry under the provisions of section 2304 must file 
with the register and receiver a certified copy of his certificate of discharge, 
showing when he enlisted and when he was discharged; or the affidavit of two 
respectable, disinterested witnesses corroborative of the allegations contained in 
the prescribed affidavit (Form 4—065) on these points, or, if neither can be pro- 
cured, his own affidavit to that effect. 

The widow or, in case of ker death or remarriage, the guardian of minor chil- 
dren may complete a filing made by the soldier or sailor as above, and patent 
will issue accordingly. 


SOLDIERS’ ADDITIONAL HOMESTEAD ENTRY. 


Any officer, soldier, sailor, or marine who served for not less than ninety days 
in the Army or Navy of the United States during said wars, who had, prior to 
June 22, 1874, the date of the approval of the Revised Statutes, made a home- 
stead entry of less than 160 acres, may enter an additional quantity of land, 
adjacent to his former entry or elsewhere, sufficient to make, with the previous 
entry, 160 acres. (Rey. Stat., 2306.) This right was extended by section 2307, 
Revised Statutes, to the widow, if unmarried; otherwise to the minor orphan 
children by proper guardian. If there be no widow, unmarried, and no minor 
orphan children, the right is held to be an asset of the soldier-entryman’s estate, 
to be disposed of by his personal representative as other personal property. (29 
L. D., 510 and 658.) An assignment by the heirs will be accepted if accompanied 
by a certificate of the proper court showing that no administration has ever been 
had on the soldier’s estate and that they are all the heirs entitled to the right. 
The right was formerly regarded as a personal one and not transferable, but 
under authority of the decision of the Supreme Court of the United States in the 
case of Webster v. Luther (163 U. 8., 331), it is now held to be assignable with- 
out restriction, and residence and cultivation are not required in its exercise, 
either by the original beneficiary or by his assignee, whether the original entry 
was perfected or abandoned (24 L. D., 502). 

It was formerly the practice, on proof of military service and original entry, 
under section 2306, Revised Statutes, to issue a certificate in the name of the 
soldier-entryman, showing his additional right and its area, but the practice 
was discontinued by circular of February 13, 1883 (1 L. D., 654), and it is 
held that there is no statutory authority for the same and that the soldier can 
obtain the right for himself and sell it to another without certification (23 L. D., 
152) 

By the act of March 3, 1893 (27 Stat. L., 593), provision is made that where 
soldiers’ additional homestead entries have been made or initiated upon a cer- 
tifieate of the Commissioner of the General Land Office of the right to make 
such entry, and the certificate of right is found to be erroneous or invalid for 
any cause, the party in interest thereunder on making proof of his purchase 
may, if there is no adverse claimant, perfect his title by payment of the Goy- 
ernment price for the land, but no person may acquire more than 160 acres 
through the location of any such certificate. 

By the act of August 18, 1894 (28 Stat. L., 397), all certificates regularly 
issued are declared to be valid, notwithstanding any attempted sale or transfer, 
and holders thereof desiring to exercise a right of entry in their own names 


34 GRASS LANDS OF THE SOUTH ALASKA COAST. 


must file such certificates in the General Land Office, together with satisfactory 
proof of ownership and of bona fide purchase for value. If, upon examination, 
the proof so filed is satisfactory, an additional certificate will be attached to 
the original authorizing the location thereof, or entry of land therewith, in 
the name of the assignee or his assigns. (Circular of October 16, 1894; 19 L. D., 
302. ) 

Existing homestead laws, while recognizing settlement upon unsuryeyed pub- 
lic lands, do not authorize the entry or the patenting thereof until the public 
surveys have been regularly extended over them. This section as amended, 
however, in terms authorizes the entry of unsurveyed lands in Alaska, and makes 
provision for a private survey for the purpose of patenting the claim, if the 
public surveys have not been extended thereto at the time it is desired to sub- 
mit proof, as is hereinafter referred to. 

In executing surveys for homestead applications the instructions now pre- 
yailing will be followed, and the limit of 160 rods as to frontage will be meas- 
ured along the meandered line of said frontage. 

The form of the tract sought to be entered, if upon unsuryeyed land, is pre- 
scribed in the act as follows: 

If any of the land * * * is unsurveyed, then the land * * * must be 
in rectangular form, not more than a mile in length, and located upon the north 
and south lines run according to the true meridian. 

That is, the boundary lines of each entry must be run in cardinal directions, 
true north-and-south and east-and-west lines by reference to a true meridian (not 
inagnetic), with the exception of the meander lines on meanderable streams 
und navigable waters forming a part of the boundary lines of the entry. Thus 
a frontage meander line, and other meander lines which form part of the bound- 
ary of a Claim, will be run according to the directions in the Manual of Sur- 
veying Instructions issued by this Office, but other boundary lines will be run 
in true east-and-west and north-and-south directions, thus forming rectangles, 
except at intersections with meander lines. A 

In other respects the rules previously adopted to govern surveys of claims 
under the act of May 14, 1898, will continue to be followed, of course taking 
into consideration the limitations as to area of claims. 

Every person who is qualified under existing laws to make a homestead entry 
of the public lands of the United States who settles or has settled upon any of 
the unsurveyed public lands of the United States in the district of Alaska with 
the intention of taking the same under the homestead law shall, within ninety 
days from date of settlement or prior to the intervention of an adverse claim, 
file the record of his location for record in the recording district in which the 
land is situated, as provided by sections 18 to 16 of the act of June 6, 1900 (31 
Stat. L., 326 to 328). 

Said record shall contain the name of the settler, the date of settlement, and 
auch description of the land settled on, by reference to some natural object or 
permanent monument as will identify the same. 

If at the expiration of the time required under sections 2291 and 2292, Re- 
vised Statutes, and as modified by section 2305, Revised Statutes, or at such 
date as the settler desires to commute under section 2301, Revised Statutes, the 
public surveys have not been extended over the land located, the locator may 
secure a patent for the land located by procuring, at his own expense, a survey 
of the land, which must be made by a deputy surveyor who has been duly ap- 
pointed by the surveyor-general, in accordance with section 10 of the act of 
May 14, 1898 (30 Stat L., 409), and the provisions of the act of March 3, 1903, as 
herein set forth. 


LAND LAWS APPLYING TO ALASKA. 35 


When the survey, either public or private, as herein provided for is approved 
by the surveyor-general under authority of this Office, the same rules should be 
followed as heretofore established governing the location of soldiers’ additional 
homestead rights, in addition to which the settler must furnish the required 
proof of residence and cultivation. 

The office of the surveyor-general of Alaska is located at Sitka. 

Section 10 of said act of May 14, 1898, also provides that all affidavits, 
testimony, proofs, and other papers provided for by this act and by said act of 
March 3, 1891, or by any departmental or Executive regulation thereunder, by 
depositions or otherwise, under commission from the register and receiver of 
the land office, which may have been or may hereafter be taken and sworn 
to anywhere in the United States, before any court, judge, or other officer 
authorized by law to administer an oath, shall be admitted in evidence as if 
taken before the register and receiver of the proper local land office. And 
thereafter such proof, together with a certified copy of the field notes and plat 
of the survey of the claim, shall be filed in the office of the surveyor-general 
of the district of Alaska, and if such survey and plat shall be approved by 
him, certified copies thereof, together with the claimant’s application, shall be 
filed in the United States land office in the land district in which the claim is 
situated, whereupon, at the expense of claimant, the register of such land 
office shall cause notice of such application to be published for at least sixty 
days in a newspaper of general circulation publshed nearest the claim within 
the district of Alaska, and the applicant shall at the time of filing such field 
notes, plat, and application to purchase in the land office aforesaid, cause a 
copy of such plat, together with the application to purchase, to be posted upon 
the claim, and such plat and application shall be kept posted in a conspicuous 
place on such claim continuously for at least sixty days, and during such period 
of posting and publication, or within thirty days thereafter, any person, cor- 
poration, or association having or asserting any adverse interest in, or claim to, 
the tract of land or any part thereof sought to be purchased, may file in the 
land office where such application is pending, under oath, an adverse claim 
setting forth the nature and extent thereof, and such adverse claimant shall, 
within sixty days after the filing of such adverse claim, begin action to quiet 
title in a court of competent jurisdiction within the district of Alaska, and 
thereafter no patent shall issue for such claim until the final adjudication of 
the rights of the parties, and such patent shall then be issued in conformity 
with the final decree of the court. 

When a settler desires to commute, the survey and homestead application 
must cover his entire claim, but only 160 acres, or less, thereof may be com- 
muted, in which event the entry will stand intact as to the portion not 
commuted, subject to future compliance with the requirements of law within 
the statutory period of seven years. 

Entrymen who commute will be required to pay, in addition to the price of 
$1.25 per acre, the same fees and commissions as in final homesteads. 

Whenever a settler or other claimant desires to make entry or submit final 
proof, he should address the register and receiver of the United States land 
office at Juneau, Alaska. 


eh tjaatels 


eabo'lis: 


oO 


DESCRIPTION OF PLATES. 


PLATE I. Map of Alaska, showing the approximate location of the grass-land 
areas in black. 

PLATE II. Fig. 1—View of the level lands at the head of Womans Bay, Kadiak 
Island. Similar areas occur at the heads of most of the inlets. Fig. 2.— 
Mowing beach rye on the Frye-Bruhn ranch. 

PLate III. Bluetop (Calamagrostis langsdorfii) on Kadiak Island, 6 feet high, 
July, 1904. The hillsides in the background were burned over during the 
preceding spring, and are covered with an equally luxuriant stand of the 
same grass. 

PuatTe LV. Fig. 1.—A view of Kadiak, November 7, 1903. <A light fall of new 
snow covers the low mountains in the background. Fig. 2.—Another view 
of Kadiak, March 26, 1904. The small snowfall of this region is made very 
clear by these two pictures. 


38 


Bul. 82, Bureau of Plant Industry, U. S, Dept. of Agriculture. PLATE Il. 


Fig. 1.—A VIEW OF THE FLAT LANDS LYING AT THE HEAD OF WOMAN’S Bay, KADIAK 
ISLAND, ALASKA. 


Fia. 2.—MOWING BEACH RYE ON KaDIAK ISLAND, ALASKA. 


> 
? 


PLATE III. 


® 
2 
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= 
3 
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42 
a 
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a 
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=) 
a 
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SI 
mo} 
= 
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s 
as 
oe 
v= 
° 
=) 
ws 
® 
2 
3 
jaa} 
aN 
oo 
Sl 
iva) 


BLUETOP (CALAMAGROSTIS LANGSDORFII) SIX FEET HIGH, ON KADIAK ISLAND, ALASKA, 


JuLy, 1904. 


The grass on the hillside in the background was just as luxuriant. 


w¥ 


peers eee 
Ritts 


—— lc tll 
. Qs r i 
= 
. ° i s 
. 
. t n 
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: i 
‘ . ‘ ' 
i 
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; . ~~ t 
2 ’ -¢ 
. “ : 
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be er nes 


Bul. 82. Bureau of Plant Industry, U. S. Dept. of Agriculture. PLATE IV, 


Fig. 1.—A VIEW OF KADIAK, ALASKA, NOVEMBER 7, 1903. 


Fig. 2.—A DIFFERENT VIEW OF KADIAK, MARCH 26, 1904. 


The small snowtall of this region is made very clear by these two pictures, 


Ee, DEPAKIMENT OF AGRICULTURE. 
BUREAU OF PLANT INDUSTRY—BULLETIN NO. 83. 


B. T. GALLOWAY, Chief of Burcau. 


VITALITY OF BURIED SHEDS. 


BY 


03. Ws ok. DU Vt, 


ASSISTANT IN THE SEED LABORATORY. 


Issuep Auaust 4, 1905. 


WASHINGTON: 
GOVERNMENT PRINTING OFFICE. 
EOD: 


BUREAU OF PLANT INDUSTRY. 


B.. T. GALLOWAY, 
Pathologist and Physiologist, and Chief of Bureau. 


VEGETABLE PATHOLOGICAL AND PHYSIOLOGICAL INVESTIGATIONS. 
Apert F. Woops, Pathologist and Physiologist in Charge, Acting Chief of 
Bureau in Absence of Chief. 


BOTANICAL INVESTIGATIONS AND EXPERIMENTS. 
Freperick V. CoviL_e, Botamist in Charge. 
GRASS AND FORAGE PLANT INVESTIGATIONS. 
W. J. SpitymMan, Agriculturist in Charge. 
POMOLOGICAL INVESTIGATIONS. 
G. B. Brackett, Pomologist in Charge. 
SEED AND PLANT INTRODUCTION AND DISTRIBUTION. 
A. J. Prerers, Botanist in Charge. 

ARLINGTON EXPERIMENTAL FARM. 

L. C. Corsert, Horticulturist in Charge. 


EXPERIMENTAL GARDENS AND GROUNDS. 
E. M. Byrnes, Superintendent. 


J. E. Rockwe.u, Editor. 
James E. Jones, Chief Clerk. 


SEED LABORATORY. 
ScreENTIFIC STAFF. 
EpGar Brown, Botanist in Charge. 


F. H. HituMan, Assistant Botanist. J. W. T. Duvet, Assistant. 


» 


LETTER OF TRANSMITTAL. 


U. S. DEPARTMENT OF AGRICULTURE, 
BurEAU OF PLant INDUSTRY, 
OFFICE OF THE CHIEF, 
Washington, D. C., May 29, 1905. ' 

Sir: I have the honor to transmit herewith, and to recommend for 
publication as Bulletin No. 83 of the series of this Bureau, the accom- 
panying technical paper entitled ‘‘ The Vitality of Buried Seeds.” 

The experiments discussed were undertaken in order to determine 
the length of time that seeds of different species of plants will retain 
their vitality when buried at various depths. Seeds of both cultivated 
and wild plants were used, but special attention was given to weed 
seeds in order to ascertain what weeds can be eradicated by deep plow- 
ing and how long the soil must remain undisturbed before the vitality 
of the seeds will be entirely destroyed. The results of the first year’s 
experiments show that the noxious character of weeds is closely 
associated with the length of time the seeds will remain viable in the 
soil, and that many weeds can be eradicated by plowing. Much 
additional information is given, showing the relative resistance of the 
seeds of cultivated plants and of those commonly designated as weeds, 
and the influence upon the preservation of vitality of the depth of 
burial, of hard seed coats, and of hulled as compared with unhulled 
seed. 

This paper was prepared by J. W. T. Duvel, Assistant in the Seed 
Laboratory, and has been submitted with a view to publication. 

The accompanying illustrations are necessary for a complete under- 
standing of the paper. 

Respectfully, 
B. T. GaLLoway, 
Chief of Bureau. 
Hon. JAMES WILson, 
Secretary of Agriculture. 


oo 


el 
e 
« 


s 


. 3 ~o) 4 * 
ee had a 


COMLEN TS. 


cement GO! PiTial £0 Vijality . 7-2-2 ee een enon s--- ane 

EIR es ns Pa se ee oe, ake ene eee en owe eee 

Pimmoeiunavaten versus wild plants...--...2...c..-22 -<c.2------.---sccce 

ee al EES SS ee eRe gg Se 

Description of plates... -- ree seach 5 Semen Sete he Doe wc atee eae soca 
5 


Page. 


LLE GS TR A TAO ae 


PLATES. 


Puate I. Fig. 1.—Bromus racemosus, smooth brome-grass. Fig. 2.—Bromus 


secalinus, cheat, or chess ...2--....-..2+-5> eee 


If. Fig. 1.—Alsine media, common chickweed. Fig. 2.—Rumev crispus, 


curled dock. Fig. 3.—Datura tatula, jimson weed........-- wean 


Ill. Fig. 1.—Elymus canadensis, nodding wild rye. Fig. 2.—Fraxinus 


americana, White ash. Fig. 3.—Phytolacca americana, poke.....- 


TEXT FIGURE. 


Fic. 1. Diagram showing order in which seeds were buried..... scours 


6 


Page. 


bo 
bo 


to 
lo 


22 


10 


B. P. I.—165. 


fee VITALITY OF BURIED SEEDS. 


INTRODUCTION. 


The preservation of the vitality of seeds when buried in the soil 
and the awakening of metabolic activity in such seeds on being 
exposed to conditions favorable to their germination are equally as 
important to the practical farmer as to the scientist. The intelligent 
farmer in order to combat noxious plants successfully should know 
how much time must elapse after heavy crops of weeds of various 
sorts are turned under before the ground can be plowed again with 
safety. He should also know what plants he can hope to eradicate in 
this way, for with many of our worst weeds this method would result 
only in failure. In fact, the reason why the majority of our most 
persistent weeds are so difficult to eradicate is because their seeds are 
capable of retaining their vitality for a number of years when buried 
in the soil. It thus becomes important to know how different species 
of seeds behave when buried under similar conditions, and how seeds 
of the same species behave when buried under different conditions. 


KINDS OF SEEDS BURIED. 


So much has already been written on the germination of seeds that 
have remained dormant in the soil for a number of years, in some 
cases even for centuries, that it seemed desirable to determine with 
some degree of accuracy the length of time that certain seeds will 
retain their vitality when buried in the soil under known conditions. 
Accordingly, in the autumn of 1902, 112 different samples of seeds 
were selected for these experiments, as follows: 


TaBLeE I.—List of seeds selected for the experiments. 


Labora- Burial 
tory number 
; = S 1 : 
test Kind of seed. ant a as given 
num- ; on 
ber. diagram. 


Poacee (grass family): 


16173 Agropyron repens (L.) Beauv. (couch grass) ...-......-..-.---------- 100 | 31 
16174 CE m OCU CELI (WALIGCOMb) lo. aasiee cesta aes Soeeteee Seine oo- Gace bees 71 |, 9 
16175 PARCTI SLIDE COOLS eects nine tae teenies cine Sina as eee one Se Siena 24 8 
16176 ETONLESSECOIUVUS Nas (Cheat. CHESS) 2. coe Soe senlecemeseae oes soem aso 34 36 
16177 Bromus racemosus L. (upright chess, smooth brome-grass).........-. 33 37 
16178 Chaetochloa verticiliata (L.) Scribn. (foxtail) ...........--.----------- 108 66 
16179 Chaetochloa glauca (L.) Seribn. (yellow foxtail)...............--....- 46 33 
16180 Chaetochloa viridis (L.) Seribn. (green foxtail)...........-..-..-....- 5 67 
16181 Eleusine indica (L.) Gaertn. (wire-grass, crab-grass) ..............-.- 36 72 
16182 EVV INUS ULF OURICUS 1a: (IV ITSINIA WIIG, TYC) acm sas cence e coon enc ne ee 77 15 
16183 biymis canadensis. o. (NOUdIng Wild. FYe)2. 2252-2... .25-- 25-0225. --- 74 13 
16184 Bhymistriticoides Buckl: (wild wheat): <. <- ce0ce<ecc a+ ccamecceneee se 69 | 14 


~I 


VITALITY OF BURIED SEEDS. 


TaB_e I.— List of seeds selected for the experiments—Continued. 


1 


Labora-| 


tory 
test 
num- 
ber. 


16189 
16190 


16191 
16192 | 
_ 16193 | 


16194 
16195 


16196 | 


16197 | 


16198 | 


16199 
16200 
16201 
16202 


16203 
16204 


16205 | 


16206 
16207 


16208 | 


16209 


16210 | 


16211 
16212 
16213 


16214 
16215 
16216 


16217 
16218 
16219 


16220 
16221 
16222 
16223 
16224 
16225 
16226 
16227 


16228 
16229 


16230 
16231 
16232 
16233 
16234 
16235 
16236 
16237 
16238 
16239 
16240 


16241 
16242 
16243 


16244 
16245 


Kind of seed. 


Poacez (grass family )—Continued. 

Festuca elatior L. (meadowitestue) cme. os ow.ce oe wav cm ioslesw sos ccmawane 

Hordeum sativum Jessen. (Bavi@y oe. 2= -. ooc 2 5-o- ow ne 2 2 om pe eee ee 

Panicum virgatum L. (tall, smooth panicum).......-----.----------- 

Phalaris arundinacea L. (reed Canary grass)..........----.---+------ 

‘Phieum pratense i. (tinmtothwi)teeeerer secee cae eens oe onan eras 

Poa pratensis. L, (Kentucky pluegrass) 2225. <5... ness cone oem eee ee 

Secale cereale Ii. (TYE) sc ac cites a cicleee steele ee oon eile ese de oe eee sees 

Sporobolus airoides Torr. (hair-grass Grop-seed) ......-.-------------- 

Sporobolus cryptandrus (Torr.) A. Gray (sand drop-seed)....-...--.-- 

Sporobolus cryptandrus(Torr.) A. Gray (sand drop-seed—hulled seed) . 

Triticum aestivum L. (wheat).....-+---- B olerelt o tcleat bas Gee eee ee 

Zea mays L. (corn—Boone County white)..............-------------- 

Zea mays L. (sweet corn—early Concord 
Cyperacez (sedge family): 

Cyperus esculentus L. (yellow nut-grass) .-.--. 22.20 00co es eeenseseman 
Liliacez (lily family): 

Alliwme cepa ls, (OWLOM) i siche cate aimless erie wets a ea te 
Conyallariacez (lily-of-the-valley family): 

Asparagus officinalis L. (asparagus) ...---..--.---.------------------- 
Moracez (mulberry family) : 

Cannabis sativa Ii. (hemp) fe 20 ste aajce opera ialelee oes ie la ae 
Urticacez (nettle family): 

Boehmeria nivea Gaud.Aramie) Ss. 22 de eeeieieoes sc oe eee ee seers 
Polygonacee (buckwheat family): 

Fagopyrum fagopyrum (L.) Karst. (buckwheat)...........----------- 

Polygonum pennsylvanicum L. (Pennsylvania persicaria, smartweed) . 

Polygonum persicaria L. (lady’s-thumb, smartweed)............--... 

Polygonum scandens L. (climbing false buckwheat) ...---..-..------ 

Rumex salicifolius Weinm. (willow-leaved dock) .........---.------- 

Rumex crispus L. (curled dock), not cleaned.........--..-.---------- 

Rumex obtusifolius L. (broad-leayed dock, bitter dock) ....-........- 
Chenopodiacez (goosefoot family): 

Axyris amaranthoides L. (Russian pigweed)........------------ ----- 

Beta vulgaris a. (Sugaribeet), 2.c-ee-22-2=<2-5-sna eee as cee 

Chenopodium album L. (lamb’s quarters, white goosefoot) .....-..... 

Chenopodium hybridum L. (maple-leayed goosefoot) .....-.---------- 
Amaranthaces (amaranth family): 

Amaranthus retroflecus (rough pigwee@) ......-..-..20-cssscecnesseace 
Phytolaceaceze (pokeweed family): 

Phytolacca americana L. (poke, pigeon berry)...--.-.---------------- 
Portulacaces (purslane family): 

Portulaca oleracea L. (purslane, pussley)..........---------------- cord 
Silenacee (pink family): 

Agrostemma githago L. (corn cockle) .. 2.0.5. 25-. 2042. seencj sian ensine 

Alsine media I.. (common chickweed) ..-.......2..0.. 2 ese ese sneeene 

Vaccaria vaccaria (1..) Britton (cowherb).........-.----+.-.----++--- 
Brassicacez (mustard family): 

Brassica nigra (L.) Koch (black mustard).............--.---+0---20- 

Brassica:oleracea Li;,(Cabbale)-.~ sens. .040ste sms aake see 

Brassica campestris 1s. (tUrnp) ci) boo cieeles een eee 

Bursa bursa-pastoris (.) Britton (shepherd’s purse) 

Erysimum cheiranthoides L. (wormseed, treacle mustard).........--- 

Neslia paniculata (L.) Desv. (ball mustard).............-....-.----+- 

Sisymbrium altissimum L. (tall sisymbrium).............+-----++--+--- 

Thiaspi arvense L. (field penny Cress) ......----- +2202 -- ee ee een ee ence 
Rosaceze (rose family): 

Potentilla monspeliensis L. (rough cinquefoil) .........-.--..---+----- 
Caesalpiniaceze (senna family ): 

Cassia marylandica L. (wild senna, American senna).......-..-----. 
Fabaces (pea family): : 

Lespedeza frutescens (L.) Britton (wand-like bush clover)............ 

Medicago sativa G. (alfalfa, lucern))..... 2.0.0.0. c02.sccescwe cena seuaene 

Phaseolus vulgarg@@e L. (DEAD) 5.625. .-cccccwceccedecgcases== enna eeenion 

Piswm sativum THC pea) 2. sais ceeds cw cwwebad sw comb cece= Seen aa eee 

Robinia pseudacacia L, (locust tree, false acacia) ............------4-- 

Trifolium hybridum L. (alsike clover) .......-...----2.-++0- seen nenee 

Trifolium pratense L. (Ted ClOVEL)..... 2. cee ece ce ccenen nnn cenensenes 

Trifolium pratense L. (red clover) harvest of 1900............-....---- 

Trifolium pratense L. (red clover) hard seed from No. 16237........-- 

Trifolium repens L. (white Clover) ......-----+-.... eee ences eens eeeee 

Vigna catjang Walp. (Lrom cowpea) ...........- +. see eee eee een eeeees 
Linacew (flax family): 

Linum usitatissimum L. (flax, lins@@d).......ce.sccc cece cncceesceceses 
Anacardiacese (Sumac family): 

Rhus glabra L. (scarlet SUMAC).... 2.2.20. c cence eens eee secescnenscsces 
Malvaces (mallow family): : 

Abutilon abutilon (L.) Rusby. (velvet leaf) ............csece ee cneeenes 

Gossypium hirsutum L, (COttON) .. 2.2... eee eee eee eee eee eee eens 

Hibiscus militaris Cay. (halberd-leaved rose mallow)............-.--. 


Sample 
number. 


Burial 
number 
as given 

on 


diagram. 


eas 


Je) 


HOW THE SEEDS WERE BURIED. 


TasLe I.— List of seeds selected for the experiments—Continued. 


Labora- : Burial 
tory . , | humber 
test Kind of seed. Sample | a5 given 

num- number. i 
ber. . diagram. 
Hypericacee (St. John’s wort family): 
16246 Ascyrum hypericoides L. (St. Andrew’s cross) .............-..--.------ 44 95 
Onagracee (evening primrose family): 
16247 Onagra biennis (L.) Scop. (common evening primrose) ...:...-....-- 8 96 
Apiacez (carrot family): 
16248 | PE ELUM ICI ae CCLCTY | 50-0 aclasiow cin = 22 ayn plates Ae naa cine a 94 | 57 
16249 Paginas sao is. (PpATSNIP, Wild) 2.522% 222.2 secs ere eee 95 56 
Oleacez (olive family): 
16250 Rraxinvus americand As. (white ash)<2- ..< 2. Vleet eee eee ee 105 21 
Convolvulacez (morning-glory family): 
16251 Convolvulus sepium L. (hedge bindweed, great bindweed)........... 56 | 23 
16252 |- Ipomoea lacunosa L. (small-flowered white morning-glory)........-- 81 Pe 
Cuscutacez (dodder family): : I 
16253 Cuscutu polygonorum Engelm. (smartweed dodder)..........-.------ 63 98 
16254 Cuscuta.eniunum Weihe. (flax dodder) .......-2. 22 -2.22-2-22 228-5 82 97 
Verbenacee (vervain family): 
16255 Werbenwnistara- - / Ide VELVaIN) 6-2. ~ 8 20 ss oe es Gees cc nessecte } 66 | 100 
16256 Verbena urticifolia L. (white vervain, nettle-leaved vervain)........ 79 | 99 
Solanacez (potato family): 
16257 Capsicum annuum L. (red pepper) ...-....-...---------- Saat See eat 39 59 
16258 Datura tatula L. (purple stramonium, jimson weed) .....-.......--- 106 61 
16259 Lycopersicon lycopersicon (L.) Karst. (tomato) 45 60 
16260 |! MaRGlranen LAnCHi Lis (LODRECO)) on etree nese ass ee oe ee eee eee 99 101 
16261 Solanum nigrum L. (black nightshade, garden nightshade) ........- 61 58 
Serophulariacee (figwort family): 
16262 Verbascum thapsus L. (great mullen)....- Bree ee De Ge oe aes Soe 76 102 
Plantaginacee (plantain family): 
16263 Plantago lanceolata L. (ribwort, ribgrass, buckhorn).........-.-.---- 88 105 
16264 | Plantago major L. (common plantain) ......-.-.------------------+-- / 91 103 
16265 | Plantago rugelii Dec. (Rugel’s plantain, broad plantain) ......-.-.--- 65 | 104 
| Cucurbitaceze (gourd family): 
16266 | Citrullus citrullus (L.) Karst. (watermelon).......--..2....-----------| 6 26 
16267 | OUEUIS INCL? bs. (TAUSKANCION))t a5 -- acs = scco-~ wwe} sem oe rcae eae sees 26 25 
16268 BUTTS COLTUILS Ls: (CHEERED) os vee oe ne nani ape nin we mnie Sos s lee ne oo 48 24 
Cichoriacez (chicory family): 
16269 Se aChuce: Sarrole Wot piaekd y-lGLbMCG) 2 eaca.0 2 e252 atten an. too t= choose 98 107 
16270 COGVT IRE aed Vig 2h 6 1 Cle) a a 8 ee 28 62 
16271 Taraxacum erythrospermum Andrz. (red-seeded dandelion).......... 90 106 
Ambrosiacez (ragweed family): | 
16272 Ambrosia artemisiaefolia L. (ragweed) ..-....---------------+-+-------- 87 63 
16273 Ambrosia trifida L. (great ragweed) .....-...--..---------------.----- 53 28 
16274 | Xanthium pennsylvanicum Wallr. (Pennsylvania clotbur, cocklebur). 51 27 
| Asteracez (aster family): | 
16275 | Arctium lappa L. (burdock, Clotbur)........-.--.---------2-2-++------ 101 112 
16276 | Bidens frondosa L. (black beggar ticks) ... 3 84 64 
16277 | Carduus arvensis (L.) Robs. (Canada thistle 80 111 
16278 | Chrysanthemum leucanthemum L. (whiteweed, oxeye daisy) 92 110 
16279 / Grindelia squarrosa (Pursh.) Dunal. (broad-leaved gum plant) ....-- 89 | 1u8 
16280 | Helianthus annuus L. (common sunflower, wild) ...-...-.-.-.---- fest 97 29 
16281 | Helianthus annuus L. (common sunflower, cultivated).......-.....-- 29 7 
16282 | Onopordon acanthium L. (cotton thistle, scotch thistle) ............--. 109 | 65 
16283 Rudbeckia hirta L:.( black-eyed Susan) .............------------------- 57 109 
| Pinacez (pine family): | 
30 


16284 | Pinus virginiana Mill. (scrub pine, Jersey pine)........-..-..-------- 36 


HOW THE SEEDS WERE BURIED. 


The foregoing list represents 109 species, 84 genera, and 34 families 
of plants. Carefully counted seeds of these samples were mixed with 
dry clay soil and packed in well-baked earthen pots (the common 
flowerpot used in greenhouses). The filled pots were covered with 
inverted clay saucers in order to prevent the seeds from being 
destroyed or becoming mixed with other seeds which might have been 
in the soil with which the pots were covered. By burying the seeds 
mixed with earth in porous clay pots of this character they were sub- 
jected to conditions almost identical with those which would exist if 
the seeds were buried either accidentally or by natural causes. The 
porous clay pots admitted of the free circulation of air and water. 

30356— No. 883—05——2 


10 VITALITY OF BURIED SEEDS. 


The pots containing these seeds were buried at three different depths. 
Eight complete sets were buried from 6 to 8 inches below the surface, 
being covered approximately the same as would result from deep 
plowing. Twelve complete sets were covered toa depth varying from 
18 to 22 inches, sufficiently deep in this latitude to be reasonably 
secure from the action of frost. Twelve more complete sets were 
buried at a depth varying from 36 to 42 inches where the conditions 
were nearly uniform, so far as the three factors which regulate the 
germination of seeds are concerned, namely, heat, moisture, and air 
(oxygen). Figure 1 shows the arrangement of the pots, which were 
of 6-inch, 4-inch, and 2-inch sizes, to accommodate the different kinds 
of seed. 


22 OOCCOOOOOOS 


DEEEEwNweon@e 
PEODS OOOO 
e320) 170 a) a ee 
QUOOSROOCORS 
Belair @enoore 

(106) (07108) /09 OOG 


Fic. 1.—Diagram showing order in which seeds were buried, 


These seeds were buried December 19 to 23, 1902, in a heayy clay 
soil on the Arlington Experimental Farm of the United States Depart- 
ment of Agriculture. With the exception of two of the duplicate 
samples of red clover, the seeds were of the harvest of 1902. 

In each case a definite number of seeds was taken. Of the larger 
kinds, such as beans, peas, corn, etc., 100 seeds were used, but for the 
majority of the samples 200 seeds were taken. The seeds selected 
were for the most part of plants with which the greater number of 
the farmers throughout the United States are more or less familiar, 
either as plants of economic importance or as weeds. 

In all 32 complete sets, representing 3,584 pots, were buried. One 
set from each of the three different depths is to be taken up as the 
conditions warrant and will be tested for vitality. The results of 
these tests are to be compared with the germination of seeds from the 
original bulk samples designated throughout this report as ‘* controls.” 


GERMINATION TESTS. 11 


The control samples are bejng stored in cloth bags in a dry room on 
the second floor of the Seed Laboratory. The first complete series of 
three sets was taken up in November, 1903, eleven months after they 
had been buried. The results of the first year’s experiment are given 
in the following pages. 


GERMINATION TESTS. 


In making the germination tests of the buried seeds the contents of 
the pots, the mixture of seed and soil, were spread on sand in ordinary 
greenhouse flats. Along with these, in the same flats, were control 
samples taken from the original bulk lot of seeds, as previously men- 
tioned. In addition, another complete set of control samples was 
tested in the germinating chambers of the Seed Laboratory. The tem- 
peratures given in the tables are those best suited for the germination 
of the different seeds. 

For convenience the results of the germination tests have been 
divided into three groups, as follows: 

A. Seeds in which the control samples, as well as those that were 
buried, gave only negative results when tested in the greenhouse. 

B. Seeds which had either decayed or germinated and afterwards 
decayed while they were buried. 

C. Seeds which had not completely lost their vitality while buried. 

The first group, i. e., those in which both the control samples and 
those which had been buried failed to germinate when planted in flats 
in the greenhouse, consists of the following species: 


. Axyris amaranthoides L. (Russian pigweed). 

. Boehmeria nivea Guad. (ramie). 

Bursa bursa-pastoris (L.) Britton (shepherd’s purse). 

. Cannabis sativa L. (hemp). 

. Chaetochloa viridis (L.) Seribn. (green foxtail), 

. Citrullus citrullus (L.) Karst. (watermelon). 

. Cyperus esculentus L. (yellow nut-grass). 

. Onagra biennis (L.) Scop. (evening primrose). 

. Polygonum pennsylvanicum L. (Pennsylvania smartweed, persicaria). 
10. Polygonum persicaria L. (lady’s-thumb, smartweed). 

11. Polygonum scandens L. (climbing false buckwheat). 

12. Sporobolus airoides Torr. (hair-grass drop-seed ). 

13. Sporobolus cryptandrus (Torr.) A. Gray (sand drop-seed—hulled seed). 


MIO OP wo DD 


Cc 3 


In this series the hemp should be discarded, as repeated tests failed 
to show any seeds from the bulk samples capable of germination. The 
control samples of the other seeds when tested in the germinating 
chambers germinated nearly as well and in some cases even better than 
the chamber tests which were made at the time the seeds were buried. 
Undoubtedly some of these seeds had decayed while buried in the soil; 
in fact, the watermelon seeds, Avyr/s amaranthoides, and Sporobolus 
airoides were marked ‘** mostly decayed” when taken up. Generally 
speaking, the results show that the failure to germinate was not in the 


12 VITALITY OF BURIED SEEDS. 


seeds, but that the conditions in the greenhouse were at fault, and 
until other tests are made these results can not be discussed with any 
degree of satisfaction. On the other hand, it is certain that some of 
the smaller seeds failed to germinate because they were covered too 
deeply when sown in the flats in the greenhouse. 

Polygonum scandens possibly should be classified in Table III, inas- 
much as some of the seeds which were buried at depths of from 18 to 
22 inches and from 36 to 42 inches showed a few sprouts at the time 
the seeds were taken up, but after being transferred to the greenhouse 
no seedlings were developed. However, the failure in the germina- 
tion of the control sample of Polygonum scandens throws it into the 
first group (A) with the other two species of the same genus, i. e., 
Polygonum pennsylvanicum and P. persicaria. 

The result of the tests of the buried seed of Sporobolus cryptandrus, 
as given in this group, should be compared with the germination of 
the unhulled seed as given in Table III, No. 64. The control sam- 
ples of both the hulled and the unhulled seed which were sown in the 
greenhouse failed to germinate, but all three samples of the unhulled 
seed that had been buried gave some germination when tested in the 
greenhouse. 


TABLE II.— Results of tests of seeds which had either decayed or germinated and afterwards 
decayed while buried. 


= Chamber tests. | Greenhouse tests in sand. 
= Labora- | 
Bs} LOL I} : | Depth of burial 
2s test Kind of seed. Tem- | Origi- | pane Con Pp E 
a~— num- | epra- <i) 
al ier, Resa anaes trol. | trol. | 68 | 1892 | 3649 
z | | ‘inches, inches. inches. 
—- = — 53 FPR ee | "| | | 
9) C. \- Perth. | Pereb / Per ct..| Per ct. | Per et.-| Per ct. 
14 16196  Zeamays(BooneCounty white ; | 
COLI sats ae wants eee 20-80 99.5 I er ae ees Be 
15 16197 | Zea mays (sweet corn) .......-| 20-30 98.5 Clo Pre ee | cee din sip alg hee metal ea eneea 
16 16217 | Agrostemma githago........-- r 20-30 | 99 |> 9855" |... Se .ccleemeeie seine ee oe 
17 16221 Brassica oleracea.............- 20 85.5 82 ||. .eael: oe ees 3] ee 
18 16244 | Gossypium hirsutum.......... | 20-80 | 77.5 7 Nan ERE pe aes) Cee 
19 16233 | Pisum sativum ................| 20-30} 99 98 | Sa secs Sp eee Sea oe ees 
20 | 16232 | Phaseolus vulgaris...........- 20-30 | 97.5 O80 lo vince alerts |e ae livin scat (a) 
21 16203 | Fagopyrum fagopyrum........ 20-30 100 98: 5) |fccxtuce (0)" 1 GG) Die aaeee 
22 16195 | Triticum aestivum ...........- ) 20, 99 96; Bibs caeen (Q) ye) Due eee (a) 
23 | 16186 | Hordeum sativum........+.... ) 20 100 98) 4 |ougeeees (dt). Po" @® (a) 
of} T6175") AvengBatlvas. mens sic tects | 20-80 | 70.5 91.5 | 95.5 0. 0 0 
25 16191, | Secale'cereale .....2.. 22-5050. 20 100 98.5] 88 0 | 0 0 
26 16267 | Cucumis melo....-.-<.s...--- 20-30 96.5 97 88 0 0 0 
97.|° 16199: | Allium cepai 0.3... ...dubuces: | 20-30} 94,251 88 | 70.5 0| 0 0 
28 | 16270 | Lactuca sativa ................ | 20-30 -100 98.5 91 0 0 0 
29 16281 | Helianthus annuus (cult.)....) 20-30 97 96.5 | 3 0) 0 0 
30 | 16241 | Linum usitatissimum ......... | 20-30) 93.5 | 95 $3.5 0° 0 0 
31 | 16245 | Hibiscus militaris............. 20-30 | 98.25] 92 94 | 0. 0 0 
32 16200 | Asparagus officinalis........... | 20-30 80 e69 74 F0 tO TO 
33 16177 | Bromus racemosus............. | 20-830 100 | 98.5 92.5 a0) 0 aod 
34 | 16176 | Bromus secalinuS.............. | 20-30 | 88.5 | 77 95.5 0 | 0 0 
35 | 16181 | Eleusineindica................ 35, |, .78.25.4.1./9)25)) "vor 0) 0 0 
36 16284 | Pinus virginiana@.............. 20-30 | 18 6.5 43.5 | 0} 0 0 
37 | 16234 | Robinia pseudacacin.......... 20-30 14 711.5 | 3.5 | 0 ) 0 0 


a Many had germinated and afterwards decayed. 

b Approximately 10 per cent had germinated; the remainder had decayed. 

e An occasional old sprout was found, 

d Approximately all had germinated and afterwards decayed. 

e Clipped, 87 per cent; not clipped, 51 per cent. 

f Practically all had sprouted; the sprouts from seeds buried at the 36-42-ineh depth were found 
matted in the bottom of the pot. 

g Clipped. 


GERMINATION TESTS. 13 


. 


The corn, sweet corn, corn cockle, cabbage, cotton, peas, beans, 
bfickwheat, wheat, and barley—the first ten samples given in the fore- 
going table—were all so unmistakably decayed when the seeds were 
taken up that the contents of the pots were thrown away, no green- 
house tests being made. The first six of these samples showed no 
trace of any remains of old sprouts; apparently all of the seeds had 
decayed before germination had taken place. If germination took 
place it must have been comparatively soon after burial, thus giving 
ample time for all of the old sprouts to decay beyond identification. 
This, -however, seems hardly probable, considering that the seeds 
were buried during the latter part of December, 1902; moreover, the 
beans, buckwheat, and barley from some or all of the different depths 
showed clearly the remains of well-developed radicles. 

The beans which were buried at depths of from 6 to 8 and from 18 to 
’ 22 inches had decayed, while many of those buried at a depth of from 
36 to 42 inches had germinated and afterwards decayed. The buck- 
wheat from the 6 to 8 inch depth showed that approximately 10 per 
cent had germinated, while at 18 to 22 inches there were only the 
remains of an occasional old sprout, and at 36 to 42 inches all of the 
seed had decayed. In the wheat the greater number of the grains 
that were buried from 6 to 8 and from 36 to 42 inches had germinated 
and then decayed, while those which were buried at a depth of from 
18 to 22 inches showed only decayed seed. Approximately all of the 
barley at the three different depths had germinated and afterwards 
decayed. 

The last fourteen species given in this table were marked ** decayed” 
when the seeds were taken up, but as the conditions were not so clearly 
indicated as in those first mentioned, germination tests were made in 
the greenhouse. 

The results of the germination tests show that none of the pots con- 
tained any viable seeds. Of this latter group only the pots containing 
the Asparagus officinalisand Bromus racemosus (Nos. 32 and 33) showed 
remains of old sprouts. The seeds in the other pots apparently had 
all decayed without any germination during the time they were buried. 
The germination of the asparagus seed had been almost perfect. The 
pot buried at the greatest depth contained only a mass of sprouts, 
many of which were still partially alive. The Bronius racemosus 
showed that germination had taken place only in the pots buried at 
6 to 8 and 36 to 42 inches, while those buried at the depth of 15 to 22 
inches had all decayed before germinating. 

It is interesting to note in this connection the behavior of the two 
species of Bromus—Bromus secalinus (cheat or chess) and 2. race- 
mosus (upright chess). The seeds of both of these species had com- 
pletely lost their vitality within the eleven months in the soil, while 
the contro] samples gave a germination of 95.5 and 92.5 per cent, 


14 VITALITY OF BURIED SEEDS. 


respectively. These differences are more clearly shown in Plate I, A 
and B. 

The results aboye stated, while perhaps not altogether conclusive, 
inasmuch as they represent only single tests of 200 seeds in each case, 
show that seeds of these two plants will not remain viable for long 
periods when buried in the soil. 

This is particularly interesting in the case of the common cheat or 
chess, which is frequently a pernicious weed in the grain fields of the 
United States. The generally accepted opinion is that the grains of 
cheat will live in the soil fora number of years, the seeds germinating 
when conditions are most favorable, the resulting plants then crowd- 
ing out the wheat. Some people even hold that in ‘‘off seasons” 
wheat turns to cheat, but fortunately such erroneous ideas are fast 
disappearing. 

The results of these experiments show that cheat, whenever found 
growing in grain fields or elsewhere, has come from seed recently 
sown and has not been lying dormant in the soil. With but few 
exceptions the unexpected appearance of cheat comes either from 
seeds that have been sown unintentionally mixed with wheat or other 
grains so that they passed unobserved, or from seeds that have been 
scattered with stable manure. 

Dr. Beal“ has also shown that buried seeds of Bromus secalinus do 
not retain their vitality for a long period of years. In Beal’s experi- 
ments the first test was at the expiration of five years, but not a single 
grain of cheat responded to the germination test at that time. 

Table II includes the majority of our more commonly cultivated 
plants of the field or garden, all of which failed to show any seeds 
capable of germination after having been buried in the soil for 
approximately one year. This statement will hold good for the major- 
ity of our cultivated plants. There are, however, a number of excep- 
tions. Many of these will be found in Table II], some showing that 
vitality was remarkably well preserved. Of these celery, parsnip, and 
tobacco (numbers 94, 95, and 99, respectively) should be mentioned in 
particular. The highest germination in each case was 64 per cent for: 
the celery from the 18 to 22 inch depth, 63 per cent for the parsnip 
from the 36 to 42 inch depth, and 70 per cent for the tobacco from 
the 18 to 22 inch depth. 


“B ulletan No. 5, Michigan Agricultural College, 1884. 


ee: Ne lal 


—— 


GERMINATION TESTS. 


TaB_eE III.—Results of tests of seeds that had not completely lost their vitality while 
buried, 


~ 


Je num- 
er, 


| aa 


PSSSEBRESSE 


SSSIFSGASSASSHARGLLVASSH 


= 
bo 
& 


16264 
16278 | 


16188 

16248 
16249 | 
16212 

16280 

16269 | 
16260 
16173 
16275 
16209 
16208 
16215 | 
16250 
16258 

16207 

16178 
16282 | 


Kind of seed. 


Chamber tests. 


Greenhouse tests in sand. 


ol 


Tem- 


Origi- 


Depth of burial. 


Con- Con- Se = — 
| Pera- | nal- trol trol 6-8 18-22 | 36-42 
vure, | paniple. inches. inches. inches. 
° C.._| Per ct..| Per ct. | Per ct..\ Per ct. | Per.ct..|Per ct. 
Mestnes elatior =. = .=-...----- 20-30 97 83 86 0.5 0.0 a0.0 
Capsicum annuum ..-.-....-.--- 20-30 §=696 97 80 .0 .0 Dd 
Brassica campestris...........- 20 90.25 86 18.5 .0 .0 b.5 
Trifolium Tepens .--.-.2.35.5.- ‘O 84.75 | 42.75 86 .0 b1 .0 
Wapit. caWjang... 520-2. 22---- 20-30 82.5 59.5 Tl oh? 0 1 .0 
Lespedeza frutescens.-.....---- 20. 15 el 2.5 .0 .0 1 
Ascyrum hypericoides .......- 30 1.5 .0 .0 1 .0 =6 
Lycopersicon lycopersicon ..... 20-30 99.25 72.5 88 .o 1 mS 
Chaetochloa glauca ......-...-- 20-30 | 55.75 - 37.5 18 1 1 1 
ES PARED 8 oo ees ose 20-35 .0 .0 .0 .0 .0 2 
CUCHMHS BATIVA 2.25.25 20.2 2: 20-30 100 98.5 62 .0 1 3 
Trifolium pratense ..........-.. 20} 89.75 | 73 85.5 2 4 4 
| Trifolium hybridum ........-.. 20 | 91.75) 84 73 P. 4 4.5 
| Xanthium pennsylyanicum .. 3015-02) 13.20.33 .0 .0 .0 a) 
| Cassia marylandica .-.......-- 30 14.5 | a98 20 3 3 5 
Axnubrosia trifida 222. 22---+.--- 20-30 , 29 52.5 48 .0 b2 6 
Trifolium pratense (10964) ...-. 2 PH Gialol==5 soe. 70 4.5 b5 6 
Vacearia vaccaria....--.....-- 20-35 | €6.5 8s 68 .0 b4 7 
| Convolvulus sepium ........-- 20-30 4 2 24 2 4 a 
Rudbeckia hirta ....-....-..-- 30. «69.5 78.5 74.5 6.5 6.5 7 
| Erysimum cheiranthoides .... 20-35 52.5 42 14.5 2 b5 bs 
Medicago sativa .-..-.....-.-- 20 | 84.5 64.5 97 b2 bg bg 
fe PnlIaspLearvense =5.2-. 2. 2-54. - 20-30 | 57.25 54.5 a) b11 8 11.5 
Solanum nigrum. ...........-- 20-30 | 97.75 91 12 9.5 10.5 1b 
Chenopodium hybridum...-.- 20-30 61 18.5 10.5 14D 9.5 13 
Cuscuta polygonorum .......- 20-30 | 12 8.5 55.5 11.5 10.5 3 
Sporobolus cryptandrus.....-- 30} 2.25 3 0 2D 1.5 13.5 
Plantago rugelii .............. 20-30 | 3.75 5.5 67.5 12 12 13.5 
Verbena hastata 2.22. s5-225=- 20-30} 9 Se Bae ae 11.5 13 14 
BIASICA DISLS os. = eo aces 20} 15 | 13.25) 34 10 b14 b14 
Trifolium pratense (hard) ..-. 20" 13a fal- 9,25.) > 18 610.5 15.5 14.5 
Elymus triticoides ..........-.- 20-30 84 75 85 1.5) 53.5 615.5 
Panicum virgatum......-...-. 20-30 | 30.5 36.5 22 7 17 16 
Avena fata .3: 252. 5h22--524<2 20-30 | 70.5 91.5 93.5 bg 68 18 
IBEGEVHIPATIO® 225. 20 - coe se ne 1. 20430) 158) 2-2 90.5 7 19.5 20 
Potentilla monspeliensis....... 20-30 41 83 73.5 b9.5 16 21.5 
Elymus canadensis........---- | 20-30 | 93.5 95.5 81 a) b7 b22 
Pom pratensis... .- 252-2525... -| 20-30: 90.75 87 59 16 224 24.5 
| Verbascum thapsus .:......... | 20-30) 82.5 98 72.5 7 7.5) 025.5 
Elymus virginicus....-.....--- | 20-35) 65.25 44 83 a2 513.5 625.5 
| Sisymbrium altissimum .....--! 20 | 88.25) 86.25) 76 610.5 VES 26 
Verbena urticifolia 30 135 .0 56.5 PSTN Woe Bs 26.5 
Carduus arvensis... - 20-30 56.75 | 68 5 74 SOS ee 28.5 
Ipomoea lacunosa...........-. 20-35 «98.5 | f 88 88 200 Fb 33 
Cuscuta epilinum ...-........- | 20-30 -0 | .0 .d 15.5 | 23.5 34 
Amaranthus retrofiexus.....-- 20-30 | 94.75 | 91 61 LSC ia 22. ob Be 
Bidens frondosa........--...-- 20-30 795 52.5 25 2925) 38 36 

eslia paniculata ...........-- 20-35 96 97 68 23 24.5 38.5 

ortulaca oleracea .....-...--. 35 | 83.75 | 91.5 16 39 38.5 | 30.5 
Ambrosia artemisiaefolia ..... 20-30 58.5 42.5 | 30.5 32 56 abe by Map 8 
Plantago lanceolata........... 82.5 78 | 67.5 A ab 41 
Grindelia squarrosa .........-. 20-30 | 25.75) 41 7.5 30.5 36 42 
Taraxacum erythrospermum -.| 20-30 85.75 | &7.5 85.5 35.5 41.5 45.5 
Pleniaro wajor...-3--:2... - 20-30 | 24 .| 78 0 39.5 43.5 |~ © 4625 
Chrysanthemum  leucanthe- } 

CLT ieee eee eee ee = 20-30 96.25 91 85.75 | 621 533 649.5 
Phalaris arundinacea ......... 20-35 | 69.25) 8 7 45 46.5| 56.5 
Apium graveolens...........-. 20-30 | 88 83.5 72.5 | 48.5 64 60 
Pastinaca sativa...........-..-- 20-30 | 55.5 67 78.5 29 51 63 
Chenopodium album.......... 20-30 (67.25 58 33.5 32 63.5 | 64.5 
Helianthus annuus (wild) ....| 20-30 100 97 86 43.5 644 66.5 
Bactoes scariola_-. 5... -.- =: 20 ~25.| 11.5 83 63.5 69 69.5 
Nicotiana tabacum............ 20-30 | 89.25 84.25] 89.25 46.5| 70 55 
Agropyron repems_...........- 20-30 | 80.24) 84 23.5. | 20.5] 673 | 66.5 
AretinM Lippas =< occas es fen - <i 20-30 99.75 | 96 88.5 | 42.5 63.5 | 73 
Rumex obtusifolius ........-... 20-30 | 97.5 95.5 80 73 | 72.6} 79.5 
MUMEX CTISPUS 25) 2255522252222 20-30 | 80.75 | 83.5 91 67.5 | 79.5 | 7 
Phytolaccaamericana......... 20-30 40.5 488.5 84.5 7.5| 66.5 80.5 
Fraxinus americanus.......... 20-30 49.5 2 | 26 -0} 0 | S4 
Dative trie 32-2525 3055 -- 20-30 99 d54 | 88 L GSGh par eage| 86.5 
Rumex salicifolius ............ 20-30 | 98.25) 96.5 | 2.5 | 88.5| 85.5 70.5 
Chaetochloa verticillata ...... 20-30 92.75 | 94.5 88.5 | b58 71 90 
Onopordon acanthium ........ 20-30' 95.5 | 31 (0.5 Shee? as. 90.5 


16 VITALITY OF BURIED SEEDS. 


Tasre III.—Results of tests of seeds that had not completely lost their vitality while 
buried—Continued. 


Chamber tests. Greenhouse tests in sand. 
Labora- : 
tes z | urial. 
test Kind of seed. | Tem- | Origi- | Gon. | con- | Depth oa 7 
bent pera- | nal | trol. | trol. | 68 | 18-22 | 3642 
sie epee inches. inches. inches. 
iC) Per ct. \Per et." || Per ices | Per ct. | Per ct. \erer che 
110.|' 16218 |\-Alsine media, --en20 se eee ee 90-801) 97 | 98.6 | 198 90.5 | 96.5 92.5 
111g] 16243 | Abutilon abutilon.....2../....)........ Beers Perr orate Pe a 
1129} 16189 | Phleum pratense .............. PEs Bee eae [parece tose | 0 = sa eS oa ee oe ftttese 
Average percentage of | 
Permingtions sles es ot eleses sees 6392") |tbFa5 53/24] sn ee 26.5 31 
; 


aMany had germinated and afterwards decayed. i 

b Fresh sprouts found when samples were taken up. These sprouted seeds were not thrown away, 
but were transplanted with the remainder of the sample and tested in sand in greenhouse, conse- 
quently those which produced seedlings are included in the percentages of germination given in the 
table. These fresh sprouts were found as follows: 


Sam- Sam- | Sam-| Sam- 


= 1 i} 
cnt Depth. | Sprouts. BAG Depth.| Sprouts. | cae Depth. Sprouts. we Depth. | Sprouts. 
ber. ber. | ber. ber. 

| Inches. Inches. | | Inches. | Inches. 
39 36-42 1 59 36-42 1 71 18-22 Many. | 92 18-22 Many. 
53 | 18-22 3 60| 6-8 1 73. AGG 4 92 | 36-42 Many. 
54 18-22 1 67 18-22 2 7A 18-22 Few. | 100 18-22 |. 1 
55 18-22 1 67 36-42 2 74 36-42 Many. || 106 6- 8 2 
58 | 18-22 10 | 68| 68 1|| 77| 13-22 io || 108| 6-8| Many. 
58 | 36-42 5 69 | 18-22 Few. || 77 | 36-42 5 | 
59 6- 8 4 69 36-42 Many. || 78} 6-8 I 
59 | 18-22 2 71| 68 Many. || 92) 6-8 Many. | 

| | | | 


eClipped seed germinated, 59 per cent. 
aClipped. 

t Gloped eed me nie en 1 NOPE r cent. 
g Tests interrupted. 

In Table III the names of the seeds are arranged in the order of 
their vitality as determined by the germination tests made in the 
greenhouse. The list. of seeds tested begins with Festuca elatior 
(meadow fescue), which showed only one viable seed, that being from 
the 18 to 22 inch depth, and ends with -A/s/ne media (common chick- 
weed), in which nearly all of the seeds retained their power of germi- 
nation throughout the entire period that they remained in the soil. 
The germination of the latter, when sown in the greenhouse, was 
almost perfect. (See Pl. II, fig. 1.) 

In many instances some of the seeds had germinated while they were 
buried. In most cases the seeds which had germinated afterwards 
decayed. In the larger seeds this could usually be determined with- 
out much difficulty, but with many of the smaller seeds no such obser- 
vations could be made. However, it is more than probable that many 
of the smaller seeds which showed a low germination when trans- 
planted in the greenhouse had germinated and afterwards decayed 
before being dug up, but this could not be satisfactorily determined 
by a hurried field examination. Many of the pots also contained fresh 


RELATION OF DEPTH OF BURIAL TO VITALITY. 16 


sprouts at the time the seeds were taken up. The number of fresh 
sprouts in each case is indicated in a footnote to Table III. 

Unlike Table II, Table III includes names of but very few of our 
cultivated plants. The majority belong to that class of plants com- 
monly known as weeds. These results show that but a limited num- 
ber of our cultivated plants produce seeds which can retain their 
vitality for any length of time when buried in soil. On the other 
hand, the seeds of the plants which are commonly known as weeds are 
of strong vitality, and many of them deteriorated but very little with 
the treatment given. This, of course, is what we would expect. By 
natural selection the wild plants which survive are just those from 
seeds which are capable of living in the soil for a period of time more 
or less extended, and ultimately this factor becomes hereditary. With 
most of the cultivated plants the seeds are gathered and carefully 
saved from year to year, resulting in the loss of these inherited 
characteristics. 

The mere fact that certain seeds retain their power of germination 
for a period of years when buried in the soil brands the plants which 
they produce as weeds. The length of time that such seeds can remain 
in the soil and still retain their power of germination largely deter- 
mines their noxiousness. In other words, it may well be said that the 
pernicious character of weeds is directly proportional to the length of 
time the seeds will remain viable when buried in the soil. For this 
reason bad weeds are difficult to eradicate once the seeds are allowed 
to mature. (See Pl. II.) 


RELATION OF DEPTH OF BURIAL TO VITALITY. 


Table III shows that many of the seeds were better preserved the 
deeper they were buried. This is probably best explained by the 
difference in the three factors which govern germination, viz, heat, 
moisture, and oxygen. At the greatest depth the amount of moisture 
is always more uniform, the supply of air is greatly lowered, and the 
temperature ismuch reduced. The temperature decreases very rapidly 
as we go below the surface of the soil, and at 33 feet is comparatively 
uniform throughout the year. Experiments conducted at McGill 
College, Montreal, Canada, by C. H. McLeod show that at a depth of 
40 inches below the surface of the soil the minimum and maximum 
temperatures through the year were approximately 35° and 60° F., 
respectively.? 

The greater number of seeds germinate best when subjected to daily 
alternations in temperature. These alternations do not take place ata 
depth of 3 feet below the surface; consequently there is a better 


@Trans. Roy. Soc., Canada, Ser. 2, Vol. 7, Sec. III, pp. 13-16, 1901. 


18 VITALITY OF BURIED SEEDS. 


preservation of vitality at that depth as a result of the more dormant 
condition of the seeds. (See Pl. III and the diagram below.) 

As was anticipated, most of the seeds which were stored in the Seed 
Laboratory preserved their vitality much better than those that were 
buried. But there are a number of cases in which the seeds were 
preserved practically as well in the soil as in the laboratory, the deteri- 
oration being very small in either case. However, with but few 
exceptions, an ample number of seeds remained germinable at the ter- 
mination of the first year to produce plants in sufficient number to keep 
the up-to-date farmer busy for a good share of the summer in sup- 
pressing them. The average percentages of germination of all samples, 
including the original test and both controls, are best shown in the 
following diagram: 


Average germination of controls and buried seeds. 


Original tests, 63.2 per cent. 


Controls (chamber), 57.5 per cent. 


59 9 


Controls (greenhouse), 53.2 per cent. 
5 ’ 


Buried 6-8 inches, 20.5 per cent. 


Buried 18-22 inches, 26.5 per cént. 


Buried 36-42 inches, 31 per cent. 


HARD SEEDS. 


An interesting point in these first results is in the behavior of the 
Trifoliums and closely related genera, including Lespedeza and Medi- 
cago. Generally speaking, these seeds are considered to be able to 
withstand very critical treatment, but the results of the first year’s 
experiments show that the seeds of all of these deteriorated very greatly 
while in the soil. 

The white clover, No. 41, germinated only 1 per cent, and showed 
one fresh sprout when taken up from the 18 to 22 inch depth and 
nothing from the shallower or deeper trenches. The red clover did 
but little better; No. 49, a sample of the harvest of 1902, germinated 
2, 4, and 4 per cent for the three different depths of 6 to 8, 18 to 22, 
and 36 to 42 inches, respectively. Another sample of red clover, No. 
54, germinated 4.5, 5, and 6 per cent, respectively, for the three 
different depths. A third sample of red clover, No. 68, germinated 
10.5, 15.5, and 14.5 per cent, respectively, from the three depths. The 
last two samples were of Oregon-grown seed of the harvest of 1900. 
The original sample of this seed, No. 54, contained 51.5 per cent of 


SUMMARY. 19 


hard seed. No. 68 includes only the hard seed selected from the 
Oregon clover by soaking in water for 18 and then for 20 hours a 
portion of the original bulk sample, using only the remaining hard 
seed. 

These results, while unsatisfactory, show clearly that it is the hard 
seeds in the clovers which remain over in the soil for some considerable 
time. The alsike clover, No. 50, behaved practically the same as the 
sample of red clover first mentioned. The Lespedeza, or bush clover, 
No. 43, gave results very similar to the white clover. The alfalfa, No. 
59, gave a germination of only 2, 9, and 9 per cent, respectively, for 
the three different depths. But in all cases a few fresh sprouts were 
present when the seeds were taken up, showing that the seeds were 
germinating and afterwards decaying. 


SEEDS OF CULTIVATED VERSUS WILD PLANTS. 


A number of interesting cases showing the greater hardiness of the 
seeds of wild plants over those of like or closely related cultivated 
forms were recorded. In //elianthus annuus (Nos. 6 and 97) the seeds 
from the cultivated plant—our common sunflower of the garden—uall 
decayed, while the seeds of the wild sunflower retained their vitality and 
germinated 43.5, 64, and 66.5 per cent, respectively, for the three dif- 
ferent depths. Similarly with Lactuca sativa and Lactuca scariola, Nos. 
5 and 98, respectively, the common garden lettuce seed had all decayed, 
while the seed of the prickly lettuce, possibly the parent of our cos 
varieties, germinated 63.5, 69, and 69.5 per cent, respectively, for the 
three different depths. Another striking example is in Avena sativa, 
No. 1, and Avena fatua, No. 71, the latter germinating 9, 8, and 18 
per cent, respectively, for the three different depths, besides showing 
many fresh sprouts in the two shallower depths at the time the seeds 
were taken up. 

Furthermore, it is not uncommon to find wide variations in different 
species of the same genus, even where all forms are wild, e. g., Elymus, 
Chaetochloa, Chenopodium, Cuscuta, Plantago, etc. But in the cases 
above mentioned of the cultivated and the closely related wild forms 
the ability of the seeds to withstand such treatment as being buried in 
the soil has been lost by long cultivation of the plants and the careful 
preservation of the seeds under artificial conditions or storage, while 
seeds from the wild forms can survive when buried in the soil, for it 
is the plants from just such seeds that have survived. 


SUMMARY. 


The length of time that seeds will retain their vitality when buried 
in the soil is of much importance in the extermination of weeds. 

The seeds of many of our pernicious weeds can be destroyed by 
deep plowing, if the soil is left undisturbed for some time. 


20 VITALITY OF BURIED SEEDS. 


Seeds of the cultivated plants, with but few exceptions, lose their 
vitality when buried in the soil. 

Seeds of the plants commonly designated as weeds retain their vital- 
ity remarkably well when buried in the soil. 

In general, the pernicious character of weeds is directly proportional 
to the length of time the seeds will remain viable when buried in the 
soil. 

The deeper seeds are buried, the better is vitality preserved. 

Hard seeds of the same species retain their vitality much better 
than those with softer seed coats. 

Unbulled seed, especially of the grasses, is more resistant than hulled 
seed, and the vitality is always better preserved. 

Seeds of plants from the same genus often retain their vitality in a 
very different degree. . 

Vitality is best preserved, even in weed seeds, when the seeds are 
carefully harvested and stored in a dry and comparatively cool place. 


DESCRIPTION OF PLATES. 


Piate I. Fig. 1.—Bromus racemosus, smooth brome grass. Fig. 2.—Bromus secalinus, 
cheat or chess. The two divisions at the right of each figure show the vigorous 
growth made by the check samples. In the three divisions at the left, A, B, and 
C, were planted the seeds which had been buried at depths of 6 to 8 inches, 18 
to 22 inches, and 36 to 42 inches, respectively. The vitality of the seeds of these 
two species, which are considered as noxious weeds in the grain fields of the 
United States, was destroyed at the expiration of eleven months. 


Piare II. Fig. 1.—Alsine media, common chickweed. Fig. 2.—Rume.x crispus, curled 
dock. Fig. 3.—Datura tatula, jimson weed. Seedlings from weed seeds which 
did not lose their vitality by burial for eleyen months, as shown in the three 
divisions at the left of each flat, the germination being practically the same as 
in the case of the two check samples shown at the right of each flat. 


Prate IIL. Fig. 1.—Llymus canadensis, nodding wild rye. A, buried 6 to 8 inches— 
all killed; B, buried 18 to 22 inches—only one seedling shows in the figure, but 
the total germination was 7 per cent, as given in the table; C, buried 36 to 42 
inches—germinated 22 per cent; the two check samples at the right made vigor- 
ous growth, germinating 81 per cent. Fig. 2.—Fraxinus americanus, white ash. 
A, buried 6 to 8 inches, and B, buried 18 to 22 inches—all killed; C, buried 
36 to 42 inches—germinated 84 per cent; the check samples germinated 26 per 
cent, but the seedlings had ‘‘damped off’’ before the photograph was taken. 
Fig. 3.—Phytolacca americana, poke. <A, buried 6 to 8 inches—germinated 7.5 
per cent; B, buried 18 to 22 inches—germinated 60.5 per cent; C, buried 36 to 
42 inches—germinated 80.5 per cent; the two check samples germinated 84.5 per 
cent. 


The illustrations show that in many cases the vitality of seeds is better preserved 


at a depth of 36 to 42 inches than at shallower depths. 
29 


O 


Bul. 83, Bureau of Plant Industry, U. S. Dept. of Agriculture. PLATE Il. 


Fic. 2.—BROMUS SECALINUS (CHEAT, OR CHESS). 


Bul. 83, Bureau of Plant Industry, U. S. Dept. of Agriculture. PLATE Il. 


Fic. 1.—ALSINE MEDIA (COMMON CHICKWEED). 


Fig. 2.—RUMEX CRISPUS (CURLED DOCk). 


Fig. 3.—DATURA TATULA (JIMSON WEED). 


™ 


a wt 


os 


Bul. 83, Bureau of Plant Industry, U. S. Dept. of Agriculture PLATE III. 


Fig. 1.—ELYMUS CANADENSIS (NODD!ING WILD RYE). 


FIG. 2.—FRAXINUS AMERICANA (WHITE ASH). 


FIG. 3.—PHYTOLACCA AMERICANA (POKE). 


“ 4 


U. S. DEPARTMENT OF AGRICULTURE. 
BUREAU OF PLANT INDUSTRY—BULLETIN NO. 84. 


B. T. GALLOWAY, Chief of Bureau. 


THE SEEDS OF THE BLUEGRASSES. 


I. THE GERMINATION, GROWING, HANDLING, AND ADULTERATION 
OF BLUEGRASS SEEDS. 


By Epa@ar Brown, Botanist in Charge of Seed Laboratory. - 


II. DESCRIPTIONS OF THE SEEDS OF THE COMMERCIAL BLUEGRASSES 
AND THEIR IMPURITIES. 


By F. H. HitiMan, Assistant Botanist, Seed Laboratory. 


IssuED NovEMBER 14, 1905. 


WASHINGTON: 
GOVERNMENT PRINTING OFFICE. 
1905. 


BUREAU OF PLANT INDUSTRY. 


B. T. GALLOWAY, 
Pathologist and Physiologist, and Chief of Bureau. 


VEGETABLE PATHOLOGICAL AND PHYSIOLOGICAL INVESTIGATIONS. 


AuBERT F. Woops, Pathologist and Physiologist in Charge, Acting Chief of 
Bureau in Absence of Chief. 


BOTANICAL INVESTIGATIONS. 
FREDERICK V. Covi1LLE, Botanist in Charge. 


FARM MANAGEMENT. 
W. J. Spruuman, Agriculturist in Charge. 


POMOLOGICAL INVESTIGATIONS. 
G. B. Brackett, Pomologist in Charge. 


SEED AND PLANT INTRODUCTION AND DISTRIBUTION. 
A. J. Prerers, Botanist in Charge. 


ARLINGTON EXPERIMENTAL FARM. 
L. C. Corsert, Horticulturist in Charge. 


INVESTIGATIONS IN THE AGRICULTURAL ECONOMY OF TROPICAL AND 
SUBTROPICAL PLANTS. 


O. F. Coox, Bionomist in Charge. 


DRUG AND POISONOUS PLANT INVESTIGATIONS, AND TEA CULTURE 
INVESTIGATIONS. 


Ropney H. True, Physiologist in Charge. 
DRY LAND AGRICULTURE AND WESTERN AGRICULTURAL EXTENSION. 
Cari 8. ScorreLp, Agriculturist in Charge. 
EXPERIMENTAL GARDENS AND GROUNDS. 


KE. M. Byrnes, Superintendent. 


SEED LABORATORY. 
Epa@ar Brown, Botanist in Charge. 


J. E. Rockwe uy, Editor. 
James KE. Jones, Chief Clerk. 


SEED LABORATORY. 
SCIENTIFIC STAFF, 
EpGcar Brown, Botanist in Charge. 


F. H. Hituman, Assistant Botanist, J.W. T, Duvet, Assistant, 
2 


LETTER OF TRANSMITTAL. 


U.S. DEPARTMENT OF AGRICULTURE, 
BureEAvu OF Puiant InNpustTry, 
OFFICE OF THE CHIEF, 
Washington, D. C., July 15, 1905. 

Str: I have the honor to transmit herewith and to recommend for 
publication as Bulletin No. 84 of the series of this Bureau the accom- 
panying technical paper entitled ‘‘The Seeds of the Bluegrasses.” 

- This paper was prepared by Mr. Edgar Brown, Botanist in Charge 
of the Seed Laboratory, and Mr. F. H. Hillman, Assistant Botanist, 
Seed Laboratory, and has been submitted with a view to publication. 

The bluegrasses are among the most important forage plants in 
many sections of the United States and Europe, and large quantities 
of seed are harvested annually for use in this country and for expor- 
tation. 

The process of cleaning the seed of the bluegrasses for market is 
such that many of the distinguishing characters are lost, and separate 
descriptions are necessary for the hand-picked and commercial seed of 
the same species. 

The seeds of the different commercial species are so nearly alike in 
general appearance that at present none but the trained observer can 
distinguish between them. This similarity of appearance has encour- 
aged the use of the cheaper and less desirable species, especially 
Canada bluegrass, for the adulteration of or substitution for the more 
expensive species. 

The descriptions and illustrations herewith given of the bluegrasses 
and of their impurities will be of great value in furnishing seedsmen 
the necessary information to enable them to distinguish the different 
species. 

The accompanying illustrations are necessary for a complete under- 
standing of the text. 

Respectfully, B. T. GatLoway, 
Chief of Bureau. 
Hon. James Witson, 
Secretary of Agriculture. 


be orermantertiinantien 


CONTENTS. 


I. THE GERMINATION, GROWING, HANDLING, AND ADULTERATION OF BLUE- 
Cras woObeEnss a by sldrar browne +sa22 22224 atone oe eh Sree 
Description of commercial and hand-gathered seeds. -..--..---------- 
Grades and quality of commercial seeds. ---...--.------------------ 
UMMM tte le oe Bathe Site Se SOS RAS NS, fe DLE aie me Se 

0 ETP Ugo L005) Sa BR Ge ao Mage Piece Pk SR aes pe 
OS EDEL  M ant 
Semmens EPPHETA NO eS Soe ere ts El eed ee Ao Jacke deel ke ae 
Pon pracensis ( Kentucky: bluegrass) 2 5.. ..-.22%..2.-2lde 2. o2 

Poa compressa’ (Canada bltiegrass) >. 2..-2........-.-2222.-2-2. 

Poa trivialis (rough-stalked meadow grass) 

Poa nemoralis (wood meadow grass) 
Paatrpors (lowl. meadow erans) 25252555. 26s 2 chee we ooo 

Poa arachnifera (Texas bluegrass ) 
aaannnda (annual blneprass) 24-2. £2. osdeus.. 2-2. es 25-38 
ooalpma (alpine meadow erass) 2. 2. 2.222. -.<-.--.----.-2--2 

1G SOTO BU ASSES eat Bk ak a rr 


II. Descriprions OF THE SEEDS OF THE COMMERCIAL BLUEGRASSES AND THEIR 
meiintisas vn. 81 AEP 6 Noo oe heh cee 250 enon es 
CODE SCTE 2 gn a eS eae Ne eee eee See 
Key to the seeds of the more common species of Poa as found 
Giniaer DPATMEN SPEGMNCHA 15 227 5. 2 eS ee le coe cece e oce 

Key to commercial bluegrass seeds after preparation for market. - 
Comparison of the principal distinguishing characters of blue- 

PLCS ES ee 2 a ee ie opt = rn Oe ee re ae ae 

Beier ip OL BDEGIER 95026 See en et ae 2 SoG ek Loe 

Poa pratensis L., Kentucky bluegrass, June grass......-..-- 

Poa compressa L., Canada bluegrass, flat-stemmed bluegrass. - 


Poa triflora Ehrh. (P. flava L., P. serotina Ehrh.), fowl 
meadow grass, false redtop.-.....--..- 
Poa arachnifera Torr., Texas bluegrass ..-......:.-...---.-- 
Poa annua L., annual meadow grass 
Poa alpina L., alpine meadow grass 
Evi SUCEICIDEVOPTICE ooo Sane en ee Fae. 2 oe Be 2S bee 
TULLE REVDD eS ot ek 8 ee Bae © Oa a a PACE 
Panicularia nervata ( Willd.) Kuntze, nerved manna grass 
sometimes called fowl meadow grass...........--.--- 
Panicularia americana (Torr.) MacM., reed meadow grass, 
. water meadow grass, tall manna grass 


? 


Page. 


oOo © 


10 
10 
11 
12 
12 
12 
13 
13 
13 
13 
14 
14 
14 
14 


15 
15 


18 
19 


20 
22 
22 
24 
24 
26 


27 
28 
29 
29 
30 
31 


31 


31 


6 CONTENTS. 


IJ. DrescrIpTions OF THE SEEDS OF THE COMMERCIAL BLUEGRASSES AND THEIR 


Impurities—Continued. Page. 
Weed seeds commonly found with commercial bluegrass seeds ----.- 32 
Bursa bursa-pastoris (L.) Britton, shepherd’s-purse .....-------- 32 
Lepidium virginicum L., peppergrass -...----------------------- 32 
Cerastium vulgatum L., mouse-ear chickweed.-....-------------- 32 
Alsine media LL., common ehickweed - --......--- = =. ae eee 32 
Alsine gramines: (::)/Britton .- -... - - 2. =. 2 = eee 33 
Carduus arvensis (L.) Robs., Canada thistle --......--..-------- 33 
Taraxacum taraxacum (L.) Karst., dandelion........----------- 34 
Matricaria inodora L., scentless camomile ........-------------- 34 
Hieracium sp., hawkweed -..--....-2.---1 J... 5s. 34 
Anthemis cotiula L., dog fennel, mayweed.......---------------- 35 
Chenopodium album L., lamb’s-quarters, pigweed....-..--------- 35 
Plantago lanceolata L., rib-grass, buckhorn, English plantain. - - - 35 
Rumez crispus L., curled dock -.-:.-----..¢025: 2.525250 36 
Rumezx acetosella L., sheep’s sorrel, sorrel.....------------------ 36 
Veronica arvensis L., corm speed well-. ........-.-:22 222 eee 36 
Juncus tenuis Willd., slender rush .--.---22--2-=2-2e sees 37 
Juncoides campestre (L.) Kuntze, field rush.............----.--- 37 
Juncoides albida DC:, wood rush”. -. 22. 222.2222 22 eee 37 
Carex cephalophora Muhl., oval-headed sedge. .-.--.---.-------- 37 
Ergot occasionally found in commercial bluegrass seed. .........---- 38 


Claviceps purpurea (Fr.) Tul., ergot. -.....----2- 22 2 oe 38 


ILLUSTRATIONS. 


TEXT FIGURES. 


meeseikelenor Pos. 2225-22-25 2- 2-2 e cSt. 
. Unrubbed Kentucky bluegrass seed (Poa pratensis) ....--.-.-------- 
. Seeds of Kentucky bluegrass (Poa pratensis) - . - - 
. Different forms of commercial seeds of Kentucky bluegrass (Poa 


FRAT) Eos Be SAE eek ee ee 


A 


. Commercial seeds of Canada bluegrass (Poa compressa) ...---.------ 


Seeds of rough-stalked meadow grass ( Poa trivialis)........--.------ 


. Seeds of wood meadow grass (Poa nemoralis) - - - - 
. Seeds of fowl meadow grass ( Poa triflora).-.-.---- 
. Seeds of Texas bluegrass (Poa arachnifera) - - -- - - 
. A cluster of Texas bluegrass seeds matted by the webby fibers. -----. 
. Seeds of annual meadow grass ( Poa annua). ----- 
. Seeds of alpine meadow grass (Poa alpina) .----- - 
BPE EO EMG RUMCICO S22 Sone Soe joe Sw ke on 
. Seeds of nerved manna grass ( Panicularia nervata) ....--.---------- 
. Seeds of water meadow grass ( Panicularia americana) ...-.---------- 
. Seeds of shepherd’s-purse ( Bursa bursa-pastoris) - 
. Seeds of peppergrass (Lepidium virginicum) ---- - 
. Seeds of mouse-ear chickweed ( Cerastium vulgatum)....------------- 
. Seeds of chickweeds ( Alsine media and A. graminea)...--..--------- 
. Seeds of Canada thistle ( Carduus arvensis) - ----- - 
. Prickles often found with bluegrass seed--.----- 
. Seeds of dandelion (Taraxacum taraxacum).--.--- 
. Seeds of scentless camomile ( Matricaria inodora) - 
. Seeds of hawkweed (Hieraciumsp.)-.----.------- 
. Seeds of dog fennel (Anthemis cotula) ......----- 
. Seeds of lamb’s-quarters (Chenopodium album) - - - 
. Seeds of rib-grass ( Plantago lanceolata) ...-.----- 
. Seeds of curled dock ( Rumex crispus) ..--------- 
. Seeds of sorrel (Rumesx acetosella).....--.------- 
. Seeds of corn speedwell ( Veronica arvensis) - -- - - - 
. Seeds of slender rush (Juncus tenwis)......------ 
. Seeds of field rush (Juncoides campestre) .--.----- - 
. Seeds of wood rush (Juncoides albida) ...-------- 
. Seeds of sedge ( Carex cephalophora) .....-------- 
. Ergot (Claviceps purpurea) of Kentucky bluegrass 


31 


32 


B. P. I.—176. 


fae SEEDS OF THE BLUEGRASSES. 


I. THE GERMINATION, GROWING, HANDLING, AND 
ADULTERATION OF BLUEGRASS SEEDS. 


By Epcar Brown, 


Botanist in Charge of Seed Laboratory. 


DESCRIPTION OF COMMERCIAL AND HAND-GATHERED SEEDS. 


Great difficulty is experienced in distinguishing the seeds of the 
species of Poa. It is especially important to be able to recognize 
them, as the species vary greatly in value and the seed of one species 
is frequently substituted for that of another. 

The descriptions of the seeds of Poa already published have been 
largely those of complete or hand-gathered specimens. But the seeds 
of some kinds as they appear on the market are more or less broken 
and have lost many of their distinguishing characters. The process 
of cleaning often rubs off the web at the base of the seed and the 
hairs along the sides and breaks the tip. On this account descriptions 
based on specimens of perfect seeds are not to be relied upon in 
identifying certain commercial Poas. 

The mutilation of seeds during the process of cleaning is especially 
marked in home-grown seed of Kentucky bluegrass (Poa pratensis). 
Even the hand-gathered seed of rough-stalked meadow grass (Poa 
trivialis) is frequently so much injured about the slender apex as to 
increase greatly the difficulty of distinguishing it from that of Ken- 
tucky bluegrass. On the other hand, the commercial seeds of wood 
meadow grass (Poa nemoralis) and fowl meadow grass (fa triflora) 
retain much of the pubescence on the glume, often the web, and are 
usually not broken on the tip. 

It is important that descriptions and illustrations to be used in prac- 
tical seed testing be taken from the commercial as well as hand- 
gathered seed and be comparative in character. Those given in this 
paper have been prepared from both hand-gathered and commercial 
seed. The term seed is here used in its popular sense. 

5813—No. 84—05 2 9 


10 THE SEEDS OF THE BLUEGRASSES. 


GRADES AND QUALITY OF COMMERCIAL SEEDS. 


The seeds of all species except Kentucky bluegrass are known to the 
American trade in only one grade. This is the so-called ‘‘faney” 
grade, which is based on relative cleanness and on the bright appear- 
ance of the seed. The quality of different samples passing under this 
grade name is not necessarily uniform, but among the more careful 
dealers a purity standard of from 80 to 90 per cent is usually maintained. 

The seeds of Kentucky bluegrass and of Canada bluegrass raised in 
this country are usually much cleaner and freer from foreign seeds 
than the European-grown seeds of rough-stalked meadow grass, wood 
meadow grass, and fowl meadow grass. 

Kentucky bluegrass seed is commonly offered in two grades— 
‘*fancy,” and ‘*extra-clean” or ‘‘extra-cleaned.” The latter names are 
a survival of the time when the seed was hand cleaned and the ‘‘ extra- 
clean” was the best seed on the market. With the advent of improved 
machinery the ‘‘fancy” grade was established and it is now the only 
grade generally accepted by the intelligent purchaser. The ‘*extra- 
clean” still on the market belies its name, since it consists of the chaff 
or cleanings from the fancy seed, and consequently contains only light 
seed. Samples of ‘‘extra-cleaned” as offered usually contain less than 
10 per cent of seed. 

In some cases the growers find a sale for the rough or uncleaned 
seed after it has been passed through a feed cutter. In this condition 
it has very much the appearance of fine-cut straw with a large per- 
centage of chaff, and can be scattered over pastures and other areas, 
seeding them as effectually as could be done by the use of fancy 
recleaned seed. If well cured, the germinating quality of such seed 
is excellent, and the mass contains from 60 to 70 per cent of pure seed. 
Except for foreign trade the percentage of germination has little to do 
with the price and grade of bluegrass seed. 

Aside from adulterated samples the purity of ‘‘ fancy” seed of all 
species of bluegrass is usually good. Of the 2,887 samples of Ken- 
tucky bluegrass tested by the Zurich Seed Control Station from 1876 
to 1903 the average purity was 86.3 percent. Of the 69 samples 
tested in the Seed Laboratory of the Department of Agriculture 
during the past year the average purity was 75.02 per cent. 


ADULTERATION. 


The seed of Canada bluegrass (Poa compressa) is the only kind 
used as an adulterant of Kentucky bluegrass in this country. During 
the year 1904 649,451 pounds of Canada bluegrass seed were imported 
from Canada, practically none of which is being sold under its true 
name. Among the samples of seed sold for Kentucky bluegrass and 
sent to the Seed Laboratory for examination a large number have 


WEIGHT PER BUSHEL. 11 


contained from 30 to 50 per cent of Canada bluegrass seed and several 
have been entirely composed of the Canada seed. 

It is significant in this connection that the price of Canada bluegrass 
seed varies with that of Kentucky bluegrass seed, being usually about 
one-half that of the latter. This adulteration is not merely a simple 
fraud by which the farmer pays for what he does not get, but the 
difference in the resulting pasture or hay crop is very important. 
Canada bluegrass, while having many good qualities in common with 
other species of Poa, is by no means a pasture grass, for which purpose 
Kentucky bluegrass is unexcelled. 

The seed of wood meadow grass (Poa nemoralis) is sometimes adul- 
terated with other species of Poa, and samples have been offered under 
this name that contain no wood meadow grass seed. One sample tested 
in the Seed Laboratory contained 59.4 per cent of Poa pratensis and 
23 per cent of Poa compressa, the remainder being chaff and dirt. 
Samples of fowl meadow grass (Poa triflora) have been examined 
which consisted largely of various common grass and clover seeds 
combined with an abundance of weed seeds. These samples contained 
small quantities of Kentucky and Canada bluegrass seeds, much chaff 
and dirt, and some of them no seeds of fowl meadow grass. 

The seed of Kentucky bluegrass is used to adulterate that of the 
higher priced Poa trivialis, pure seed of the latter species usually 
being hard to obtain. Some of the German authorities say that it is 
necessary for every farmer to save his own seed of this grass in order 
to be sure that itis pure. Hunter” says: 

Previously to 1883 good and genuine seed of this species (Poa trivialis) could not 
be obtained in this country [England]. Seed of the Poa pratensis was commonly 
supplied for it. It is now less difficult to procure genuine seed, but large quantities 
of seed of Poa pratensis (which usually costs about one-third the price) are prepared 


to resemble and are sold for Poa trivialis, and it is only by careful microscopic exam- 
ination that the nature of the seed can be determined 


WEIGHT PER BUSHEL. 


The standard weight of a bushel of bluegrass seed of any grade is 14 
pounds. The actual weight, however, varies from 6 to 8 pounds in the 
case of *‘extra cleaned” to 27 pounds or more for especially good sam- 
ples of fancy recleaned seed. In the bluegrass region of Kentucky itis 
the usual practice to sell the seed fresh from the strippers or cured in 
the chaff by the bushel of 14 pounds, but it is always weighed, not 
measured. The cleaned seed is always sold by the pound. As the 
weight per bushel of bluegrass seed depends directly on its purity, 
it is customary in quoting the price of *‘fancy” seed to accompany it 
with a statement as to the weight per bushel. 


4Treatise on Permanent Pasture Grasses, James Hunter. Chester, England, 1901. 


12 THE SEEDS OF THE BLUEGRASSES. 


The foreign trade is much more critical than the domestic trade, and 
the seed exported usually weighs from 22 to 24 pounds per bushel, 
while the domestic trade is content with seed weighing from 18 to 20 
pounds. The heavier seed costs more per pound than the lighter seed, 
since there is more labor in its preparation, but it is cheaper for the 
purchaser. 

GERMINATION. 


The germination of commercial bluegrass seed is often poor. At 
the Zurich Seed Control Station the average percentage from 3,069 
samples of Kentucky bluegrass seed tested from 1876 to 1904 was 65 
per cent, while 908 samples of Poa trivialis tested showed an average 
of 72 per cent. The quality of Kentucky bluegrass seed as respects 
germination appears, however, to be improving. Last year’s tests at 
the Zurich station gave an average of 68 per cent, while a few years 
ago 50 per cent was considered fair or satisfactory. Only the best 
seed goes to Europe, and consequently the percentage of germination 
of that offered in this country is low. As carefully cured seed will 
germinate from 80 to 90 per cent, the cause for the poor quality of 
commercial seed is doubtless to be found in the way it is harvested and 
cured.” The usual process is to pile the freshly stripped seed in ricks, 
either outdoors or in barns. This mass heats quickly if not stirred 
often during the first few days. One pile left without stirring reached 
a temperature of 140° F. in sixteen hours, killing all the seed. 


GROWING AND HANDLING. 


With the exception of our native western species, more or less seed 
of all the commercial Poas is gathered in Europe, where they are 
found wild. The harvesting is done by hand from the natural mead- 
ows, woods, or other uncultivated areas. The seed is cleaned by hand 
and carried to market in small quantities and collected by dealers who 
supply the trade. The United States furnishes Europe with Kentucky 
bluegrass seed, and Europe furnishes the seed of rough-stalked meadow 
grass and wood meadow grass, as well as of the other commercial 
species of Poa used in this country. 

Poa pratensis (Kentucky bluegrass).—The bulk of the Kentucky 
bluegrass seed comes from a limited area known as the bluegrass 
region of Kentucky. The counties of Bourbon, Scott, Fayette, Clark, 
and Woodford furnish most of it, although there is a small quantity 
saved in Shelby County. Some is harvested in southwestern Illinois, 
and there is another area on the border between Missouri and Iowa 
where a considerable amount of seed is saved. The seed is gathered 
from the natural woodland pastures as well as from those where it has 


“See Bulletin No. 19, Bureau of Plant Industry, ‘‘ Kentucky Bluegrass Seed: Har- 
vesting, Curing, and Cleaning.”’ 


GROWING AND HANDLING. 13 


been sown. It is customary to graze cattle on it nearly the entire 
year, as they do not materially injure the crop of seed if they are kept 
out for two or three weeks immediately before gathering. The seed 
is harvested by pulling the heads off with a stripper, the grass not 
being cut for hay. The cleaning is a rather difficult process, as it is 
necessary to rub the heads thoroughly in order to separate the seed 
from the web at the base. The last of the chaff and dirt which is 
blown out during the cleaning process is sold as ** extra-cleaned” seed. 

Poa compressa (Canada bluegrass).—The seed of Canada bluegrass 
is mostly produced in the Province of Ontario, along the north shore 
of the eastern half of Lake Erie. The soil is a heavy clay on lime- 
stone. In this section Canada bluegrass is not sown, but appears as a 
volunteer in any fields that are not kept under cultivation, making a 
thick growth and crowding out other grasses and weeds. It is nearly 
always found in wheat fields when the wheat crop is a partial failure. 
In this case the seed, ripening as it does at the same time as the 
wheat, is thrashed with it and screened out in cleaning. Where the 
seed is harvested alone the grass is cut with a mowing machine and 
cured the same as ordinary hay, and afterwards thrashed with a 
clover huller or grain separator. The hay is bright green, even when 
not cut until after the seed is ripe, and is well liked by some farmers 
as feed, while it is considered hard and of little value by others. A 
good crop is from 200 to 300 pounds of clean seed per acre. There 
has been some demand for this seed in the Southeastern States under 
the name of Virginia bluegrass. The seed is easily cleaned, as it is 
comparatively free from wool at the base and does not require rub- 
bing, as does Kentucky bluegrass seed. No special machinery is used 
except rather long sieves to insure sufficient screening. 

Poa trivialis (rough-stalked meadow grass).—The wholesale trade 
in the seed of rough-stalked meadow grass is largely confined to the 
city of Hamburg, Germany. ‘The seed is collected in the neighbor- 
hood of that city and in the marshes of the Elbe. Seed of good 
quality is also supplied from Denmark, where in one locality this 
grass is grown especially for seed, and it is said to yield as much as 
400 pounds to the acre. The seed is stripped or the grass is cut and 
the seed allowed to after-ripen, when it is cleaned by hand. 

Poa nemoralis (wood meadow grass).—The seed of wood meadow 
grass is gathered by hand in the woods of Germany, and cleaned in 
the same manner as is the seed of Poa trivialis. 

Poa trifora (fowl meadow grass).—Though widely distributed 
throughout the northern portion of the United States, this species is 
chiefly a natural meadow grass of lowlands, and is usually so mixed 
with sedges and other grasses that seed collection on a commercial 
scale has not thus far been undertaken in this country. The seed of 
this species on the market comes from Europe and is very poor. 


14 THE SEEDS OF THE BLUEGRASSES. 


Prof. L. R. Jones, of the Vermont Agricultural Experiment Station, 
reports the seed production from a nearly pure stand of this grass as 
amounting to 6 bushels of 19 pounds each per acre. A small plat 
yielded seed at the rate of over 7 bushels per acre. The seed is pro- 
duced abundantly and ripens evenly. In Vermont it is harvested in 
the latter part of July. The name fowl meadow grass is often applied 
to another lowland grass, Panicularia nervata. 

Poa arachnifera (Texas bluegrass).—The seed of Texas bluegrass is 
gathered by hand in northern Texas. It is cleaned by rubbing 
between the hands, and, owing to the long, woolly hairs at the base of 
the seed, it is never ‘‘ fancy clean.” The best seed is produced on 
rich, black, waxy soil, and is ripe about May 1 to 15. Only a small 
quantity is gathered each year, and consequently it is high priced and 
can not be considered as a commercial seed at the present time. 

Poa annua (annual bluegrass).—The seed of the annual bluegrass is 
not on the market in this country, though the plant is common about 
dwellings, especially in the South and East, and ripens its seed 
throughout the summer. The seeds do not ripen evenly, the upper 
ones falling before the lower flowers have opened. The seed is 
gathered and used to some extent in Europe. 

Poa alpina (alpine meadow grass).—Alpine meadow grass is best 
known in Switzerland, where the seed ripens from the end of June to 
the middle of July. The viviparous form can be propagated by scat- 
tering the buds during the hot weather. 

Poa sudetica.—The seed of Poa sudetica, which is a European grass, 
is rare in the market, but is occasionally quoted by French and by 
German firms. It is sometimes mixed to some extent with the seeds 
of the meadow grasses, particularly water meadow grass (Pandeularia 
americand). 


In addition to the foregoing, other species of Poa occur in the 
western and northwestern United States, where they contribute to 
the native forage of the stock ranges. The seeds of these species, 
however, are not found in commerce. 


1ieDESCRIPTIONS OF THE SEEDS OF THE COMMER- 
CIAL BLUEGRASSES AND THEIR IMPURITIES. 


By F. H. Hrtiman, 


Assistant Botanist, Seed Laboratory. 


THE BLUEGRASSES. 


The ‘‘seeds” of the species of Poa, or the bluegrasses, are the 
ripened florets or individual parts of the smaller clusters, or spikelets, 
of the general floral system of the plant. The number of florets in 
each spikelet varies from two to nine in the different kinds of Poa 
commonly found in commerce. There is some variation in the num- 
ber of florets in the spikelets of each species. The florets separate 
readily at maturity, and well-cleaned samples of seed contain few 
whole or partial spikelets. 

A complete, mature spikelet embraces, besides its several florets, a 
pair of chaffy scales, termed empty glumes, between which the florets, 
or at least the lower ones, rest. The empty glumes, while somewhat 


Fic. 1.—I.—A spikelet of Poa: a, stem of spikelet; b, empty glumes; c, florets, or “ seeds.”’ II.—Single 
floret, back view: a, callus; b, keel; c, intermediate veins; d, marginal veins; e, hyaline portion of 
glume. III.—Single floret, side view: a, callus; b, rachilla segment; c, keel; d, intermediate vein; 
e, marginal vein; f, margin of glume. IV.—Single floret, front view: a, rachilla segment; b, mar- 
ginal fold; c, palea; d, keelsof palea. V.—Terminal floret, front view: a, rachilla segment; b, aborted 
floret; c, palea. VI.—Caryopsis, or grain: a, location of embryo; b, keeled face; c, grooved face. 


dissimilar, are keeled, acute, and one or three veined. The keel of 
each is usually hispid-ciliate above the middle. A portion of the stem 
of the spikelet often remains attached to the base of the empty glumes 
when these are found in commercial samples. 

Each mature, well-developed floret or seed consists of a caryopsis, 
commonly called grain, two inclosing scales which, together with 
the empty glumes, constitute the chaff, and a slender appendage, 


the rachilla segment. (Fig. 1.) 
15 


16 THE SEEDS OF THE BLUEGRASSES. 


The caryopsis corresponds to an individual grain in wheat, rye, and 
barley, and consists almost entirely of the seed proper, to which is 
added only the thin wall of the seed vessel. This is intimately blended 
with the seed coat, the two forming the covering of the true seed. 
The caryopsis is spindle-shaped and often broadest between the middle 
and the base. It is often bluntly keeled along one face and more or 
less evidently grooved along the opposite face. In the commercial 
bluegrass seeds the grain is amber-colored or dull wine-colored and 
semitranslucent. The surface is finely granular and dull. The kernel 
of the seed forms that part of the grain within the seed and seed- 
vessel walls. It consists of the embryo and endosperm, the latter 
forming the greater part. The embryo is situated at the basal 
extremity of the grain and is evident externally as a small ridge, often 
within a slight depression, on the keeled face. The grain adheres 
along its grooved face to the palea in some species in which free grains 
are not common in well-cleaned commercial seed. 

The two chaffy scales of the floret differ chiefly in size, form, rela- 
tive position, venation, and texture. The larger one, called the flower- 
ing glume or simply the glume, incloses the edges of the other, termed 
palea. The grain rests between the glume and palea, its keeled face 
lying against the glume. The rachilla segment is at the base of the 
palea and opposite the glume. It is one of the articulating sections of 
the rachilla, or axis of the spikelet. 

The characters by which the different kinds of bluegrass seeds are 
distinguished one from another are afforded by the glume, palea, and 
rachilla s¢gment, and involve size, form, color, veins of the glume, 
form and texture of the apex of the glume, and the pubescence. 

The glume is stiffish and more or less pointed at the ends. Its base 
is marked by the presence of a small, somewhat knob-like appendage, 
the callus. The latter bears the scar of attachment of the floret and, 
in certain species, a more or less pronounced tuft of webby hairs. 
The back of the glume is more or less keeled along its longitudinal 
center. Besides the fold forming the keel, the edges of the glume are 
infolded along the marginal veins. ‘The marginal folds often are most 
pronounced within and sometimes are confined to the lower half of the 
glume, in which event the upper margins usually diverge and become 
spreading or flaring at the apex. The keel is strongly arched length- 
wise in some species and in others is nearly straight. Five veins 
traverse the glume longitudinally; one occupies the keel, two are at 
the marginal folds and are termed the marginal veins, while the other 
two are situated midway between the keel and marginal veins and are 
called intermediate or, by some authors, lateral veins. The interme- 
diate veins exhibit considerable variation in distinctness in the differ- 
ent species. The vein occupying the keel extends to the apex. The 
apex and often the upper part of the lateral margins of the glume in 


THE BLUEGRASSES. uty 


most species are thin and translucent, or hyaline. The extent of the 
hyaline portion of the apex has much to do with the form of the latter 
and is variable in the different species. 

The palea is commonly more delicate in texture than the glume, 
being partially hyaline. It usually is shorter than the glume, but in 
some species equals or exceeds it in length. The difference in length 
usually is most evident in the lower florets of the spikelet. Two veins 
traverse the palea lengthwise and nearly meet at its apex. The mar- 
gins of the palea are more or less acutely infolded along these veins, 
which are called the keels of the palea. The keels are mostly covered 
by the glume in some species, while in others they are almost wholly 
exposed. There is some variation in this respect, however, among 
seeds of the same species. The apex of the palea is often notched. 

The rachilla segment is nearly cylindrical and usually somewhat 
curved. It is slightly expanded at the apex, which is obliquely trun- 
cate, its terminal surface constituting the scar of at- 
tachment to the succeeding floret. Different florets in 
the same spikeletin certain species exhibit a marked 
variation in the length of the rachilla segment, which 
is shortest in the lower florets and conspicuously 
longer in the terminal one, where it usually bears an 
_ aborted floret as a small, pointed appendage. 

The surface of the florets of different species of Poa 
is subject to considerable variation. Some florets are 
smooth, or glabrous; others bear numerous minute, 
stiffish hairs, rendering the surface rough, or scabrous; ral Tha a 
and some have a fine, appressed pubescence covering grass seed (Poa 
a part of the surface. Most of the species have a tacts een pea 
more or less silky pubescence on the keel and mar- marginal vein; ¢, 
ginal veins below the middle or somewhat higher on = PUDS*eenee oF 
the keel. The intermediate veins are more rarely 
pubescent. The keels of the palea are usually fringed with minute 
hairs, or are hispid-ciliate, but in some species they are silky pubes- 
cent. The basal web is wholly wanting in some species and in others 
varies from a few fibers to a copious tuft. It readily separates from 
the floret in most species. The rachilla segment is usually smooth, 
but in some species it is appressed pubescent. The presence of the 
hairs on the marginal veins often necessitates that care be used in 
examining the rachilla segment with respect to pubescence. (Fig. 2.) 

The color of mature seeds varies from very light brown to dark 
brown. Sterile seeds are usually lighter or straw colored. Immature 
seeds are more or less tinged with green; some are purplish. In cer- 
tain species the glume is tinged with golden yellow near the apex. 
The aborted terminal floret and all the hairs are white. The rachilla 
segment is lighter colored than the glume or palea. 

5813—No. 84053 


15 THE SEEDS OF THE BLUEGRASSES. 


Poorly cleaned samples are apt to contain many sterile florets. 
These are slender, sometimes shrunken, and usually lighter colored 
than the grain-bearing florets, which are comparatively plump and 
often dark colored, owing to the color of 
the grain appearing through the thin palea. 
(Fig. 3.) 

The recognition of the several species of Poa, 
when the identity is questionable, requires the 
use of a good lens and a knowledge of the 

vs AN principal distinguishing characters. A sample 
oe ates ds of Kentucky Under examination should be spread thinly on 
bluegrass (Poa pratensis): a, a Sheet of paper, or, better still, on a black sur- 
poe Sree face. With a good light and means for turn- 
ing the seeds over while under the lens, they can easily be examined 
with reference to size, color, distinctness of veins, character of pubes- 
cence, the condition of the margins of the glume, etc. Exposing the 
different sides of the florets to the light while under examination is 
often absolutely essential in determining the nature of the veins and 
pubescence. 


KEY TO THE SEEDS OF THE MORE COMMON SPECIES OF POA AS FOUND 
ON HERBARIUM SPECIMENS. 
Basal web present. 
Web very persistent and conspicuous -...--.--.--+-----c-,sssnas P. arachnifera. 
Web easily removed, small; keel of the glume pubescent. 
Intermediate veins distinct. 
Intermediate veins sharply defined as narrow ridges; glume margins narrow, 
not easily seen from the side; marginal veins usually smooth. P. trivialis. 
Intermediate veins usually not sharply defined; glume margins broader, easily 
seen from the side in fertile florets; marginal veins pubescent. P. pratensis. 
Intermediate veins indistinct. 
Rachilla segment smooth or nearly so; florets 2-23 mm. long. 
Florets usually broader above than below the middle; apex usually flaring; 


rachilla'sesment smooth 22222022: Sen. 2- eee eee P. compressa. 
Florets not evidently broader above than below the middle; apex usually 
golden yellow; rachilla segment sometimes rough ........-.-- P. triflora. 
Rachilla segment usually pubescent. 
Florets 25-3 mm. long, usually not yellow at the apex. ..... P. nemoralis. 


3asal web not present. 
Florets strongly pubescent. 
Intermediate veins distinct; palea keels prominent, often arched forward. 
P. annua. 
Intermediate veins’indistinct; palea keels not arched. ...........--- P. alpina. 
Hlorets not pubescent se... .--- 2 6. see. eels eee ee P. sudetica. 


KEY TO COMMERCIAL SEEDS. 19 


KEY TO COMMERCIAL BLUEGRASS SEEDS AFTER PREPARATION FOR 
MARKET. 


Seeds 46 mm. long; web longer than glume, forming a woolly tuft and causing the 
seeds to cling-in bunches in the sample.....-----.---------------- P. arachnifera. 
Seeds 2-27 mm. long, usually rubbed free from hairs and disconnected in the sample, 
often more or less torn at the apex; commonest commercial kinds. 
Intermediate veins distinct; seeds contracted at the apex and not wider above than 
below the middle; hyaline margin of apex seldom present in rubbed seed. 
P. pratensis. 
Intermediate veins very indistinct; seeds broader above than below the middle; hyaline 
margin of apex usually evident and flaring -.......------------- ’ P. compressa, 
Seeds 2-3 mm. long, chiefly unrubbed; disconnected or clinging somewhat in the 
sample; usually not torn at the apex; smooth or the pubescence on the veins 
and the web more or less evident. 
Intermediate veins indistinct. 

Rachilla segment usually pubescent; long, sterile racbilla segments conspicuously 
common, intermediate veins scarcely evident; keeland marginal veins pubescent; 
apex of seed often flaring; seed 23-3 mm. long ................ P. nemoralis, 

Rachilla segment smooth; intermediate veins but slightly evident; keel and marginal 
veins pubescent; apex ofseed sometimes flaring; seed 2-23mm.long. P. triflora. 

Intermediate veins very distinct. 

Rachilla segment smooth and slender; keel pubescent, marginal veins usually 
smooth; apex of seed acute and compressed; seeds often distinctly curved as 
viewed from the side -.-.-.---- Se ta ae reese i ade eae ese P. trivialis. 


THE SEEDS OF THE BLUEGRASSES. 


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OF DISTINGUISHING CHARACTERS. 21 


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22 THE SEEDS OF THE BLUEGRASSES. 


DESCRIPTIONS OF SPECIES. 
Poa pratensis L. 
KENTUCKY BLUEGRASS, JUNE GRASS. 


Spikelets 3-5 flowered; florets 2-2? mm., rarely 3 mm., long, lanceolate or fusiform 
as viewed from the back, lanceolate or ovate-lanceolate as viewed from the side, 
mostly acute or the terminal floret sometimes acuminate at the apex, glabrous 
between the veins, varying from light brown to dark brown, sometimes tinged with 
purple, sterile florets lighter; glume usually sharply keeled quite to the apex and 
often strongly arched, particularly at the base; its marginal folds comparatively 
broad, extending from the base nearly or quite to the apex, becoming hyaline-edged 
above the middle in the lower florets, usually not expanded or flaring at the apex, 
the edges nearly meeting in sterile florets, separated and usually distended forward 
in fertile lower florets, often scarcely covering the palea keels of fertile terminal 
florets, the hyaline edge more or less torn away and the margins jagged at the apex 
in rubbed commercial seed; intermediate veins distinct and glabrous; keel and mar- 
ginal veins silky pubescent below the middle or somewhat higher on the keel; basal 
web well developed; pubescence and web wanting, except occasional traces of the 
former, in well-rubbed commercial seed; palea nearly or quite as long as the glume, 
its keels finely hispid-ciliate and usually covered for the greater part oi their length 
by the margins of the glume; rachilla segment slender, glabrous, varying from about 
one-sixth of the length of the glume in the lower florets to one-half its length in the 
terminal one; aborted floret of the sterile rachilla segment minute; grain 13 mm. 
long, somewhat keeled and grooved, often broadest below the middle, reddish brown » 
or darker about the embryo, and semitranslucent. (Fig. 4.) 


a 


Sepa Pe 
etl 


2 


- Coe 


b 


Z 


Fig. 4.—Different forms of commercial seeds of Kentucky bluegrass (Poa pratensis): a and b, back 
views; c-f, side views; g-j, front views; j, a terminal floret. 


Commercial Kentucky bluegrass seed is mostly free from the silky 
and webby hairs present in hand-gathered samples, owing to the rubbing 
process to which it is subjected before being marketed. The severe 
rubbing results in more or less injury to the thin margins of the 
glume, particularly at the apex, which is usually found to be more or 
less torn when examined with a lens. Seeds of a well-rubbed sample 
do not tend to cling in small bunches as do those which are unrubbed 
or hand-gathered. Well-developed seeds are rather robust and have 
the glume margins well separated and evidently distended forward. 
Sterile seeds, or such as have the grain wanting or poorly developed, 
are generally lighter colored, slenderer, and more compressed, while 
the glume margins more nearly meet and are but slightly or searcely 
distended. Such are much lighter in weight than well-developed 
seeds and consequently are mostly blown out with other chaff in well- 
cleaned seed, . 


DESCRIPTIONS OF SPECIES. a5 


Kentucky bluegrass seed is most readily confounded with that of 
Canada bluegrass (Loa compressa) and rough-stalked meadow grass 
(Poa trivialis). Owing to the difference in cost, Pow compressa is 
sometimes mixed with or substituted for Kentucky bluegrass, while 
the latter is sometimes similarly employed with respect to Poa 


trivialis. 


The characteristic differences between Kentucky bluegrass seed and 
that of Canada bluegrass, as exhibited by the bulk samples and by 
individual seeds under the lens, may be compared as follows: 


KENTUCKY BLUEGRASS (Poa pratensis). 


The usual, well-cleaned bulk samples are 
brown in color. 

Individual, well-matured seeds exhibit 
the same brown color of the bulk sam- 


ple. 


Nearly all the seeds taper from the cen- 
ter to both ends and are not broader at 
the apex than at the base. 


The apex of commercial seeds is usually 
torn, obtusely pointed, keeled, and 
scarcely hyaline. 


The intermediate veins are almost in- 


CANADA BLUEGRASS (Poa compressa). 


Ayerage samples lighter colored than 
those of Kentucky bluegrass. 

The lighter color of individual seeds af- 
fords the principal character for the 
preliminary recognition of these seeds 
in mixtures. 

Most of the seeds are broader at the apex 
thanat the base, often distinctly broader 
at the apex than at the middle. 


Apex of commercial seeds often torn, 
mostly expanded or flaring, often but 
slightly keeled. 

The intermediate veins are very indis- 


variably distinct. tinct or apparently wanting. 


A number of the samples of Kentucky bluegrass seed examined con- 
tained seed of the Canada bluegrass. As the latter seed found in com- 
merce usually contains the prickles or even the seeds of Canada thistle 
( Carduus arvensis), these are often found in samples of Kentucky blue- 
grass seed containing the Canada bluegrass seed. Their presence indi- 
cates the admixture, since the Canada thistle does not grow in the 
seed-producing localities of Kentucky, while it is abundant in Canada, 
where the Canada bluegrass is produced. Samples of pure Kentucky 
bluegrass seed are apt to contain the prickles of horse nettle (Solanum 
carolinense), sometimes wrongly called bull thistle, a prickly plant 
common in the bluegrass region of Kentucky. These prickles are 
similar to those of the Canada thistle, but may be distinguished, as 
shown hereafter in this paper in describing the impurities of the blue- 
grass seeds. The fact that Canada bluegrass only begins to flower at 
the time Kentucky bluegrass is ripe precludes the possibility of the 
mixture of the two kinds of seed owing to the fact of growth together. 
Such mixture can occur only after the seed is gathered, through acci- 
dent or intent. 


24 THE SEEDS OF THE BLUEGRASSES. 
Poa compressa L. 


CANADA BLUEGRASS, FLAT-STEMMED BLUEGRASS. 


Spikelets 3-9 flowered; florets 2-23 mm. long, oblong-obovate or the terminal one 
lanceolate as viewed from the back, somewhat narrowly oblong as viewed from the 
side, obtuse or the terminal one acute, smooth between the veins, straw colored or 
light brown; glume somewhat arched, especially at the base, and strongly keeled at 
the back, the keel often less pronounced at the apex than at the base; margins. 
infolded from the base for about three-fourths the length of the floret in the lower 
florets and nearly to the apex in the upper ones, hyaline-edged above the middle, 
often broadly so at the apex, which is more or less flaring in the lower florets, the 
thin apex often torn and jagged in commercial seed; intermediate veins very indis- 
tinct or not evident, glabrous; keel and marginal veins silky pubescent below the mid- 
dle; basal web present, slight; palea nearly or quite equaling the glume, finely hispid- 
ciliate on the keels, which are usually more orless exposed above the middle, sometimes 
from the base; rachilla segment glabrous, varying from about one-fifth the length 
of the glume in the 
lower florets to one- 
half its length in the 
terminal one; aborted 
floret of the sterile 
rachilla segment min- 
‘ ute; grain 1-13 mm. 

long, keeled and 
slightly grooved, 
’ semitranslucent. 

(Fig. 5.) 

The seed of Canada bluegrass is the cheapest of the bluegrass seeds, 
and is therefore not adulterated with other Poas, although it is itself 
used as an adulterant to a considerable extent. 

Pure samples of Canada bluegrass seed almost always contain the 
prickles and sometimes the seeds of Canada thistle (Cardwus arvensis); 
therefore, the occurrence of these prickles with other kinds indicates 
the use of this speciesas an adulterant. Their occurrence with seed 
of Poa trivialis without evidence of the presence of Canada bluegrass 
seed is noted under the discussion of P. trivialis. 


@ 


Fic. 5.—Commercial seeds of Canada bluegrass (Poa compressa): a and b 
back views; c-e, side views; f-i, front views of florets; 7,a terminal floret. 


Poa trivialis L. 
ROUGH-STALKED MEADOW GRASS. 


Spikelets 2 or 3 flowered; florets 2-23 mm., rarely 3 mm., long, narrowly lanceolate 
or the fertile terminal one ovate-lanceolate as viewed from the back, usually lanceolate 
and curved as viewed from the sidé, laterally compressed as compared with other spe- 
cies, straw colored or light brown and sometimes purplish, sharply keeled, the keel 
somewhat arched; margins of the glume scarcely or but slightly distended, narrowly 
and rather sharply infolded nearly or quite to the apex, which is hyaline-edged, very 
acute and rarely expanded; intermediate veins very distinct as narrow and sharply 
defined ridges; keel slightly pubescent below the middle, or rarely smooth; marginal 
veins smooth or sometimes pubescent, basal web present; palea nearly equal to the 
glume, its keelssmooth or finely hispid-ciliate near the apex and mostly covered by 
the margins of the glume except in the larger terminal florets; rachilla segment very 
slender, glabrous, varying from one-fourth to one-half the length of the glume; 
grain 1-14 mm. long, keeled and grooved, semitranslucent, reddish brown. (Fig. 6.) 


DESCRIPTIONS OF SPECIES. 95 


Rough-stalked meadow grass is chiefly hand gathered; consequently 
the commercial seed is apt to bear more or less of the web as well as 
the silky pubescence on the keel. In many samples, however, both 
are rubbed away. 

This seed resem- 
bles that of Poa 
pratensis and that 
of Poa compressa 


so closely that both 
are employed as 
adulterants, th € Fig. 6.—Seeds of rough-stalked meadow grass (Poa trivialis): a and b, 


former apparently back views; c-e, side views; f and g, front views; g, a terminal floret. 


to considerable extent, since it has frequently been found to constitute 
a considerable part of samples of so-called rough-stalked meadow 
grass. One sample examined marked ‘‘/oa trivialis” from Europe 
consisted almost wholly of oa compressa. Several samples from 
Europe contained prickles of Canada thistle, but no seeds of Canada 


bluegrass were found. 


The principal distinguishing characters of the three species may be 


compared as follows: 


ROUGH-STALKED MEADOW GRASS 
(Poa trivialis). 


Commercial seeds are usu- 
ally pubescent on the 
keel vein, usually smooth 
on the marginal veins 
and bear more or less of 
the webby hairs, conse- 
quently cling together in 
masses. 

As viewed from the side, 
the seeds are somewhat 
curved, much narrower 
than the others, the 
glume margins usually 
only slightly evident. 


Apex of the glume usually 
uninjured, strongly 
keeled, acute, slightly 
hyaline-margined, often 
curved. 


Intermediate veins sharply 
defined as narrow ridges. 


Rachilla segment very slen- 
der and less variable in 
length than in P. pra- 
tensis or P. compressa. 


KENTUCKY BLUEGRASS 
(Poa pratensis). 


CANADA BLUEGRASS 
(Poa compressa). 


Commercial seeds rarely pubescent on the veins and 
the webby hairs wanting; consequently mobile in bulk, 
not clinging in masses; unrubbed seed pubescent on 
the marginal and keel veins. 


Seeds mostly straight as 
viewed from the side, 
glume margins’ cften 
strongly distended. 


Apex of the glume more or 
less torn in commercial 
seed; keeled, sharply 
pointed, hyaline-edged 
and not curved in un- 
rubbed seed, 


Intermediate veins dis- 
tinct as rather coarse 
ridges. 


Seeds straight, the glume 
margins somewhat evi- 
dent from the side. 


Apex of the glume often 
torn, otherwise some- 
what keeled, obtusely 
pointed, broader than the 
base, hyaline-edged. 


Intermediate veins indis- 
tinct or apparently want- 
ing. 


Rachilla segment coarser than in P. trivialis and often 


very short. 


26 THE SEEDS OF THE BLUEGRASSES. 


Poa nemoralis L. 
WOOD MEADOW GRASS. 


Spikelets 2 or 3 flowered; florets 23-3 mm. long, lanceolate or ovate-lanceolate, 
mostly acute at the apex, light brown, sometimes yellowish tinged near the apex; 
glume rather broadly keeled and somewhat arched at the back; margins of the 
glume narrowly infolded quite to the apex or hyaline-edged and often flaring above 
the middle; intermediate veins very indistinct; keel and marginal veins silky 
pubescent below the middle; basal web slight; surface between the veins glabrous; 
palea nearly equal to the glume, evidently shorter in florets having a flaring apex, 
its keels hispid-ciliate and usually covered by the margins of the glume; rachilla 
segment varying from one-fourth to three-fourths of the length of the glume, the 
sterile rachilla segment very uniformly much longer than the others, more or less 
appressed pubescent, the pubescence somewhat variable and sometimes nearly want- 
ing; aborted floret of the sterile rachilla segment often one-half as long as the seg- 
ment; grain 14 mm. long, rather slender, semitranslucent. (Fig. 7.) 


Fic. 7.—Seeds of wood meadow grass (Poa nemoralis): a-c, back views; d and e, side 
views; j-j, front views; j, a terminal floret. 


Commercial wood meadow grass seed is not rubbed in preparation 
for market, and therefore possesses much of its rather persistent and 
prominent silky pubescence, and the thin tips of the florets are mostly 
uninjured. The pubescence of the rachilla segment is persistent and 
present in most of the seeds of all pure samples of this species. It 
affords the most marked characteristic by which the seeds of P. nemo- 
ralis may be distinguished from those of other commercial species of 
Poa. The conspicuously longer rachilla segments of the terminal 
florets are noticeably abundant in samples of this species, since these 
florets constitute from one-third to one-half of all the seed. The 
abundance of the long rachilla segments is helpful in distinguishing 
these seeds from those of other Poas. 

Commercial seed of 2. nemoralis is apt to be very much adulterated 
with other species of Poa. Of a number of samples examined less 
than half were true to name. One was nearly pure Canada bluegrass 
seed, and the rest consisted in part of one or all of the following 
species: P. pratensis, P. compressa, and P. trivialis. 

The following comparison of characters should render it compara- 
tively easy to distinguish the seeds of 7. nemoral/s from those of the 
other species. 


DESCRIPTIONS OF 


Woop MEADOW GRASS 
(Poa nemoralis). 


Silky pubescence of the 
veins mostly present and 
prominent. 

Apex of the glume slen- 
derly pointed or nar- 
rowly flaring. 

Intermediate veins indis- 
tinct. 

Rachilla segment pubes- 
cent, often more than 


SPECIES. 27 


CANADA BLUEGRASS 
(Poa compressa) . 


KENTUCKY BLUEGRASS (Pod 
pratensis); ROUGH-STALKED 
MEADOW GRASS (Poa trivi- 
alis). 


Silky pubescence of the veins wanting or but slight. 


Apex of the glume acute. Apex of the glume broadly 


flaring. 
Intermediate veins dis- Intermediate veins indis- 
tinct. tinct. 


Rachilla segment smooth, not exceeding half the length 
of the glume. 


half the length of the 
glume. 


Poa triflora Ehrh. (P. flava L., P. serotina Ehrh.). 
FOWL MEADOW GRASS, FALSE REDTOP. 


Spikelets 2-4 flowered; florets 2-23 mm.long, lanceolate or ovate-lanceolate as 
viewed from the back, broadly keeled and strongly arched at the back, light brown 
and usually strongly tinged with yellow above the middle, sometimes purplish, 
margins of the glume narrowly infolded below the middle or quite to the apex, 
which is hyaline-edged, expanded but scarcely flaring; intermediate veins indis- 
tinct; keel and marginal 
veins silky pubescent below 
the middle; basal web 
slight; palea nearly or quite 
equal to the glume, finely 
hispid-ciliate on the keels, 
which are mainly covered 
by the glume margins in the 
lower florets; rachilla seg- 
ment slender, glabrous or 
sometimes slightly scabrous, from one-fourth to one-half or two-thirds the length of 
the glume; aborted floret of the sterile rachilla segment often prominent and nearly 
as long as the rachilla segment; grain 1 mm. long, comparatively robust and smooth, 
scarcely keeled or grooved, semitranslucent. (Fig. 8.) 


Fic. 8.—Seeds of fowl meadow grass (Poa triflora): a-c, back 
views; d and e, side views; j-h, front views; h, a terminal floret. 


Most, if not all, of the seed of P. triflora on the market appears to 
be of foreign production. The samples examined have proved to be 
the worst found among the bluegrasses. It is probable that a better 
grade of seed could be secured from the natural meadows in this coun- 
try where this species often constitutes the principal grass. The 
seeds of P. triflora are very similar to those of Canada bluegrass and 
wood meadow grass. 


28 THE 


SEEDS 


OF THE BLUEGRASSES. 


The prineipal distinguishing characters of the three kinds are as 


follows: 


FOWL MEADOW GRASS 
(Poa triflora). 


Seeds 2-23 mm. long. 

Seeds mostly narrower at 
the apex than at the cen- 
ter. 

Seeds usually yellowish at 
the apex. 

Intermediate veins usually 
evident but indistinct. 


Pubescence of the veins 
and the web often pres- 
ent in commercial seed. 


Rachilla segment mostly 
smooth,sometimesslight- 
ly rough, often  two- 
thirds the length of the 
glume. 


CANADA BLUEGRASS 


( Poa compressa). 


Seeds 2-23 mm. long. 

Seeds mostly broader at the 
apex than at the center 
or base, 

Seeds not yellowish at the 
apex. 

Intermediate veins indis- 
tinct or more commonly 
not evident. 

Pubescence of the veins and 
the web mostly absent in 
commercial seed. 

Rachilla segment smooth, 
not exceeding one-half of 
the length of the glume. 


WooD MEADOW GRASS 


(Poa nemoralis). 


Seeds 23-3 mm. long. 

Seeds narrower or not 
broader at the apex than 
at the center. 

Seeds sometimes yellowish 
at the apex. 


Intermediate veins indis- 
tinct. 


Pubescence of the veins 
usually present in com- 
mercial seed. 

Rachilla segment pubes- 
cent or sometimes only 
rough, oftenthree-fourths 
the length of the glume. 


The name fowl meadow grass is often applied, both by seedsmen 
and by writers upon grasses, to Panicularia nervata. 


Poa arachnifera Torr. 


TEXAS BLUEGRASS. 


Spikelets 4 or 5 flowered; florets 4-6 mm. long, narrowly lanceolate, acuminate, 


straw colored or light brown; glume strongly keeled quite to the apex and somewhat 
arched; margins narrowly infolded below and becoming broadly hyaline above the 
middle, not widely flaring at the apex; marginal and keel veins strongly pubes- 


C 


Fig. 9.—Seeds of Texas bluegrass (Poa arachnifera): a and b, back views, seeds showing the long 
hairs of the web; cand d, side views; e-g, front views; g, a terminal floret. 


cent with long, silky hairs; basal web copious, often twice as long as the floret, 
very persistent; surface between the veins glabrous, the keel hispid-ciliate above the 
middle; palea from three-fourths to four-fifths the length of the glume, its keels 
more or less exposed, silky pubescent to the middle and hispid-ciliate at the apex; 
* rachilla segment varying from about one-sixth to one-third the length of the glume, 
glabrous; aborted floret of the sterile rachilla segment minute; grain slender, 14-3 
mm. long, oblong-fusiform, nearly opaque, distinctly grooved and keeled. (Fig. 9. ) 


DESCRIPTIONS OF SPECIES. 29 


Texas bluegrass seed in commerce is unrubbed, and as the silky 
pubescence and web are very persistent they are always present. The 
hairs are so long and copious that the seeds cling in loosely matted, 
woolly bunches, and thus are easily distinguished 
from all the other commercial Poas. (Fig. 10.) 


Poa annua L. 


ANNUAL MEADOW GRASS. 


Spikelets 3-5 flowered; florets 13-3 mm. long, ovate or 
ovate-lanceolate and relatively robust, strongly keeled and 
arched at the back, more or less densely pubescent, light 
brown or dark brown and often purplish or yellowish; 
margins of the glume very narrowly infolded below the 
middle, thin and broadly hyaline above the middle inthe Fig. 10.—A cluster of Texas 
lower florets, flaring, gaping, or infolded at the apex; inter- bluegrass seeds matted by 
mediate veins usually distinct as narrow ridges extending “© webby fibers. 
from the base to the margin of the apex, glabrous or pubescent; marginal veins 
and keel densely soft-pubescent below the middle; surface between the veins some- 
times more or less pubescent at the base; web wanting; palea somewhat shorter than 
the glume, except in the terminal floret; keels of the palea coarse and prominent, 
mostly exposed, usually arched forward and exposed to side view in florets having a 
well-developed grain, often contracted toward the rachilla segment at the base, silky 
pubescent from near the base nearly to the apex; rachilla segment glabrous, from 
one-fourth to one-third the length of the glume, aborted floret of the sterile rachilla 
segment minute; grain 1-13 mm. long, robust, distinctly granular, keeled and grooved, 
slightly translucent. (Fig. 11.) 


The seed of Poa annua is not in the trade and is not apt to become 
mixed with the commercial bluegrass seeds. It may be readily distin- 
guished from the common commercial species of Poa by its abundant 


Fie. 11.—Seeds of annual meadow grass (Poa annua): a and b, back views; c-e, side views; j-i, front 
views; i, a terminal floret. 

pubescence, arched and silky pubescent keels of the palea, and robust 
form. The seed most closely resembles that of Poa alpina, from which 
it is distinguished in individual seeds by its distinct intermediate veins 
and prominent, arched, and silky pubescent but not hispid-ciliate palea 
keels. 

Poa alpina L. 


ALPINE MEADOW GRASS. 


Spikelets 3-6 flowered; florets 23-33 mm. long, ovate-lanceolate or obovate, the 
uppermost lanceolate, broadly keeled, arched, acute, or obtuse, light brown, some- 
times purplish, and often yellowish tinged at the apex; margins narrowly infolded 
below the middle and becoming broadly hyaline at the apex; intermediate veins 


30 THE SEEDS OF THE BLUEGRASSES. 


indistinct or evident only below the middle; keel and marginal veins silky pubescent 
below the middle or higher on the keel, which is hispid at the apex; surface between 
the marginal veins and keel appressed pubescent at the base; web wanting; palea 
nearly or quite equal to the glume, its keels not arched as in Poa annua, slightly 
silky pubescent below the middle and hispid-ciliate above; rachilla segment glabrous, 
varying from no longer than wide to one-third the length of the glume; aborted 
floret of the sterile rachilla segment minute; grain 14 mm. long, keeled and grooved, 
semitranslucent, dark reddish brown, granular. (Fig. 12.) 


Fic. 12.—Seeds of alpine meadow grass (Poa alpina): a and b, back views; c-e, side views; j-h, front 
views; h, a terminal floret. 


The seed of Poa alpina is not on the market and is not likely to be 
found in commercial seeds. Individual seeds of P. alpina closely 
resemble those of P. annua, but are to be distinguished by the indis- 
tinct intermediate veins of the glume, the variable rachilla segment, 
and especially by the keels of the palea, which are slenderer, not 
arched, less pubescent, and strictly hispid-ciliate above. The plant is 
alpine and occurs in the northern part of the United States as far 
west as Colorado, in Canada and Alaska, and in Europe and Asia. 


Poa sudetica Haenke. 


Spikelets 2 or 3 flowered; florets 3-4 mm. long, lanceelate or ovate-lanceolate; 
apex acute or acuminate; glume somewhat arched and strongly keeled at the back, 
light brown or dark brown, sometimes tinged with purple; margins of the glume 
narrowly infolded below the middle, narrowly hyaline-edged above the middle and 
not flaring at the apex; all the veins distinct, never silky pubescent, usually hispid; 


Fig. 13.—Seeds of Poa sudetica; a and b, back views; c-e, side views; f and g, front views; g,a terminal 
floret. 


general surface scabrous or sometimes glabrous; web not present; palea equaling or 
somewhat exceeding the glume and often separated from it at the apex in florets 
having a well-developed grain; keels of the palea hispid-ciliate, mostly exposed and 
more or less evident from the side; rachilla segment varying from one-fifth to one- 
third or even one-half the length of the glume, glabrous or scabrous, sterile rachilla 


DESCRIPTIONS OF SPECIES. 31 


segment tapering to the apex, the aborted floret usually minute, but sometimes con- 
spicuous and nearly equal to the rachilla segment; grain about 2 mm. long, robust, 
light brown, slightly keeled and grooved, semitranslucent. (Fig. 13.) 


This is a European species not found in the American market. 
Panicularia spp. 


Owing to the fact that Panicularia nervata is sometimes sold as fowl 
meadow grass, a description of its seed is presented. A description 
of the closely allied P. americana, which is often associated with 
P. nervata, is added as an aid in comparing the two species. 


Panicularia nervata (Willd.) Kuntze. 
NERVED MANNA GRASS, SOMETIMES CALLED FOWL MEADOW GRASS. 


Florets 1-14 mm. long, robust, ovate (obovate with reference to the plant), light 
brown, purplish and sometimes greenish when immature; glume rounded at the 
back, prominently seven-veined, its margins 
somewhat infolded at the base and not flaring at 
the apex, which is sometimes narrowly hyaline; 
surface smooth, except the veins, which are 
sometimes scabrous; palea equal to or sometimes 
longer than the glume, broad, the keels exposed, 
prominent and nearly meeting at the rounded 
and sometimes slightly notched apex, usually F!s. 14—Seeds of nerved manna grass 
scabrous above the middle; rachilla segment one- ve ‘ereteark saphaes oat Pee ea 
fifth to one-fourth the length of the glume, sub- ah 5 
cylindrical and scarcely expanded at the apex, the terminal one somewhat longer 
than the others and tipped by a minute, aborted floret; grain loosely held by the 
stiffish glume and palea, obovate, slightly flattened, {-1 mm. long, smooth, some- 
what polished, very dark brown or black, sometimes slightly translucent. (Fig. 14.) 


Panicularia americana (Torr. ) MacM. 
REED MEADOW GRASS, WATER MEADOW GRASS, TALL MANNA GRASS. 


Florets 3-3} mm. long, elliptical-oblong as viewed from the front or back, some- 
what spindle-shaped as viewed from the side, obtuse at the apex, brown, or purplish 
before complete maturi- 
ty; glume rounded at the 
back, distinctly seven- 
veined, its margins nar- 
rowly infolded at the 
base and not flaring at 
the apex; surface smooth 
between the scabrous 
veins; palea equal to the 
glume, concave, its keels 
exposed, nearly meeting 
Fic. 15.—Seeds of water meadow grass (Panicularia americana): a,b, at the apex, very finely 

and c, back, side, and front views of seeds: d, grain. 


: hispid-ciliate; rachilla 
segment one-fifth to one-fourth the length of the glume, subeylindrical, somewhat 
expanded at the apex, that of the terminal floret slightly longer and tipped by a 
minute, aborted floret; grain broadly oblong, 13-2 mm. long, somewhat flattened, 
very dark brown, slightly translucent, smooth, and somewhat polished when fully 
developed. (Fig. 15.) 


32 THE SEEDS OF THE BLUEGRASSES. 


WEED SEEDS COMMONLY FOUND WITH COMMERCIAL BLUE- 
GRASS SEEDS. 


The following weed seeds are those most frequently found with the 
various kinds of bluegrass seed. 


Bursa bursa-pastoris (L.) Britton. 
SHEPHERD’ S-PURSE. 


Seeds 3-1 mm. long, oval-oblong, one extremity often pointed by the whitish tissue 
of the scar, flattened with rounded edges; faces similar and usually presenting two shal- 
low grooves; color yellowish orreddish brown, 
usually darker near the scar; surface nearly 
smooth; endosperm absent; embryo curved 
upon itself, the cotyledons incumbent; seeds 
Cc developing a coat of transparent mucilage 
when placed in water. (Fig. 16.) 

Seldom found abundantly, but occurring 
frequently in all of the commercial bluegrass 


Fic. 16.—Seeds of shepherd’s-purse (Bursa 
bursa-pastoris): a, side view; b, edge view; 
c, natural size of seeds. seeds. 


Lepidium virginicum L. 
PEPPERGRASS. 


Seeds 14 mm. long, much flattened, ovate with one edge straight and thicker than 
the other, the curved edge narrowly margined, the mar- 


gin usually hyaline and broadest at the broad end of the “eo 
seed; faces similar, each nearly crossed lengthwise by a yay 
curved groove; scar at the small extremity, marked by Chr 

a small, whitish tissue; surface smooth, dull, and red- ec wf 


dish yellow; endosperm wanting; embryo curved upon the) ieee 
itself, the cotyledons accumbent; seeds developing a Lepidus vleeiadenan tne tie 
copious coat of transparent mucilage when placed in view; b, edge view; c, natural 
water. (Fig. 17.) size of seeds. 
Frequently found in home-grown seed and sometimes very abundant, especially in 
poorly cleaned seed. 
Cerastium vulgatum L. 


MOUSE-EAR CHICKWEED. 


Seeds about 4 mm. long, flattened but not thin, rounded or triangular, the broad 

edge rounded, the narrow edge notched; surface roughened by small tubercles 

or very short ridges, dull, and reddish- 

o, « brown; embryo cylindrical, curved about 

oo» the endosperm, its extremities nearly 

5 meeting at the notch in the seed coat. 
(Fig. 18.) 

ic. 18. Seeds of motile-cartibickweed (Ce. Found frequently; sometimes abundant in 


rastium vulgatum): a, side views; b, natural poorly cleaned seed. 
size of seeds. 


Alsine media L. 
COMMON CHICKWEED. 


Seeds circular-ovate, about 1 mm. in diameter with little variation in size, flattened 
with plane faces and flattened edges; scar in a small notch in the edge; surface dull, 
slightly tubercled, the tubercles in rows on the edges and in more or less evidently 


WEED SEEDS FOUND WITH BLUEGRASS SEEDS. 33 


concentric rows on the similar faces; color brown, or reddish in immature seeds: 
embryo cylindrical, curved about the endosperm, its extremities nearly meeting at 
the sear. (Fig. 19, a.) 

Alsine media is very common in the United States, but is so low-growing that the 
American method of seed stripping prevents the occurrence of its seeds in abundance 
in commercial bluegrass seeds. Its seeds 
are common in European bluegrass seeds, 


e e 
particularly those of rough-stalked meadow e 
grass. ; °° e 
> 
Alsine graminea (L.) Britton. & 
2 Fie. 19.—Seeds of chickweeds: a, Alsine me- 
Seeds similar to those of Alsine media, ex- _—_ dia; b, A. graminea; c, natural size of seeds. 


cept in form and surface markings; usually 
circular or oval; faces and edges somewhat rounded, finely roughened by short, inter- 
lacing ridges which are arranged more or less concentrically on the faces and 
parallel on the edges; surface dull; color grayish-brown, immature seeds reddish. 
(Fig. 19, 5.) Z 

Not found in American seed; frequent, although not abundant, in European seed. 


Carduus arvensis (L.) Robs. 


> 
CANADA THISTLE. 
19 
2 Seeds (akenes) 2-3 mm. long, 


oblong-lanceolate, flattened with 
obtuse edges, slightly ridged along 
each face, straight or curved edge- 
wise, sometimes facewise; apex 
truncate, often obliquely so, con- 
cave with a ring-like border; corolla scar represented by a central, conical projection; 
surface dull and mostly smooth, sometimes with several narrow, longitudinal grooves; 
color brown, the apical margin usually lighter 
and sometimes yellowish. (Fig. 20.) 

Prickles of Canada thistle and horse nettle 
(Solanum carolinense) often occur in certain 
bluegrass seeds. While the presence of the 
former is significant with respect to adultera- 
tion, the two kinds are apt to be confounded. 

The prickles of Canada thistle are 2-6 mm. 
long, very slender, yellowish, usually expanded 
and laterally flattened at the base, which con- 
sists of a portion of the leaf tissue and is darker 
colored than the rest of the prickle, somewhat 
* rounded or angular in form and jagged-edged. 
(Fig. 21, ce and d), 

The prickles of horse nettle (Solanum caro- Fic. 21.—Prickles often found with blue- 


Fie. 20.—Seeds of Canada thistle ( Cardwus arvensis): a, well- 
matured seeds; b, natural size of seeds; c,a shriveled seed. 


linense) are coarser, 4-8 mm. in length, light grass seed: a and 6, horse nettle (Sola- 
yellow in color, usually not darker at the base ee 

i ’ ’ : size; c and d, Canada thistle (Carduus 
They are produced on the stems and the coarse arvensis) enlarged and natural size; 1 and 
midribs of the leaves, and on breaking off have 2, characteristic forms of the bases of 
a transversely flattened scar. They occur fre- "Be two kinds of prickles. 


quently in samples of Kentucky-grown Poa pratensis and are easily mistaken for 
those of Canada thistle. (Fig. 21, a and b.) 

Matured seeds, shriveled seeds, and prickles from the leaves and stems of Canada 
thistle are frequently found in Canada bluegrass seed. The presence of the prickles 


34 THE SEEDS OF THE BLUEGRASSES. 


in the more expensive kinds of bluegrass seed indicates the probable use of Canada 
bluegrass seed as an adulterant. These prickles have been found, however, in rough- 
stalked meadow grass seed in which no trace of Canada bluegrass seed appeared. 
Owing to the troublesome nature of Canada thistle. care should be taken not to 
introduce its seeds with those of the bluegrasses. 


Taraxacum taraxacum (L.) Karst. 
DANDELION. 


Seeds (akenes) 3-4 mm. long, including the per- 
sistent base of the beak, which forms the pointed 
apex of the seed, lance-shaped or broadly so, 
straight or curved, flattened or slightly four- 
angled with similar faces, barbed in the upper, 
Fic. 22.—Seeds of dandelion (Taraza- broader half; teeth directed toward the apex, 

cum taraxacum): a, side views; b, nat- prominent on the edges and arranged in about 

unal size of needs. five rows on each face, which has two slender 
grooves with three rows of teeth between them; surface dull; color light brown or 
dark brown. (Fig. 22.) 

Occurring occasionally in both American and European seed, these seeds have 
appeared most frequently in Kentucky bluegrass and rough-stalked meadow grass 
seeds. 


Matricaria inodora L. 
SCENTLESS CAMOMILE. 


Seeds (akenes) 14-2 mm. long, slender or 
robust, oblong with obtuse extremities, taper- 
ing somewhat from the truncate apex to the 
base, slightly flattened; faces dissimilar, one 
having three prominent, longitudinal ribs 7 tris eo dia 
joined at the apex, the lateral ribs and a partial elie sie <i troaee WE AS 
one joined to them at the apex appearing on 
the opposite face, which also presents two small cavities separated by the partial 
ridge; surface between the ridges transversely roughened, dark brown or black and 
darker than the brown or yellowish ridges. (Fig. 23.) 

Found only in foreign-grown seed, chiefly in rough- 
stalked meadow grass and wood meadow grass seeds. 


Hieracium sp. 


ee HAWKWEED. 
4, fina Seeds (akenes) 1-3 mm. long, cylindrical, pointed at 
b the base; apex truncate, bearing a small tuft of short, 
whitish, marginal bristles (the remnants of the pappus 
Fic. 24.—Seeds of hawkweed bristles); surface lightly ten-ridged lengthwise; color 
(Hieraciumsp.): a, side views; brown or black, reddish in immature seeds. (Fig. 24.) 
serene pias oh rege Found most frequently in wood meadow grass seed. 
The seeds of several species of hawkweed, occurring in both America and Europe, 
are practically indistinguishable. Specific determinations can not be made by exam- 
ination with a lens. The troublesome character of orange hawkweed (Hieracium 
aurantiacum), whose seeds are 1}-1} mm. long, justifies care in the use of seed 
containing seeds of any species of hawkweed. 


WEED SEEDS FOUND WITH BLUEGRASS SEEDS. 35 


Anthemis cotula L. 
DOG FENNEL, MAYWEED. 


Seeds (akenes) cylindrical, broadly club-shaped, 13-2 mm. long, straight or 
curved; surface dull and usually roughened by 
many small tubercles more or less distinctly 
arranged in ten rows, indistinctly few-tubercled 
or nearly smooth, but commonly more or less 
evidently ten-ribbed; base tipped by the rounded, 
whitish scar; apex rounded or slightly pointed; a 
color varying from light to dark brown. (Fig. 25.) — yg. 25.—Seeds of dog fennel (Anthemis 

Found occasionally, but never abundantly, in eotula): a, side views; b, natural size 
both American and European bluegrass seed. of seeds. 


Chenopodium album L. 
LAMB’S-QUARTERS, PIGWEED. 


Seeds nearly circular, lens-shaped, with blunt edges, 1-13 mm. in diameter, occur- 
ring in commercial seeds as free seeds or as fruits, the seeds proper being invested by 
the thin pericarp; free seeds jet black, smooth 
or nearly so, and highly polished; scar occu- 
pying a curved groove extending from the cen- 
ter to the edge of one face and usually evident 
as a light-colored line; fruits only slightly 
larger than the seeds, mostly gray or black and 
dull; pericarp wall often broken away, expos- 
ing theshining black surface of theseed; again, 
this wall and the seed coat are often broken, 
exposing the yellowish or whitish embryo and 
endosperm; embryo cylindrical, occupying 
the border of the seed and surrounding the endosperm, its extremities almost meet- 
ing, the tip of the caulicle occupying an extension of the seed coat at the edge beside 
the sear. (Fig. 26.) 

Found chiefly in Kentucky bluegrass and Canada 
bluegrass seeds, but not frequently and never abun- 
dantly. 


Fig. 26.—Seeds of lamb’s-quarters ( Chenopo- 
dium album): a, various forms of seeds; b, 
natural size of seeds. 


iy 


Plantago lanceolata L. 


RIB-GRASS, BUCKHORN, ENGLISH PLANTAIN. 


Seeds oval-oblong, 13-3 mm. long, flattened, one 
face convex, the other having a deep groove and Fie. 27.—Seeds of rib-grass (Plan- 
rounded, infolded edges which scarcely meet at one —*@go /anceolata): a, front and back 
end; surface smooth or slightly uneven, shining in Maa eee aha at 
fresh seed, brown or somewhat amber-colored; scar 
situated at the center of the grooved face; embryo straight, in the center of the 
endosperm, usually evident through the somewhat transparent endosperm and seed 
coat. When placed in water the seeds develop a coat of transparent mucilage. 
(Fig. 27.) 

Small seeds are found to some extent in both American and European seed; more 
commonly in Kentucky bluegrass than in Canada bluegrass seed. 


36 THE SEEDS OF THE BLUEGRASSES. 


Rumex crispus L. 
CURLED DOCK. 


Seeds (akenes) 13-23 mm. long, triangular with equal faces and broadly ovate- 
lanceolate; color dark reddish brown; surface smooth, polished; apex acute; base 
obtuse, contracted, and narrowly truncate at the scar; edges narrowly margined; 

faces longitudinally concave in poorly developed 
seeds; true seed coat thin; embryo cylindrical, rest- 

$ ing in the center of one face of the endosperm; 
® 9 caulicle pointing to the base of the akene. (Fig. 28.) 
b Found occasionally, especially in Kentucky blue- 
grass and in Canada bluegrass seeds; small, imper- 

Fic. 28.—Seeds of curled dock (Ru- fectly developed seed more commonly found than 
mex crispus): a, broad and narrow Jarge, heavy seed. Their sharply three-angled, 

forms; b, natural size of seeds. Pare 2 

beechnut-like form distinguishes them from other 
impurities, except one or two other kinds of dock. The docks are destructive weeds, 
and care should be taken to prevent the sowing of their seeds. 


Rumex acetosella L. 


SHEEP’S SORREL, SORREL. 


Seeds (fruits) acutely oval, three-angled, with equal faces, 1-1} mm. long; repre- 
sented in commercial seed by the seed-like akene only or by the akene covered by 
the thin, closely fitting perianth segments, which are six in number, three broad 
ones covering the sides of the 
akene and three small ones cover- 
ing the angles at the base; covered 
by the perianth, the seeds are 
finely roughened, dull, and red- 


dish brown; venation of the three 
Fic. 29.—Seeds of sorrel (Rumex acetosella): a, b, and e, 


broad segments evident; small seed enveloped by the perianth; d, seed with perianth 
segments at the basal angles often removed; e, natural size of seeds. 


broken away; akenes but slightly 
smaller than when covered by the perianth, bluntly three-angled; surface smooth, 
somewhat polished, reddish brown or wine colored, often semitranslucent; angles 
dark at the apex; internal structure essentially the same as in Rumex crispus. 
(Fig. 29.) 

One of the commonest impurities in commercial 
seed, found in all seed of the cultivated bluegrasses. 


@ ° . * 
* ee Veronica arvensis L. 
° 2 ; 

d . CORN SPEEDWELL. 


Seeds }-} mm. long, flattened and thin, more or 
FIG. 30.—Seeds of corn speedwell less regularly oval, plane or sometimes curved face- 

( Veronica arvensis): aand b, front wise; center of the inner face marked by the relatively 

views, ¢, back view; d, natural large, raised chalaza, which is united by a narrow 

chy hy omnes ridge (the raphe) to the scar on the edge of the 
smaller extremity of the seed; external face slightly ridged longitudinally, indicat- 
ing the position of the embryo, which is surrounded by the endosperm; surface dull, 
finely roughened by somewhat radially-disposed ridges, and reddish yellow. (Fig. 
30. ) 


WEED SEEDS FOUND WITH BLUEGRASS SEEDS. 37 


Found in bluegrass seed of various species, especially common in seed of Kentucky 
bluegrass. The relatively prominent chalaza and the radially uneven surface dis- 
tinguish them from the seed of the closely allied Veronica 
peregrina, which sometimes occurs in commercial seed. 


Juncus tenuis Willd. o> 
a g 


SLENDER RUSH. 


by 
E 
Seeds very minute, about 4} mm. long, broadly spindle- 
shaped, the extremities usually slightly curved; surface Fic. 31.—Seeds of slender rush 
(as seen under a lens) nearly smooth; color reddish = (7™eus fenuis): a, seeds en- 
res y : larged; 6b, natural size of 
yellow, darker at the extremities, which sometimes bear ....4. 
a small white tissue. (Fig. 31.) > 
Often quite abundant in poorly cleaned Kentucky bluegrass seed, sometimes cling- 
ing in bunches of several seeds each. 


Juncoides campestre (L.) Kuntze. 
FIELD RUSH. 


a r) Seeds 14-13 mm. long, oval, not flattened, the ex- 
et -« tremities unequally pointed, the basal extremity turned 
® slightly to one side and consisting of soft white or 
b yellowish tissue; a narrow and often indistinctly de- 
ee ees. oe Geld ruall (Jun: fined whitish ridge extends from the base to the apex; 
coides campestre): a, different body of the seed wine-colored and semitranslucent or 
views; b, natural size of seeds. grayish. (Fig. 32.) 
Found frequently in the seed of wood meadow 
grass and of the Poa sudetica of European origin. 


Juncoides albida DC 


WOOD RUSH. 


Seeds 1-1} mm. long, narrowly oval, not flattened; base without an appendage of 
soft tissue; apex more acutely pointed than the base; 
a distinct brown or reddish brown ridge joins the base 
and apex; body of the seed reddish brown or wine- 
colored, often semitranslucent. (Fig. 33.) 

Found in various species of European-grown blue- 
grass seed. The usually smaller size, absence of the “* 
basal appendage, and more distinct and constant red= BEI Ae 
dish-brown lateral ridge serve to distinguish these — goides athida): a, different views: 
from the seeds of Juncoides campestre. b, natural size of seeds. 


Carex cephalophora Muhl. 


OVAL-HEADED SEDGE. 


Seeds (akenes) 13-2 mm. long, lens-shaped and broadly ovate, contracted at the 
base and tipped at the apex by a conical appendage (the base of the style); surface 
smooth and dull; color varying from light to dark brown; apical appendage often 
broken away in seeds found in commercial samples; perigynium broadly ovate- 
lanceolate, plano-convex, the tapering extremity usually rough-edged ‘and notched at 


38 THE SEEDS OF THE BLUEGRASSES. 


the apex; surface sometimes slightly grooved or ridged lengthwise, otherwise smooth; 
color varying from light brown to greenish or dark brown. (Fig. 34.) 


Fic. 34.—Seeds of sedge ( Carex cephalophora): a,seeds 
inclosed by the perigynium; } and c, seeds with 
perigynium removed; d, natural size of seeds. 


ERGOT OCCASIONALLY FOUND IN 
BLUEGRASS SEED. 


Claviceps purpurea (Fr.) Tul. 


ERGOT. 


This is a fungus growth affecting the grain (caryopsis) of 
many grasses. It is very common in the seed of redtop and 
other species of Agrostis, and occasionally occurs in bluegrass 
seed. The grain of the seed becomes elongated, extending 


Seeds of sedge (Carex) are found in 
both American and European bluegrass 
seed. Owing to the wide area of their 
production, the seeds of various species 
of Carex occur in commercial blue- 
grass seed. The seeds of Carex are 
fruits (akenes) and occur free or in- 
closed within a sae-like covering (the 
perigynium). Carex cephalophora is 
the species most commonly found in 
Kentucky bluegrass seed. 


COMMERCIAL 


Fic. 35.—Ergot (Clavi- 


beyond the glume and palea, attains about twice the length of — ceps purpurea) of Ken- 


the glume, and is club-shaped, straight, or, 


more commonly, _ tucky bluegrass: a,en- 


somewhat curved. It is black, dull, and somewhat grooved 1#78¢4; >,naturalsize. 


lengthwise. (Fig. 35.) 
O 


Bul. 85, Bureau of Plant Industry, U. S. Dept. of Agriculture. 


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U. S. DEPARTMENT OF AGRICULTURE. 


BUREAU OF PLANT INDUSTRY—BULLETIN NO. 85. 
B. T. GALLOWAY, Chief of Bureau. { 


THE PRINCIPLES OF MUSHROOM GROWING 
AND MUSHROOM SPAWN MAKING. - 


BY 


B. M. DUGGAR, 
PROFESSOR OF BOTANY IN THE UNIVERSITY OF MIssouRI, AND 
COLLABORATOR OF THE BUREAU OF PLANT INDUSTRY. 


VEGETABLE PATHOLOGICAL AND PHYSIOLOGICAL 
INVESTIGATIONS. 


IssUED NOVEMBER 15, 1905. 


. a A y 
i —— 


Ss 


WASHINGTON: 
GOVERNMENT PRINTING OFFICE. 
1905. 


BUREAU OF PLANT INDUSTRY. 


B. T. GALLOWAY, 
Pathologist and Physiologist, and Chief of Bureau. 


VEGETABLE PATHOLOGICAL AND PHYSIOLOGICAL INVESTIGATIONS. 
ALBERT F. Woops, Pathologist and Physiologist in Charge, Acting Chief of Bureau in 
Absence of Chief. 

BOTANICAL INVESTIGATIONS. 

FREDERICK V. COVILLE, Botanist in Charge. 


FARM MANAGEMENT. 
W. J. SPILLMAN, Agriculturist in Charge. 


POMOLOGICAL INVESTIGATIONS. 
G. B. Brackett, Pomologist in Charge. 


SEED AND PLANT INTRODUCTION AND DISTRIBUTION. 
A. J. PieTERS, Botanist in Charge. 


ARLINGTON EXPERIMENTAL FARM. 
L. C. CorBpett, Horticulturist in Charge. 


INVESTIGATIONS IN THE AGRICULTURAL ECONOMY OF TROPICAL AND SUB- 
TROPICAL PLANTS. 


O. F. Cook, Bionomist in Charge. 


DRUG AND POISONOUS PLANT INVESTIGATIONS AND TEA CULTURE 
INVESTIGATIONS. 


RopNrEy H. Trung, Physiologist in Charge. 


WESTERN AGRICULTURAL EXTENSION. 
CarL S. Scorre tp, Agriculturist in Charge. 


EXPERIMENTAL GARDENS AND GROUNDS. 
E. M. Byrnes, Superintendent. 


SEED LABORATORY. 
Epcar Brown, Botanist in Charge. 


J. E. ROCKWELL, Editor. 
JAMES E. JONES, Chief Clerk. 


VEGETABLE PATHOLOGICAL AND PHYSIOLOGICAL INVESTIGATIONS. 
SCIENTIFIC STAFF. 
ALBERT F. Woops, Pathologist and Physiologist in Charge. 


ERWIN F. Situ, Pathologist in Charge of Laboratory of Plant Pathology. 

Hereert J. Wesser, Physiologist in Charge of Laboratory of Plant Breeding. 

WALTER T. SWINGLE, Physiologist in Charge of Laboratory of Plant Life History. 

NEWTON B. Pierce, Pathologist in Charge of Pacific Coast Laboratory. 

M. B. WaITE, Pathologist in Charge of Investigations of Diseases of Orchard Fruits. 

MARK ALFRED CARLETON, Cerealist in Charge of Cereal Laboratory. 

HERMANN VON SCHRENK, in Charge of Mississippi Valley Laboratory. 

P. H. Rours, Pathologist in Charge of Subtropical Laboratory. 

C. O. TOWNSEND, Pathologist in Charge of Sugar Beet Investigations. 

T. H. Kearney, A. D. SHAMEL, Physiologists; Plant Breeding. 

P. H. Dorsert,t CoRNELIUS L. SHEAR, WILLIAM A. ORTON, W. M. Scott, ERNST A. 
3HSSEY, BE. M. FREBMAN, Pathologists. 

E. C. Cuitcorr, Expert in Cultivating Methods, Cereal Laboratory. 

Cc. R. BAL, Assistant Agrostologist, Cereal Laboratory. 

JoserH B. CHAMBERLAIN,’ J. AnrHur Le Cuerc,e Physiological Chemists. 

FLora W. Parrerson, Mycologist. 

CuARLES P. HARTLEY, KARL F. KELLERMAN, JESSE B. Norton, CHARLES J. BRAND, 
T. RALPH Roprinson, Assistants in Physiology. 

DEANE B. SwWINGLE, Grorce G. Hepecock, Assistants in Pathology. 

PERLEY SPAULDING, P. J. O'GARA, FLORENCE Hepers, Henry A. MILLER, ERNEST B. 
Brown, LESLIE A. Frrz, LEoNARD L. Harter, JoHN O. Merwin, A. H. Lerpien, H. F. 
BLANCHARD, Scientific Assistants. 

W. W. Cosry, Tobacco Barpert. 

JOuUN VAN LEENHOFY, Jr., T. D. BeckwitH, Laperts. 


“ Detailed to Seed and Plant Introduction and Distribution. 
+ Detailed to Bureau of Chemistry. 
¢ Detailed from Bureau of Chemistry. 


wy 


LETTER OF TRANSMITTAL. 


U. S. DeparTMENT oF AGRICULTURE, 
Bureau or Piuant Inpustry, 
OFFICE OF THE CHIEF, 
Washington, D. C., August 21, 1905. 


Str: I have the honor to transmit herewith a paper entitled “ The 
Principles of Mushroom Growing and Mushroom Spawn Making,” 
and to recommend that it be published as Bulletin No. 85 of the series 
of this Bureau. 

This paper was prepared by Dr. B. M. Duggar, Professor of 
Botany in the University of Missouri and Collaborator with the 
Office of Vegetable Pathological and Physiological Investigations of 
this Bureau. Under the direction of the Pathologist and Physiolo- 
gist, Doctor Duggar has been engaged for several years in the inves- 
tigation of mushroom culture in all of its phases, and great advances 
have been made, especially in the production of purer and _ better 
spawns. 

The accompanying illustrations are necessary to a complete under- 
standing of the text of this bulletin. 

Respectfully, B. T. Gattoway, 
Chief of Bureau. 

Hon. James WILson, 

Secretary of Agriculture. 


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PRETAGE. 


The bulletin submitted herewith presents the results of the work 
up to the present time on the problems of mushroom culture and 
spawn making. The first publication on the subject from the stand- 
point of pure culture was Bulletin No. 16 of the Bureau of Plant 
Industry. This was followed by a Farmers’ Bulletin (No. 204) on 
mushroom culture, presenting the results of our work for the use of 
the practical grower. As an outcome of the work Doctor Duggar 
has already accomplished, spawn of pure-culture origin is now being 
produced on a very large scale by several growers and is giving excel- 
lent results. This method enables the grower to improve and main- 
tain the most desirable varieties of mushrooms in the same manner as 
is possible with other plants propagated from cuttings or buds. 
Information which would enable a grower to accomplish this has not 
been up to this time available. The general method of securing pure 
cultures as here described will enable the experimenter to cultivate 
spawn of other edible species of mushrooms in case it should be found 
desirable to cultivate them. The methods described differ radically 
from any hitherto used. They are of more general application and 
give far better results. 

For the past three years this work has been carried on in coopera- 
tion with the University of Missouri, Doctor Duggar having left the 
Department to accept the professorship of botany in that institution. 
We wish to express our appreciation of the facilities furnished by 
the university for continuing this work. 

Apert F. Woops, 
Pathologist and Physiologist. 
OrFice oF VEvETABLE PATHOLOGICAL 
AND PuysyoLocicaL INVESTIGATIONS, 
Washington, D. C., June 16, 1905. 


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CON TENTS. 


LUE DUVET Se) a cane 
General considerations 
MEIC PTEECAS OTIC Se eh re een ee mee meen eee ee Ree 
(2 ELS OYE Gpnere Ge ee a ee ee oe ee ee ae 
fmeviewiOr CATIIeT WOTrk | _-\ 2 -2semeaseeane en eee -- 
TL SESE gs 3s pee Dl Lg det lial kee oot i aged pig = ae ed = ha 
netrere ey ees SE Tay al  SeC ee: se SLU ICiiAL Pee! 
LOR Cin ge SO ae a eam re Pre i eee ee ee eee ge eer ae ye ae yoete 
Growth on manure and other complex media 
Growth on chemically known media 
See aOM OlsSPeCIal TESMLUS oe on oe an ae apg aie a apg ee = 
Pectdernidemicaling mein = ses tk SSE ree! at Pe. Le ae 
eranare and momture._-— =f 2st oe a 
reimenOmOr hie COMPOS... _ 2. 4 2 | ae oo bie. 8 ela ee cee hoe 
DUS LASTi ih CLP L200 saeco een i Dp eae alls 5 a, ina, eee 
eNO CAECERESS CERES OCS a i = 
LSE LEP OL SiGe iet glee teen baal heise ear ennai aed. Shel cari ame a ee 
SeREneHAES hb Olid, MO. ms a os OT ne 
Variability in mushrooms grown under different conditions 
The cultivation of various species of mushrooms____________________.- 
OT GUSTER NTS) Sag DET ea ETT FS Pp Se a ap Se a Aer Ee De 
pemeerrartsies 1 GE UiibOd SLALCS 8 ne me ee ee 
(gL bSDG TATU ATI 22 genl? ett tingling OA» al pute agement nl a hee 
MEANS SIT GLIA VE INCA TT eros a kt. bet TER OSTA OS Pets ae A a ee 
Pare hancer praeL NOG ss) yerytel fey oes eee ay weg Le 
Pe Gein ee sINGD NOG foes ates ype AS oe ge eh the he eet ee wes 
(8 SESS reg 62) Ee ee ee ae en ae pan 
The tissue-culture method 
Mise HITHET Cre LOCCSA ttn = aye ee enn a eee Coo ee Se ew 
The vitality of mushroom spawn 


or or Gr cr or cr ¢ 
or me & WH 


io 6) 


PuatTE I. 


a 


III. 


VI. 


Vil. 


ILLUSTRATIONS. 


Page. 
A fine bed of mushrooms grown from spawn of pure-culture 
i gt | a eae eer SPENT Frontispiece 
Fig. 1.—A fine cluster of Agaricus campestris, the horticultural 
variety Columbia. Fig. 2.—Morels (Morchella esculenta), one 
of the finést edible fungi --____.___- 2... 2... 4322 eee 60 
Fig. 1.—Agaricus fabaceus, the almond-flavored mushroom. Fig. 
2.— Agaricus viliaticus, a promising species, fleshy and prolific. 60 


. Fig. 1.—A young specimen of the common puffball (Calvatia 


craniiformis). Fig. 2.—The oyster mushroom (Pleurotus os- 
treatus), growing on decayed willow log_--_------------------- 60 


. Fig. 1.—A mushroom house provided with gas-piping framework 


for shelf beds. Fig. 2.—The preparation of compost-----...--- 60 
Fig. 1.—A large mushrcom establishment—a common form of 
mushroom house. Fig. 2.—The method of making pure cul- 
tures, showing the apparatus and materials - _---- i 60 
Fig. 1.—Mushrooms prepared for the American market. Fig. 
2.—Good (** well-run’) mushroom spawn, brick form ------ 60 


a) 
s 


S 


B. P. I.—182. Vv. P. P. I.—142. 


THE PRINCIPLES OF MUSHROOM GROWING AND 
MUSHROOM SPAWN MAKING 


INTRODUCTION. 


For a number of years there has been an increasing demand in the 
United States for information concerning mushroom growing. In 
the horticultural and agricultural press many individual practices 
have been presented; but in order to give rational encouragement to 
mushroom growing in favorable sections of this country it was recog- 
nized at the outset of the investigations undertaken by the writer that 
much experimental work would be required. Bearing upon the cul- 
ture of Agaricus campestris* a number of physiological questions 
were demanding attention, for it was desirable to ascertain (1) the 
conditions of spore germination, in order that “ virgin ” spawn might 
be propagated and the principle of selection attempted; (2) the 
relation of this fungus to nutrients, or a determination of the sub- 
stances or compounds which might best serve as food materials; and 
(3) the relation of the growing mycelium and of mushroom produc- 
tion to temperature, moisture, and other conditions of the environ- 
ment. In the next place it would be necessary to determine the 
application of any physiological principles established to the practice 
of mushroom growing and mushroom spawn making. 

In connection with a presentation of the results of the experimental 
work ” it seems desirable to include also a more or less comprehensive 
account of the present status of mushroom growing at home and 
abroad. 


«Throughout this paper the writer has employed the generic name Agaricus 
in the sense in which it is usually understood by those interested in the practical 
side of the work. 

b During 1903-4 the writer was assisted in the experimental work by Mr. A. 
M. Ferguson, instructor in botany in the University of Texas, at that time special 
agent of the Department of Agriculture, and during 1904-5 similar assistance 
has been rendered by Mr. L. F. Childers, student assistant. Through the assist- 
ance thus given it has been possible to complete an unusual amount of 
experimental work, only a portion of which can be described in detail, although 
it has all been taken into consideration in the conclusions drawn. 


10 MUSHROOM GROWING AND SPAWN MAKING, 


It is not possible at this time to give more than a few brief sugges- 
tions concerning the possibility of cultivating other edible species 
than Agaricus campestris. The determination of the fundamental 
needs of diverse species will require study during a term of years. 


GENERAL CONSIDERATIONS. 


The propagation of Agaricus campestris does not seem to have been 
undertaken to any extent by the ancient Greeks or Romans. The 
occasional references to mushrooms in the classics are very general, 
as a rule, and do not suggest that artificial propagation was attempted. 
In the vicinity of Paris Agaricus campestris has been cultivated for 
several centuries, and the plants have certainly been sold on the open 
market quite as long. It has not been possible to ascertain whether 
the methods now in vogue are essentially the same as those employed 
a few centuries ago. It is very probable, however, that the methods 
have been gradually improved. It would appear that the cultivation 
in caves is comparatively recent. The earliest records obtainablé 
concerning the cultivation of mushrooms in the underground quarries 
indicate that this practice was not common previous to the nineteenth 
century. 

Mushrooms are to-day extensively grown in England and France, 
and to a limited extent in Belgium, in Germany, and in many other 
countries. Paris remains, however, the center of commercial pro- 
duction. In the vicinity of that city the culture of mushrooms is now 
almost entirely confined to the underground limestone quarries or 
cement mines. The caves used for this purpose are termed “ carriéres * 
or * champignoniéres.” These caves may consist of a labyrinth of gal- 
leries, or halls, ranging from 5 to 50 feet in width and from 5 to 30 
feet in height. In some regions the earth is practically honey- 
combed by them, and the extent of the cave space used by the larger 
growers may be measured by miles. For the most part the ventila- 
ting system is perfect, every cave system possessing numerous air 
shafts, protected at the surface by wooden towers. Artificial parti- 
tions in the caves themselves enable the operator to control the venti- 
lation. Until recent times the cultural methods have been more or 
less sacredly guarded by the growers, and even to-day it is not easy 
to get permission to make a casual visit to the champignoniéres. In 
many cases the work has been followed from generation to generation 
within the same family. There are at present, however, large corpo- 
rations in control of some of the most fanyous caves. 


ain a painting of the early seventeenth century (that of a Fishmonger’s and 
Poulterer’s Shop, by Jordaens and Van Utrecht, in the Gallery of Old Pictures, 
Brussels) Agaricus campestris and Boletus are shown on sale as a conspicuous 
part of a market scene. 


MARKET CONDITIONS. 11 


In the United States fresh mushrooms have only recently been of 
any importance commercially, although florists and gardeners of 
English and French training have long been successful growers on 
a small scale. Nevertheless, during the past decade or so, the record 
of failures has been most conspicuous, and it is certain that of the 
many who attempted this work only a few, relatively, were uniformly 
successful. 

The conditions under which mushrooms may be successfully grown 
are limited, and intelligent attention is therefore essential. It must 
be said, moreover, that the majority of failures may be directly 
traced to erroneous ideas as to the cultural requisites, or to a reckless 
disregard of. conditions. The essential conditions will be subse- 
quently defined in detail, but it may be stated here that failures are 
usually due to one or more of the following causes: (1) Poor spawn; 
(2) very poor manure; (3) unfavorable temperature; and (4) heavy 
watering during the early stages of growth. 

Under suitable conditions mushrooms may be grown with assur- 
ance of success. Ordinarily they are grown only where the condi- 
tions may be controlled, and success should therefore be invariable. 


MARKET CONDITIONS. 


In the vicinity of Paris the mushroom industry has been remark- 
ably developed during the past eight or ten years. The total product 
sold through the central market of Paris in 1898 was nearly 4,000,000 
pounds; the quantity for 1900 is given as approximately 8,500,000 
pounds, and for 1901 nearly 10,000,000 pounds. These figures show 
most convincingly the present status of the mushroom industry in 
France. It may be safely assumed that more than one-third of this 
quantity is consumed in a fresh state in and about the city. The 
growth of the canning industry during this period has also been 
remarkable. In 1898 about 1,800,000 pounds were preserved, while in 
1901 the canned product amounted to nearly 6,200,000 pounds. Dur- 
ing 1901 the approximate monthly production of mushrooms ranged 
from 651,000 pounds to 985,000 pounds, from which it is evident that 
these caves yield heavily throughout the year. In some instances 
growers are able to get a crop every four or five months. 

It is extremely difficult to estimate the quantity of mushrooms 
grown in the United States. It is certain, however, that the produc- 
tion has increased very greatly, and particularly within the last four 
cr five years. In the vicinity of several of our larger cities there are 
to-day individual growers who produce more than the total commer- 
cial output in the neighborhood of those cities ten years ago. 

There is now a very good open market for fresh mushrooms in a 
few of the larger cities, although many large growers continue to 


12 MUSHROOM GROWING AND SPAWN MAKING. 


sell entirely by contract or by special orders to hotels and restaurants. 
With such an enormous comparative consumption of the canned 
product, there is every reason to believe that fresh mushrooms can 
be sold in much greater quantity as soon as this product becomes a 
certain factor in the market. With canning factories to take the 
surplus product, growers could afford to accept a smaller margin of 
profit, and this would place mushrooms within reach of many who 
may not be able to purchase them at present average prices. Agaricus 
campestris and its varieties and allied species are perhaps the only 
fresh mushrooms commonly salable in the markets of American 
cities. Throughout practically the whole of Europe several other 
species are legitimate market products. The more delicate or fleshy 
forms of the latter are sold as fresh mushrooms; others are dried, 
and some of these, being tougher, are used only for soups, sauces, and 
gravies. Besides the various species of truffle and morel, any special 
mention of which will be omitted here, the French market to-day 
legalizes the sale of five or six other species of mushrooms. 


GERMINATION STUDIES. 


Review of earlier work.—In a small way the germination of the 
spores of Basidiomycetes has received attention from the earliest 
times. A complete historical review of the literature dealing with 
spore germination will be found in Bulletin No. 16 of the Bureau of 
Plant Industry. It will be seen that most of the early work fur- 
nishes only incidental references to spore germination. By far the 
most important contributions made by early workers to this particu- 
lar subject were several papers by Hoffmann.* It is not to be expected 
that the method employed by him would yield accurate results. 
Nevertheless, the work of Hoffmann is comprehensiye for that time. 
Brefeld,’ in his extensive reports upon the Basidiomycetes, gives the 
results of germination studies with a large number of the fleshy fungi. 
More than 200 species were used in his various experiments, and suc- 
cessful germination is recorded for about 160 species. 

In 1898 the writer became interested in some attempts to germinate 
the spores of certain Basidiomycetes. Subsequently the problem 
received incidental attention in connection with some general studies 
on the physiology of spore germination.° The work progressed only 


aHoffmann, H. Ueber Pilzkeimungen. Botan. Zeitg., 19: 209-214, 217-219. 
1859. Beitriige zur Entwickelungsgeschichte und Anatomie der Agaricineen. 
Botan. Zeitg., 18: 389-395, 397-404. 1860. Untersuchungen tiber die Keimung 
der Pilzsporen. Jahrb. f. wiss. Botanik, 2: 267-837. 1860. 

> Brefeld, O. Botanische Untersuchungen iiber Schimmelpilze. Basidiomy- 
ceten, I, Bd. I, H. 3. 1877. Untersuch. a. d. Gesammtgebiete der Mykologie. 
Basidiomyceten, II, H. 7; III, H. 8 1888-89. 

eDuggar, B. M. Physiological Studies with Reference to the Germination of 
Certain Fungous Spores. Bot. Gaz., 31: 38-66. 


GERMINATION STUDIES. 13 


far enough to suggest that an investigation of the factors influencing 
germination might yield studies of special interest. During 1900- 
1901 Dr. Margaret C. Ferguson undertook a systematic investiga- 
tion of the relation of stimuli to germination in certain species. 
The results* have made it evident that the problems involved are 
not the well-known simple nutrient or physical factors. Miss Fer- 
guson spent much time in experimenting with a great variety of 
nutrient media and special stimuli. Several thousand cultures were 
made. In the majority of these cultures Agaricus campestris was 
used, and it is shown that from the known ecological relationships 
of this fungus one could not possibly predicate the probable stimulus 
for germination. In fact, with no known nutrient medium or special 
chemical stimulus employed, was there anything more than erratic 
germination. Nevertheless, the work was finally very successful 
in the discovery that almost a perfect percentage of germination 
could be secured by the influence of the living hyphe of Agaricus 
campestris upon the spores, as announced in the statement that “if 
a few spores are able to germinate under the cultural conditions, 
or if a bit of the mycelium of Agaricus campestris be introduced into 
the culture, the growth resulting will in either case cause or make 
possible the germination of nearly all the spores of the culture, pro- 
vided, of course, that the other conditions are not such as to inhibit 
germination.” 

The stimulus would seem to be of enzymatic nature. No other 
mycelium tested produced a similar effect. This was a distinct 
advance in our knowledge of factors influencing germination. The 
stimulus, however, could only be looked upon as perhaps a substitu- 
tion stimulus. It did not seem possible that it could obtain in nature, 
nor could it be looked upon as wholly satisfactory from a practical 
point of view. 

Miss Ferguson’s results offered encouragement; but, nevertheless, 
the problems with Agaricus campestris and related species were left 
open for further investigation. It should, perhaps, be emphasized 
that prior to 1902 no method had been published, so far as can be 
learned, whereby one might be able to obtain with uniformity the 
germination of Agaricus campestris. It is quite certain that Chev- 
reul and others obtained at best only erratic results. Nevertheless, 
as early as 1893 Costantin and Matruchot ® announced that a method 
had been developed by them whereby they were able to germinate the 


@¥Ferguson, M. C. A Preliminary Study of the Germination of the Spores of 
Agaricus Campestris and Other Basidiomycetous Fungi. Bulletin No. 16, Bu- 
reau of Plant Industry, U. S. Dept. Agriculture, pp. 1-48. 1902. 

>Costantin and Matruchot. Nouveau procédé de culture du champignon de 
couche. Compt. Rend. de l’Acad. des Sci., 117 (2): 70-72. (Compare, also, 
Bul. Soe. de Biol., 2 December, 1893.) 


14 MUSHROOM GROWING AND SPAWN MAKING. 


spores and to grow in pure culture the mycelium of Agaricus cam- 
pestris. Information concerning the details of the method employed 
was avoided in the reports of this announcement and in subsequent 
references to the process.* In the first announcement the method is 
stated as follows: 


Method followed.—The spores are collected free from contaminations, and 
in order to preserve them in that condition are sown on a certain sterilized 
nutritive medium. We obtain in this manner a twisted mycelium which con-~ 
stitutes pure spawn. By repeated cultures on an identical substratum the 
spawn can be multiplied indefinitely, and is transferred at a proper time to 
sterilized manure, where it develops abundantly in several weeks. At that 
stage it possesses the characteristic appearance and odor of natural spawn. 
It can then be sown in a bed of ordinary manure, to which it adheres and where 
it grows and fruits normally. 


In the later paper cited, writing of the recent improvements in 
mushroom culture, Costantin expresses himself as follows: 


We have succeeded in manufacturing an artificial spawn obtained from the 
spore germinated on a medium free from contamination. It is then pure 
spawn. We can state further that it is virgin spawn. 


In 1897 Répin ” claimed to have independently arrived at results 
similar to those obtained by Costantin and Matruchot. Concerning 
his germination studies he says: 


It is only recently that the study of this question has been renewed, inde- 
pendently and simultaneously by Costantin and Matruchot. 

There is nothing unusual in the germination of the spores of Agaricus. 
Spores can be germinated on media such as used in bacteriology, on wet sand, 
or in moist air as well aS on manure. Without doubt, germination is not pro- 
duced with the same spontaneity and rapidity as in the case of the spores of 
lower fungi, which fact makes it necessary to promote the process by some 
artifices, but they are only sleight-of-hand tricks, variable according to the 
operators, and which are acquired after some unsuccessful attempts. The 
spores which should germinate (and these are always in the minority) begin 
by swelling. This very simple method makes it possible to obtain virgin 
spawn at pleasure. It is applied industrially in the manufacture of spawn of 
Agaricus from cultures which I have made. 


So far as the writer has been able to ascertain, therefore, no descrip- 
tion of the method employed by the above writers is to be found. The 
veport of Miss Ferguson’s work is accordingly the only available 
scientific record defining the conditions under which germination 
had been constantly obtained up to this time. 

Experimental worl:.—The writer has been able to confirm Miss Fer- 
guson’s work repeatedly, and at the same time numerous series of 
experiments have been made to test further the possibility of influenc- 


¢Costantin, J. La culture du champignon de couche et ses récent perfection- 
nements. Extrait du Revue Scientifique. April, 1894. 

’Reépin, C. Le blane vierge de semis pour la culture du champignon d-¢ 
couche. Revue Générale des Sciences. (September 15, 1897.) 


GERMINATION STUDIES. 15 


ing germination by chemical stimuli. In distilled water, on the one 
hand, and in plant decoctions (such as decoctions of beans, sugar 
beets, mushrooms, potatoes, etc.) and in bouillon, on the other hand, 
there have been tested a large number of inorganic and organic salts, 
carbohydrates, nitrogenous compounds, and active enzymes. 

The results of one series of experiments are tabulated in detail. 
In general, it has been found that dulcite, monobasic magnesium 
phosphate, magnesium phosphite, magnesium potassium ammonium 
phosphate, ammonium molybdate, magnesium lactophosphate, dibasic 
calcium phosphate, and other salts, especially phosphates, have in one 
medium or another been more or less effective as stimuli for germina- 
tion. Unfortunately, none of the substances mentioned, apparently, 
are very strong stimuli; they are unable to cause invariable germina- 
tion in all nutrient media. Moreover, in subsequent series, where the 
conditions have been the same, within experimental possibilities, 
wholly analogous results have not always been obtained. No account 
has been taken, however, of the particular variety of Agaricus cam- 
pestris from which the spores were obtained, and it may be that this 
will influence the results. 

It is to be noted from the following table that Miss Ferguson’s 
method of employing living bits of mycelium was modified by the use 
of small pieces of the inner tissue of a young mushroom taken under 
sterile conditions. It was found that often a new growth of mycelium 
was developed from this tissue. Whenever this growth appeared, 
the influence upon spores in the drop culture was, as might be 
expected, the same as had been demonstrated for the living mycelium. 
Frequently a few spores germinated within from three to five days. 
The most interesting conclusion, however; which could be drawn from 
the cultures in which small bits of tissue were used was the following: 
Under favorable conditions a small piece of the inner growing tissue 
of a mushroom is capable of producing a mycelium with great readi- 
ness. This fact has been utilized, as shown in detail later, in the 
development of a new and effective method of securing pure cultures 
of fleshy fungi in general. ‘ 


TABLE I.—E«tent of germination. 


No. Media. After 3 days. After 5 days. 


Distilled water ..........--------- {eos arteries, Gets As before. 


GUM a a eee 
$+ per cent KH.PO,.-....--..---.-- 
4 per cent KH»PO, in bouillon - 

Ppericent Wall Oye. 0 2733 22 
4 per cent K.HPO, i in bouillon - 
+ per cent NasHPQ, --....-...--- 
4 per cent NaoHPO, in bouillon_. 
4 per cent (NH4)2.HPO, eee et 
+ aa cent (NH,)sHPO, in bou- 


Sommer lo 


23994—No. 85—06 m——2 


16 


MUSHROOM GROWING AND SPAWN MAKING, 


TABLE I.—Extent of germination—Continued. 


Media. After 3 days After 5 days. 
3 per cent MgH,(PO,4)o -----.----- RU DOLGS 2 coo oS 3 ne Sate 50 per cent. ~ 
| + per cent MgH,(PO,). with bou- {on BOERS LLECES. Bis see 3 per cent. 
illon. . eNOS esse, ho ee one 
3 per cent MgHPQy. -.....---.---- mabe O22 03.2 OE {i = eae 
3 per cent MgHPO, in bouillon --| a—10 spores -_--...----.----- Conia spores badly 
injured. 
+ per cent Mg(NHy)PO,---..----- Moric eet selprosicler. fe None. 
Ps = cent Mg(NH,)PO, in bou- { 2 spores _.....-.-..--------- As before; injured. 
1110n. 
+ per cent MgK(NH,)PO,--..--_- Few spores __...__--.-.----| 5 per cent. 
4 percent MgK(NHy,)Ho(PO,4)oin |_....do -_.-._._-.-.---.------ Do. 


bouillon. 


a—Note 2:-o2te-9 BGs Few spores. 

+ per cent (NHy)2CsH40¢ spre dies {$ —NONG.-. 9 eo ae eee 10 percent. 

4 ie cent (NHy4)oCyHyO, in bou- | 10 spores __....----.-------- As before; injured. 
illon. 

4 ne cent magnesium lactophos- |-_--- Go ¢ .o28 se -seast 2-2 se 5-10 per cent. 
phate. 

+ per cent magnesium lactophos- |--.-- G0 2es49-h£--) 43 se 2-5 per cent. 
phate in bouillon. 

4 per cent CayHo(PO4)s -----.----- IN ONO)... 423 hee ee 1-2 per cent. 

3 per cent CasH»(PO,).in bouillon) 10 spores -_-...---....-.--.-] Injured. 

4+ per cent KCHO>, J. ...- 2-222. --=- Worle. -t¢-c¢ saze-ect od Pees 1-2 per cent. 

+ per cent MgHROs. = 2222: 10 spotes’:. 2icsence-h aban 10-59 per cent. 

+ per cent MgHPO; in bouillon--|_---- €0.25..4414.2t22 ees 1 per cent. 

+ per cent MgK(NH,)H2(PO,)o |_---- GO ences 2 ve eee eee 1-2 per cent. 
in mushroom decoction. 

+ per cent KH»PO, in mushroom | None_-_--...__-...-----.---- None 
decoction. 

4 per cent KeHPO, in mushroom |____- dof 2310; eee Do. 
decoction. 

+ per cent Na,HPO, in mush- | Few spores _-_....--...-----| Injured. 


room decoction. : 
3 per cent (NH,)oHPO, in mush- 
room decoction. 


|} per cent MgHPO, in mush- 


room decoction. 


coms tig tat Ae Bie eee ee 


+ per cent Mg(NH,)PO,in mush- 
room decoction. 


| ¢per cent (NHy)oC,H,O, in mush- 


| room decoction. 
| } per cent magnesium lactophos- 
phate in mushroom decoction. 
+ per cent CasHs(PO,4). in mush- 
| yvoom decoction. 
+ per cent KCHO, in mushroom 


decoction. ; 
+ per cent MgHPO; in mush- 
room decoction. 


Decoction of mushrooms ____-___- 


_ Living tissue of mushroom in 
mushroom decoction. 


lor 2 spores __.22--__-_-- = 


10-50 per ents. - 2.522 2 
Pew Sporesern 24 = ok - 


ib-2 per cént_cice hi iit 


\fa—2 per cent ..-.5..--2.f2. 
{pov ery few spores - 

| a—Few spores @ -_- 
| b—None? _____- 


|\fa—Few spores@......_-.... 


| \b—None > 


Few spores. 


Hew Spores © -22esee7 Seat. -8 2-5 per cent. 


10-20 per cent. 
Contaminated. 


2 Per coltet soso 33-0  oeee Contaminated: 50 per 
cent, but injured. 

10 spores) feed e 3-5 per cent: injured. 

5-8 per. cent.......--....---; 10 percent. 

2-3 per, Con has S.-ce- esa 10-20 per cent. 


a—5 per cent. 
b—Contaminated 
2 per cent. 

1-2 per cent. 


--] 12 spores. 


As before. 
None. 


“In this cell the tissue developed a new growth. 
» No growth from tissue introduced. 


On account of the fact that magnesium phosphite and magnesium 
potassium ammonium phosphate had in most cases proved to be 
stimuli for germination, experiments were next made to determine 
the efficiency of these salts on various media, as indicated in the 
table on the following page. 


‘GERMINATION STUDIES. 17 


TABLE II.—#fficiency of salts on various media. 


Nature of compost. |Appearance after 26 Grace |l | Nature of compost. “Appearance after 25 days. 
Well-rotted stable | No growth. Well-rotted cow ma-)| Good growth. 
manure.@ ||  nure. 0 
10th See ipa lta Fine growth. | Peaty mmDke aes agen No growth. 
Half-rotted stable | One, fair growth: one, | 1D Yo <a ee Do. 
manure. @ good growth. Maryland peata_____ Do. 
a ace No growth. 1D Topsy eae Do. 
| Well- oad Ginkgo Do. 
Fresh stablemanure@| One, good growth; one, leaf mold.« 
slight growth. | Ho.0. a eee 0. 
108 ae eee One, good growth; one, | Cotton-seed motes@_| One, no growth; one, fine 
fine growth. ! growth. 
Wellrotted cow ma- | One, good growth; one, | 1D Yo 5a eee Fine growth. 
nure. 4 slight growth. | 


«Watered with concentrated solution of magnesium phosphite. 
+ Watered with strong solution of magnesium potassium ammonium phosphate. 


Large test tubes were used in these experiments, and duplicate 
cultures were made in every instance. From these and from numer- 
ous other cultures it was ascertained that germination could not be 
obtained invariably, even on favorable media and under pure-culture 
conditions, by the use of these partial stimuli. Nevertheless, the 
percentage of failures has usually been small. By means of the 
stimulus given by magnesium phosphite it has also been possible to 
get growth from the spores in test-tube cultures with gray filter 
paper as the solid substratum and various plant decoctions and cul- 
ture solutions as the nutrients. Details of these results, however, 
may be omitted. 

In many cases it has been possible to obtain growth from the 
spores by the use of the stimulating salts which have been mentioned 
im connection with the germination studies. Where it is desired to 
make experiments along this line the writer has found it more 
practicable to use spores from a mushroom as young as possible. 
If one takes a mushroom just at the time that the veil is breaking, 
inoculations may be readily made from the spores and few contami- 
nations will result. In this case, by means of a sterile needle, or 
scalpel, a few spores may be removed from the’ spore-bearing, or 
gill, surface and these may be transferred to the tubes in the same 
way as were bits of the fresh tissue. It is also possible to secure a 
spore print from a mushroom the gill surface of which has not been 
exposed to germs of the atmosphere. In the latter case it is desir- 
able to remove stem and partial veil, peel off the incurved edges of 
the cap which have been in contact with the soil, and place the cap, 
gill surface downward, in a sterilized dish or on sterile paper. If 
this is then kept free from dust, a spore print may be obtained, which 
should not be contaminated by foreign germs. This print may then 
be used in making a large number of spore cultures. 

Experiments were also made in which pots of unsterile composts 
and manures were inoculated, on the one hand, with spores, and, on 


18 MUSHROOM GROWING AND SPAWN MAKING. 


the other hand, for control purposes, with spawn from pure cultures. 
The duration of the experiments was two months. Some of these 
pots were watered with a mineral nutrient solution including as one 
constituent magnesium phosphite, designated 1, others with the same 
solution to which was also added a small quantity of dried blood, 
designated }’, and the remainder with pure water. The results are 
tabulated as follows: 


TABLE II1.—Hartent of growth of spores and spawn in pots. 


| | 
Cette mp “'ianare | ,staule "manure. Qlastable| pertlizer 
Rspairba $I eV Name db a) do 
Spores:\b0) AS a te dee ee 
Spawn. me Ee Daee us. dor crea ceente tae 
sk SS a ae || SIR WORE ORES” 55 foe 
Spawn Were Det dost of Bena ee 
| | 


TISSUE CULTURES. 


The suggestion which had presented itself of using bits of living 
tissue from a sporophore instead of spores seemed also, from general 
observations, to be of sufficient importance to warrant a thorough 
trial. During moist weather, or in a moist cellar where mushrooms 
are being grown, one will frequently find that an injury in a young 
mushroom is rapidly healed by a growth of hyphe from the edges 
of the injured area. The same thing had been noted in the open in 
the case of puffballs. In many instances, moreover, pure cultures 
of fungi in other groups have been obtained by the use of small bits 
of a sclerotial mass of tissue. Accordingly, a young sporophore of 
Agaricus campestris was obtained, and after breaking it open longi- 
tudinally a number of pieces of tissue from within were carefully 
removed witb a sterile scalpel to a sterile Petri dish. A number of 
cultures were then made by this tissue-culture method on a variety 
of nutrient media, such as bean pods, manure, leaf mold, ete. From 
this and from numerous other similar tests it was ascertained that 
when the mushrooms, from which the nocules of tissue are taken, 
are young and healthy, there is seldom an instance in which growth 
does not result. It was easily shown that failure to grow was gen- 
erally due to the advanced age of the mushroom used, to an unfavor- 
able medium, or to bacterial contamination. 

The first suecessful pure cultures were made by this method during 
the early spring of 1902 from mushrooms grown indoors. During 


TISSUE CULTURES. 19 


the following summer, or as other fleshy fungi appeared in the open, 
cultures were made from other forms in order to determine the 
general applicability of the method. The experiments were success- 
ful in most cases, although it was found almost impossible to obtain 
certain species of fungi in a condition young enough to be free from 
bacterial infestation. In general, the method seemed to commend 
itself strongly as a means of procuring pure cultures of desirable 
edible species, particularly of those species the spores of which could 
not be obtained pure or which could not be readily germinated. 
During the two subsequent seasons this method has been employed 
with a great variety of fungi representing many natural orders. No 
systematic endeavor has been made to determine the limitations of 
the tissue-culture method as applied to Basidiomycetes, but, inci- 
dental to the general studies, cultures have been made from forms 
differing very widely, not only in relationship but also in texture and 
in habitat. aed 
In all there is a record of 69 species having been tested upon one 
or another medium. In a few cases the cultures have invariably been 
contaminated, and it is to be supposed, perhaps, that the plants were 
collected in a condition too old for the purpose in hand. In only 
about ten forms has it seemed that there is no evident reason for the 
failure to develop mycelium. Of the remainder fully 40 have grown 
promptly on the media employed. The table following indicates the 
names of the species employed and the results obtained. It must be 
said, however, that cultures of a number of species were made of 
which no record was kept; among these, also, some grew and some 


failed. 3 


TABLE 1V.—fesults obtained from different species. 


Num- | 
Fungus. beret | Substratum. Result. 
tures. 

Agaricus arvensis -__..___..-.---- Few. Beans, manure, leaves, ete._....| Rapid growth. 
Agaricus augustus -_---.--------- i hig) 37ee lee en ae oe oe Contaminated. 
Agaricus campestris (various oc | Beans, manure, leaves, etc--____- Rapid growth. 

varieties). . 
MP aTiCHS fHUACOUS—.----2-.-5--.-| OG |-.--- GIS peeras pe MOR ach elle ea Sale mete Do. 
Agaricus fabaceus var ---_-------- basa i Ate oe eh gd ey ye nek Do. 
Agaricus placomyces- -----.------ ESTAS ere See ees eect ht eee ot Some growth. 
Agaricus villaticus--_-____------.-- . | Manure, leaves, ete-=--..----.--.- Rapid growth. 
Amanita frostiana --_....---.---- [UGC Gr tes | eae ne Se ES ES Contaminated. 
Amanita muscaria -_---.._-.___-- Reams ty eet eee os Goto rt 3 Do. 
Pere Vertn= ss 2 eet The 2 pe RE ene Se ee eee Do. 
Amanitopsis vaginata -___...._-- rs) Ba Gia ats syria _vlhees Do. 
armnlistis melled _-_.-. 2.2. Beans, leaves, dead wood, etc__.| Rapid growth. 
Boletinus porosus ---__.---------- oa | paris eL EU Ee IG Bhd) sae Slow growth. 
Te UE UnLRELLUG (EG Sieeumie Sos athe asin ial ila (ORY SoS ate al Ses ae Lael No growth. 

oletus miniato-violaceus ea tew CAE SEO SES 1d Do. 
Boletus peckil: _ 2. = .-.------5 5 eee lste Sal ole es eee Contaminated. 
Bovistella ohiensis____-_...._----- . | Beans, leaves, etc__......-..----. Rapid growth. 
Calvatia craniiformis -__..._____- MRE ls at Loe eee Sh ees Do. 
Calvatia cyathiforme -_. | Beans, leaves, soil, etc. __.-.._-_- Do. 
Calvatia rubro-flava__-- uJ . | Beans, leaves, etc_-.. 22. +.-.-_-- Do. 
Cantharellus cibarius ___._____._- Beans, se wes A FAM 271203 J No growth. 
EeGHE HE TOMNIOND = 2 22222222 -2-22-|°-- ~ I>. 22 GOL maw Be eereeeececncn eccecs oO. 


. e@cc indicates an indefinite nu-nber. 


20 MUSHROOM GROWING AND SPAWN MAKING, 


TABLE LV.—Results obtained from different species—Continued. 


Fungus. Substratum. . Result. 


AZHILOCY be LULUGGDS <2 ose cee ee =a Home growth. 
Clitocybe sp.?----- ..| Rapid growth. 
Clito Nias prunulus - Few. | Some grew well. 
Colly bia platyphylla - 1 No growth. 
Colly bi a radicata-__- Few*-|-_--- d Page growth. 
Collybia velutipes __.___- if eee GeO.ss si. bes eee Do. 
Coprinus atramentarius - cc | Beans, leaves, manure, etc ae | Rapid growth. 
Coprinus comatus ------ oc | Beans, manure, leaves, etc Do. 
Coprinus fimetarius- - Few. | Beans, leaves -_-..._--.-..- Do. 
Coprinus micaceus ---_- oc | Beans, leaves, manure, ete | Do. 
Cortinarius armillatus - d, 
Cortinarius castaneus 1 Do. 
Cortinarius sp.?. ---- Few. | Beans, leaves, manure -___- | Good growth. 
Daedalia quercina-_------- Few. | Beans, leaves, manure, etc --| Rapid growth. 
Hydnum caput medusae. 1 Beans _- _ Good growth. 
Hydnum coralloides-- weeds z Oud Do. 
Hydnum erinaceum - Pe Lee Oe eae ees 
Lactarius corrugis (?) Few. Slight growth, one. 
Lactarius piperatus 2 yet Saber 9 (a ie No growth. 
Lactarius volemus - 1 | Acid beans | Some growth 
\heienont eae eae ae oc | Beans -- No plas 
Lepiota americana - 1 LF eee do... 22. eee eet S 
Lepiota morgani -_-- A eS do... 2 Fase 2 eee | Bamse growth. 
Lepiota procera ---.--- oc | Beans, leaves, etc -..---.-----.-- | ieee 2 growth. 
Lepiota rhacodes - t Few. |..--- CO's. 2. Gel. 2 ee eee 
Lycoperdon gemmatum ee 2,| Beans. 2: 22.2 de - eds Oe | Good ye 
Lycoperdon wrightii. ---.-------- W002 eco coe ere Do. 
Morchella esculenta ----..----_--- 2 | Beans and leaves --..-...-..-==-. Do. 
Pluteus cervinus - Bec ye a I T'S 5 NB as OO 8 Bee sees ee ee Some growth. 
Pleurotus ostreatus __....__-.__-- cc | Beans, leaves, manure, etc_----- | Rapid growth. 
Pleurotus ulmarins -..2°2..-- 2-3 1") Beans “= 202 ee eee eee Do. 
Pholiota adiposa......--.-.------- 1 | Beans and leaves.........-----.- No gro owth. 
Polyporus betulinus_____--- ------ A“ Beatis! $9522) oe ee ee | Slow growth. 
Polyporus intybaceus ----.------- 1 Sees (ee 2 oe ee OE Do. 
Polyporus sulphureus_-_.-___----- 2 | Beans and leaves _____...-___---- "Rapid growth. 
Polystictus cinnabarinus -------- 2 leoaes (oo ae NS gm! Good growth. 
PUSSIES asi: oF eas ee 1 | "Beans ee eee | No growth. 
Bussula emetica__ i. .= +4. oc. | J8éans, 66G. Koad ree oe Often contaminated 
but some grew 
SEMASUE Otc tee etapa ae ce Now) 1 does... 2 5-c 2a pease No growth. 
Russnlasordiag: = 5° -3 52 “Secenk 3 1 hen (ep he GOS. 22 ee ee eee Do. 
Russula virescens ......---.-.£.-- Few. |_.--- GO. see ea ep eo Do. 
Secotium acuminatum __________- rh ee SS dow > .JAMS* . 2 AM Rees Slow growth. 
Strobilomyces strobilaceus -_-___- i ie) (ee G02. ee eee No growth. 
Deropnaria Spices oS <sseh ee i es dow. = dawk hee eee Contaminated. 
Tremella mycetophila --.--_.-__- Lj»? Goi & thle Als eS Se No growth. 
Tricholoma p2rsonatum ---.. ---- 1 | Beans and manure -.-.-..--.----..- Good growth. 


ALTICHOMONIA TUSSUUE oe. oes 2 | ‘Beans 4nd lGaves —._ 2. ---5e ae Do. 


It is not to be understood that the failures recorded in the forego- 
ing table indicate that these species will not grow. The evidence is 
that they did not grow upon the media used, and it is very probable 
that most of these could be propagated in culture by this method if 
a systematic attempt were made to determine what substrata are 
desirable. The writer believes that this statement holds true particu- 
larly in the case of certain species of Boletus. No attempt was 
made to cultivate Boleti in any other way than upon bean pods. A 
few mycelial threads were developed in such cases, but these failed 
to grow upon the bean, apparently dying before even the nutrients 
in the fragment of tissue were exhausted. 

It is interesting to note that many of the fungi which have given 
good growth have not hitherto been grown in pure culture. Accord- 
ing to Costantin and Matruchot,* Van Tieghem (1876) produced the 

a Costantin and Matruchot. Sur la production du mycelium des champignon 
sup¢rieurs. Extrait Compt. Rend. d. Séances de la Soc. de Biologie. January, 
1896, 


TISSUE CULTURES. ra | 


mycelium of Coprinus in pure culture. Later, Brefeld’ accom- 
plished the same result with many species of Coprinus, and also with 
Armillaria mellea. Costantin has also published a number of brief 
papers, or announcements, of successful cultures upon artificial media 
of the mycelium of several fleshy fungi. Besides Agaricus campestris, 
he has grown the mycelium of Amanita rubescens, Armillaria mellea, 
Collybia velutipes, Lepiota procera, Marasmius oreades, Tricholoma 
nudum, Pleurotus ostreatus, Pholiota aegerita, Coprinus comatus, 
Polyporus tuberaster, P. frondosus, Hydnum coralloides, Morchella 
esculenta, and perhaps a few others. He has also grown to maturity 
sporophores of Agaricus campestris, Coprinus comatus, and Tri- 
choloma nudum. Unfortunately, Costantin seldom indicates the 
substratum upon which his cultures were made. Falck? reports 
having produced in culture the sporophores of Collybia velutipes, 
Phlebia merismoides, Hypholoma fasciculare, Chalymotta campanu- 
lata, and Coprinus ephemerus in his studies upon the connection of 
oidial stages with perfect forms of the Basidiomycetes. In the work 
of the writer so far no special attempt has been made to obtain the 
sporophores of the fungi cultivated except in the case of Agaricus 
campestris. Nevertheless, the following species have fruited in pure 
’ culture upon the media indicated: 


Medium. 
Pee AMIVeSULIN: = 22 => oes oe Eee y WL ea LL E Manure. 
Perens rap ACES: tt)! Ae ee esi pe a ae Pee! Ss Manure: 
Mar riciswamycdalinust iver _ fet) sett ay pti Manure. 
PamaIe Ee), MOU CAs els eet ow Lee cope le pee eee: Beans. 
SITLL ToS Ce a ee SO ee OSS er. Soil. 
PTC MARIMOITOLWMNG | = — en 2 Se Soil. 
DEAS Reet Da 0 Naa cS Se: eS eS ER a Soil. 
PRMRMERAICTISE Sf) See ne ee Se eee ee ae ee cree Soil. 
Sees s COT tues POET. ANd nie. Pet SO ikl Leaves. 
MeV RMSE SIN CEATITIG) #2. 5 ho por oie eh eyed cere bet rere rt Leaves. 
Smee UP INES SCO DSIRE 1012, ) ns ES i Leaves, ete. 
CEC MRC TESIEIESE UID) ge Mee ok ee ee Leaves, ete. 
PPC UOMICR eae os eA ee eS eR Beans. 
ernie rrr iii) see ee ae 7a ee ees PR Peet Ase Soil. 
EIGUSGLNSUOSUECALUG A! Lene tip! ry Sibi li POLO iis Sel > Beans and manure. 
PACU OMNIS SPLINT TUS pp eet bee ree as eget) pea eects 2 npr oes oo Manure. 


In some instances the sporophores have been minute, owing to the 
small quantity of the culture medium. P 


« Brefeld, O. Unters. aus d. Gesammtgebiete d. Mykologie, 8, 9, 10. 

b Falck, R. Die Cultur der Oidien und die Riickfiihrung in die h6here Fruchr- 
form bei den Basidiomyceten. Cohn’s Beitrige zur Biologie der Pflanzen, 8: 
507-346 (Pls. 12-17). 


“22 MUSHROOM GROWING AND SPAWN MAKING. 


From the standpoint of obtaining pure cultures, the tissue-culture 
method is capable of very general application. Three considerations 
render it particularly important, as follows: (1) When a suitable 
culture medium is at hand, a pure culture may be obtained almost 
invariably from a young, healthy plant. (2) Cultures may be made 
from fungi the spores of which have never been brought to germina- 
tion. (3) Pure cultures are made by direct inoculation; that is, 
dilution cultures are rendered wholly superfluous. In the case of 
Agaricus campestris and other Basidiomycetes, in which the gill- 
bearing surface is protected until the spores are produced, it is pos- 
sible, with the precautions previously mentioned, to obtain the spores 
pure, or practically pure, and at the same time in considerable quan- 
tity. This is not possible with the great majority of fleshy fungi, 
which are truly gymnocarpous. Again, members of the genus Cop- 
rinus are deliquescent, and here it is impracticable to secure spores 
by the spore-print method. In the Lycoperdacez and other Gaster- 
omycetes it has been found that bacteria are frequently present in the 
tissues by the time the spores are formed, and, even if the spores 
could be germinated, direct cultures would perhaps be seldom possi- 
ble. By the tissue method it is only necessary that the plant shall be 
so young that the cells of the tissue are capable of growth and that 
there are no foreign organisms present in the tissue. In this connec- 
tion it may be stated that in the Phallinez, Hymenogastrinex, and 
Lycoperdinez no representative has been germinated, while in the 
Plectobasidinee germination is known only in the case of Sphaero- 
bolus stellatus and Pisolithus crassipes. 

When the natural conditions of germination shall have been more 
definitely ascertained, direct spore-culture methods should in prac- 
tice, perhaps, replace the pure tissue-culture methods in making 
virgin spawn. This would render unnecessary a tedious portion of 
the work, and the process of spawn making would be thereby made 
less expensive. 

A discussion of the respective practical merits of the spore and tissue 
methods would not be complete without reference to the comparative 
vigor, or productive power, of the resulting mycelium. Inthe growth 
of the mycelium no difference could be detected. The writer has also 
grown mushrooms from spawn produced by both of these methods; 
but the results do not indicate that there is any advantage for the one 
over the other. It is believed, therefore, that the processes of basidial 
and spore formation are in this regard relatively unessential, or at 
least do not intensify whatever invigoration may, in general, result 
from mere sporophore production. It is to be expected, perhaps, that 
any and all cells of the sporophore may. be invigorated by whatever 
is to be gained by the assemblage, or concentration, in the differen- 


NUTRITION. 23 


tiated sporophore, of food products collected by a ramifying myce- 
lum. According to the studies of Harper,* Maire,” and others, there 
is no sexual fusion in the case of the Basidiomycetes which have been 
studied. Two nuclei are present in the cells of the sporophore, but 
these are associated conjugate nuclei, and the fusion of these in the 
basidium is generally considered in no sense an act of fertilization, 
but rather a form of nuclear reduction. Maire states that the cells of 
the mycelium obtained by the germination of the basidiospore are 
uninucleate. It has not yet been ascertained when or how the binu- 
cleate condition arises. 


NUTRITION. 


Although Agaricus campestris has been cultivated for so long a 
time, it does not seem that it has previously been subjected to careful 
experimentation from the point of view of nutrition. The belief 
generally prevalent is that the most essential factor in the nutrition 
of the mushroom is the “ammonia” of the manure or compost. 
Again, it is claimed that organic waste products, such as those indi- 
cated, must undergo a process of fermentation, or “ preparation,” 
in order to furnish the necessary nutrients for the growing mycelium. 
This idea, as will be seen later, is merely based upon casual observa- 
tions “in nature,” and it is found wholly erroneous when tested for 
its fundamental worth by the elimination of other factors of the com- 
post environment. 

Growth on manure and other complex media.—Early in this inves- 
tigation it was ascertained that the mycelium of Agaricus campestris 
in pure cultures would grow luxuriantly on fresh stable manure, and 
as a rule upon the same product in any stage of fermentation or 
decomposition. In some instances, undoubtedly, fresh manure may 
contain injurious compounds; somewhat oftener the same is true for 
the fermented product. In some instances it is desirable to dry or 
thoroughly air the fresh manure before use. Fresh manure from 
grass-fed animals is not to be recommended. The mycelium also 
grows luxuriantly on bean stems or pods, on half-rotted leaves of 
deciduous trees, on rich soil, on well-rotted sawdust, and on a variety 
of other substances. It does not grow readily upon peaty products. 

Some of the more promising edible species were cultivated in 
various media in order to obtain an idea of the comparative value of 
these media in furnishing a nutrient to particular forms. It is not 
possible, of course, to base definite conclusions upon results obtained 


a@Harper, R. A. Binucleate cells in certain Hymenomycetes. Bot. Gaz., 
63: 1-25. 1902. 

Maire, R. Recherches cytologiques et taxonomiques sur les Basidiomycetes. 
Bul. Soc. Myc. de France, 18: 1-209. 1902. 


24 MUSHROOM GROWING AND SPAWN MAKING. 


from pure cultures, since the presence of particular foreign organisms 
in the substratum under natural conditions is perhaps quite as im- 
portant a consideration as that of the specific nutrient value of the 
substratum. The following results are, however, suggestive: 


1. Agaricus campestris. 
Leaves—good growth throughout. 
Soil—fair growth, with tendency to become threaded early. 
Manure—good growth throughout. 
Beans—good growth throughout. 
Sugar beet—fair growth, spreading very slowly. 
Potato—slight growth, spreading very slowly. 
Corn meal—slight growth, spreading slowly, soon becoming brown. 
2. Agaricus fabaceus. 
Leaves—very good growth, rapidly filling tube. 
Soil—good growth, but slower than the above. 
Manure—good growth, but slower than the above. 
Beans—yvery dense growth, soon filling whole tube. 
Sugar beet—good growth; somewhat less rapid and abundant than the 
above. 
3. Agaricus villaticus. 
Practically the same as Agaricus campestris. 
4. Agaricus fabaceus var. 
Practically the same as Agaricus fabaceus. 
5d. Bovistella ohiensis. 
Leaves—good growth throughout. 
Soil—growth throughout, but sparse and threadlike. 
Manure—good growth throughout. 
Beans—good growth; appressed. 
Sugar beet—very slight growth. 
6. Calvatia cyathiforme. 
Leaves—very good growth throughout. 
Soil—good growth; quite as rapid as above. 
Manure—practically no growth. 
Beans—good growth, but spreads very slowly. 
Sugar beet—slight growth. 
Calvatia craniiformis. 
Practically the same as above. 
Calvatia rubro-flava. 
* Practically the same as in the other species of this genus, but spreads 
somewhat more slowly on soil. 
9. Coprinus atramentarius. ; 
Leaves—very good growth throughout. 
Soil—slight growth. 
Manure—fair growth, but very slow. 
Beans—very good growth. 
10. Coprinus comatus, 
Leaves-—very good growth throughout; rapid. 
Soil—good growth. 
Manure—very good growth throughout; rapid. 
Beans—very good growth throughout; rapid. 
Sugar beet—very slow growth. 


ba | 


Ce 


NUTRITION, 25 


11. Lepiota rhacodes. 
Leaves—very good growth. 
Soil—slight growth. 
Manure—slight growth. 
Beans—very good growth throughout. 
Sugar beet—very good growth throughout. 
12. Morchella esculenta. 
Leaves—very good growth; mycelium never dense. 
Soil—very little growth. 
Manure—very slight growth. 
Beans—very good growth. 
Sugar beet—good growth, but slower than above. 
13. Pleurotus ostreatus. 
Leaves—yery good growth; rapid. 
Seil—fair growth. 
Manure—good growth. 
Beans—very good growth; rapid. 
Sugar beet—slight growth; very slow. 
14. Pleurotus ulmarius. 
Practically the same as Pleurotus ostreatus. 
15. Polyporus sulphureus. 
Leaves—fair growth; abundant, filling tube. 
Soil—fair growth. 
Manure—fair growth, but very slow. 
Beans—very good growth, rapidly filling tube. 
Sugar beet—fair growth; much lighter mycelium than the above, with 
prompt oidial development. y 
16. Tricholoma personatum. 
Leaves—very good growth throughout. 
Soil—very good growth throughout. 
Manure—growth slow, but eventually good. 
Beans—good growth throughout. 


Plates II, I11, and IV show some of the more important of these 
species. 

Taking into consideration the variable quality of the stable manure 
which may be obtained at all seasons, the value of half-rotted decid- 
uous leaves as a substratum for Basidiomycetes is worthy of special 
emphasis. The writer has found such material more readily sterilized 
than manure, and usually more prompt than the latter to give growth. 

In order to test in pure cultures the probable effect of fertilizers 
as indicated by any marked increase in the rapidity of growth of the 
mycelium, experiments were made by adding a small quantity of 
ordinary nutrient salts to test tubes containing manure. A chlorid 
and a nitrate of the following salts were employed, viz, ammonium, 
calcium, magnesium, and potassium. In addition, dibasic potassium 
phosphate and also sodium chlorid, as well as control cultures, were 
used. Three tubes were employed with each of the compounds men- 
tioned. There was no marked difference in the amount or rapidity 
of the growth noted, as found by comparing the averages of growth. 


26 MUSHROOM GROWING AND SPAWN MAKING. 


It seemed possible, however, that some slight advantage resulted from 
the calcium compounds, but there was no pronounced benefit in any 
tube. Further reference is made to the use of nutrient salts in mush- 
room growing in another chapter. 

Growth on chemically known media.—tIn an attempt to determine 
somewhat more accurately the value of different compounds as 
nutrients, particularly carbohydrate and nitrogenous substances, sev- 
eral series of extensive tests have been made with Agaricus campes- 
tris, and also with Agaricus fabaceus and Coprinus comatus. These 
fungi do not grow readily in liquid media, and it has been difficult 
to obtain a wholly reliable and satisfactory substratum, one which 
would itself be practically pure, or well known, chemically, and at 
the same time effective for its purpose. After unsatisfactory at- 
tempts with various gelatinous solid media, with charcoal, ete., it was 
decided that the commercial gray filter paper had more to recom- 
mend it than any other substance suggested. Accordingly, all ex- 
periments were made in Erlenmeyer flasks of 150 c. ¢. capacity, and 
in each flask was placed about 6 grams of this paper wadded into 
pellets. The latter was moistened in each case with the nutrient 
solution used. The flasks were subsequently sterilized in the auto- 
clave and then inoculated with a very minute fragment of straw 
with the fresh mycelium from a pure culture on manure. 

Tabulation of special results —In the following tables are given 
the results of two out of several series of experiments, which have 
been conducted in order to throw some dight on the point just dis- 
cussed. These tables include, also, many cultures on media of un- 
known composition. 


TABLE V.—fesults of growth on media—First series of experiments. 


No. Medium. | Extent of growth. 
/ : 
ib He ae Wieteadrs are, saiee ab hae hc ee a "Bo. #* 
| meepemoner aise tan L& 
ab fer A g: inl sugar, 1} per cent - wane ne ne seen eee eeeeeeee \ 7. Do. 
eee s WEEN mr 
bb i Muon A apd laGese, Li Per CEN. .. 256s ooenk a ecewcce nse ceeeee \ Do. 
6b f LRCLOBS, Ld DOE COWS oncus achenn'< aac oneness sagen cokes ee oe ) Do: 
nb {Solution A and pi@erin, 12 pet Cont... ..<qsc0ca.) sss, 0kcgeuee { a 
on )Glycerin, LE CEI s Feo = ew ta aan ated anal opiate ae J 
on }Solution A and starch paste, 4 per cent. .... 22.5. sc ib cee ee eee | a aha 
7 {Starch PASbG FOr CONG Se. cen ccncitrk sade seine ed pdee aan Oa 
= \solution A and starch, } per cent, and diastase, trace -.._.---- ee | discarded. 
7D {Starch, + per cent, and diastase, trace - ne ener eee eis 
- Solution A and Gextrose, 1) Per CONG. wo ewe essen denn acunnd bie pe 


* Solution A.—Water, 2,000 cubic centimeters; MgSO,, 5 grams; KsHPO,, 10 grams; 
NaCl, 1 gram; CaCl, 0.25 gram; KNO;, 20 grams; FeCl;, trace. 


NUTRITION, 


27 


TABLE V.—Results of growth on media—First series of experiments—Continued. 


No. Medium. Extent of growth. 
i \Dextrose, PPE POM CGI i 5 che a3 eee reine a/R ee ee ye: Heroes ¥ 
cs fSolution A and mannite, 1} per cent__....--.-...-..-.---.------ Very slight. 
16a |\y,7. . 
16b | Mannite, PPSVeISCOM G26 don ah yo eee a ns Saree at tS Lost. 
lia Contaminated with Asper- 
» Vib }Solution AVandimaltose; ie percents: 22225. 2222S te ee peeks Fair growth, yel- 
wish in color. 
Eos |Maltose, CER OIE oer hu ne Ee alas setae yu ateaneausass= art 
ts }Solution A and potassium tartrate, } per cent_________________- {ouere. 
ae }Potassium tartrate, + per cent - crease wiapaaies seas ae { me 
kts }Solution A and magnesium tartrate, } per cent_.__......------ { a 
2 | Magnesium ini ebruees 2 per Cento. ——sepe ces Se Th? URE 3h { Bp. 
ears \Sotution A and ammonium tartrate, } Wer COMun o.--2 a5 scene { ae 
24a * : Do. 
24b }Ammonium EEA Or COGN = aes eae one las oe oer et { Bie 
ae \Solution A and potassium lactate, } per cent.-__....---.-------- { ee 
2b \Potassium LAG ianCMGL COMG on52..06e5s sone se astee oe re ne {sieht} tovfair, 
an }Solution A and magnesium lactate, } per cent -_--.-.---------- joes 
ae |Magnesium PIC beR Er DCR CONU Mae <2 9a o oe nee oe oe an = oa i nas 
oo Solution A and ammonium lactate, } per cent__..________--.--- iat ES good. 
a MAMI IACLALG. 1 per CONG. -—... .-.2.5-2--0- -202----2-—-2-< (endo 
a \Solution A and calcium hippurate, } per cent__...._2....---.-- = ened 
Ps \caleium etn DELO POr COIN bs 5-1 2.26. 2255 sete sae tos ce J olen 
3b }Solution Pang asparagin-.t DemCent.=—--— 4 ss-ccssa-dee4an45--2 { he 
3b \Asparagin, “ATER CAST Repeat ek Malpelo nao kd 1a a J ae 
oe \Solution Sean pepoong. + Der COW... .--5 oss Seats wagence een k lage 
ba \Peptone, Lu [OVSES a7 DUET, 2 RR cree Se EY Soh Cr pela ad ee deat, slight. 
pe {Solution PNAC Hay G CRTS(S Ta PNS Bh of ss Gye) a ae a a J very ss good. 
38a, f Do. 
38b }Casein, 4 POUL COU hee. 2h opens a5 Saws ae ns fu Bi 2 oe Pe ee ae 1. Do. 
ae }Solution MARSDEN, + POM CONbt= = 2.2... ass seasenen sds ln =e ponerse fair. 
‘ob \Pepsin, AGRI @SUaER cc tape eae bape seal ep es teh dah ae eae { De. 
a : 0. 
Alb jSolution FES eR ee en 8 Oe ah a rah nek aes mee ste is Do. 
io Solution B and asparagin, } pey cent -_.-...........-----.------ peer 
Ass Solution B and peptone, } per cent ___.....-.._--.---2-.-.----.- Nite slight. 
a }Solution AA GeCAS OI. 2 POl CONba. 22 occ seo ne ee Wess good. 
Ps \Solution Eandenepsin: 4 percent... <=. 1) os eee Cultur e lost. 
46a, ‘ Slight to fair. 
46b \Bouillon BEMPSE NO OTE ice oe Sas eae ae ne os eee ete aD 
rs \Bean (SkSCOYEAFCos papa NO Daal Ree Ame De abe 2. [ey cs te ie a More es sod. 
ae \Beet FRG geet SOT ROS ETE RODE So 31 ie ay Gace po WarY, good. 
ec {Manure GEICO), a a Ae et Meme vege ‘good. 
ae \Manure SEER Be ee See en ee Serna eee me ett ew Bea D Bo. 
PmMMPOOU TIS Tray Set ooo ac onan os ceew co oee eee ele ost. 
Lo BainwWonsA andwinheat Straw —._-/.2-.---. 5---2- cose ese -caee- { Be 
bos \Solution en NVOE Aa SUL AW 2 aoe foe a eo Sanh cet Ree e { nt ’ 
oa \Solution B and NH,NO; and cane sugar. --......----.---------- eames to fair. 
= \Solution B and cane sugar and Ca(NOs)s, $ per cent..________- wat slight. 


« Solution B.—Water, 1.000 cubic centimeters; MgSQ,, 2.5 


CaCls, 0.25 gram; NaCl, 0.5 gram. 


grams; K.,.HPO,, 5 grams; 


28 


MUSHROOM GROWING AND SPAWN MAKING. 


TABLE V.—Results of growth on media—First series of experiments—Continued. 


Medium. 


Solution B and sugar and Mg(NOz)o, + per cent 


}Solution B and sugar and NH,Cl, } per cent 
}Solution B and NH,NO;, } per cent 


Solution B and Ca(NOz)s, + per cent 


Solution B and Mg(NOsz)o, } per cent 


Solution’B and NHAC, + per cént. 2 222.5. Ss. .22. tee 


Mushroom decoction 


} 
Extent of growth. 


TABLE VI.—Results of growth on media—Second series of experiments. 


Medium. 


\Presh horse manure (grass-fed animals) 


Fresh horse manure, thoroughly washed, residue only used-.- 
iltrate, or liquid resulting from washing No. 2_-_--...---.---- 


Decoction of fresh horse manure, as in No. 1 


ermented horse manure, thoroughly washed----.-- Sarr ee 


iltrate or washing from No. 5 


Rotted stable manure 


bP 
i 
Pe 
P 


ecoction of green timothy hay 


pBeridne from decoction in No. 8 


{Strong D@an G0ICe ha sa oo a oe lea eee ee 
jWeak DEAD JUICO. eo). 22k ek on 8 ee eee 
{Strong decoction ofmushrooms ..-5--- =. -eenees es no ee 
{One-half strength decoction of mushrooms ---_---.-----~------ 
}Weak decoctioniof mushrooms .; =. ..2-.25 2202s 22 ae 
jOat straw 


ila straw 


f 
ie 
\, 
j* 
\y 
{" 


\Corn meal 


gram cane sugar in 25 c.c. solution A 


ram milk sugar in 25 c.c. solution A 


gram galactose in 25 c.c. solution A 


My gram cornstarch in 25 ¢c. c. solution A 
} strength albumen (egg) 


My gram glucose in 25 c. c. SOU GOD. Ao. Ane courier { 


M4 gram dextrose in 25 c. c. solution A 


SOMMTION A. nan namin dae dae 


gram mannite in 25c. c. 


} gram glycogen in 25 c. c. solution A 
hy gram maltesent 25 0. c. solution Acuu.:. scot 
ie gram levulosein 26.6, c. solution: A. siiaocesc.. Sobed ene sn one ns 


pi gram glycerin in 2c. c. solution A 


Extent of growth. 


None. 
Slight. 
Contaminated. 
Good. 


od. 
\Contaminatea. 


(shen. 
Contaminated. 
Good. 


1 
J oo 
istient, 


Contaminated. 


Slight. 
Do. 
loornaate. 


\fSlig ht. 
\Confined to nocules. 


/\Siiees throughout. 


Slight, but contaminated, 
(Slight at top. 
'\Lost. 
Confined to nocules. 
ope at top. 


ieee throughout. 
\{Fair. 

HP ah. onbags 
ey contaminated. 
Slight at top. 

| Do. 


Slight. 
Slight at top. 


Confined to nocules. 


NUTRITION. 


29 


TABLE VI.—Results of growth on media—Second series of experiments—Cont'd. 


Medium. 


Extent of growth. 


4 oid flake spawn 


| 
j 
j 
j 


1 gram urea in solution A 


+ gram peptone in solution A 


+ gram casein in solution A 


\ gram potassium lactate in 25 c. c. solution A 


lh gram magnesium tartrate in 25 c. ce. solution A 
} gram calcium hippurate in 25 c. c. solution A 


i gram asparagin in solution A 


\ gram potassium tartrate in 25 c. c. solution A__....---------- 


+ gram potassium lactophosphate in 25 c. c. solution A-------- 
My gram magnesium citrate in 25 c. c. solution A_--..._--------- 


1 gram magnesium malate in 25 c. c. solution A__-------------- 4 


bh PianCaselt 11) Aer GC. C. BOMMGION Ib — =. 22-2. snk one eee ee ees seems 


| Asbestos with bean decoction 
| 


f pine sacle contaminated. 


oO. 
grees to nocules. 
Do. 
Do. 
Do. 
Yeuent at top. 
O. 


G at 
ood top. 
D 


0. 

sSlight. 

\ Do. 

{ Confined to nocules. 
0. 

{Fair at top. 

{" Do. : 

sPair throughout. 


( Pratnpenrz0le Aclo 1 SOlUbION A _----------.----2---2-- ce cen ee Is one. 

tt Pram benzoic acid im'solution, At? 5222-. 28. £52 so eS { et 

(a a aim tae Siegen nade eenpemace rai {Confined to nocules. 
{Solution 1B ccs SS pe Re ce ll a ar ae Rte Mi te { _ 
(EU glare jell satan ate te ae - 

|Decoction ELOMUCOCuCLiVveOld node ee eee acta 
}Oak Bawaust, only sheitly rotted'....---------- 2. es-e nate to nocules. 
}G@luten PiPeceeET EW it ese ee ee eee ere eee 
}Cotton-seea TERN GR ALeIN eee ae eee eee teen aig capa 
\Cotton-seed Sl yuh mecegshte pes at dled - Ne A i. spacer Mal J cee moire 2g 

\i Pram aaparaein. imsolution Bl...) ese ee eee haga top. 

\ Pa AAS PATRI I SOMLUION Bs... ---- 2202 Aan Meee aan ae a oo Ae: 

hi erica en sOlinOnes foe eee eh ee poe 

\ PranMr eat SOON Ere. te ee agree ye top. 

hs PraAnnnres im solution G 22.20. 9 — sae ee ee dais ngage pis 
hh gram peptone in 25 c. c. solution B_--.--_---- eee eee oe top. 

he gram peptone in 25 c. c. solution B.__...-..--.-- cgeidi nt Sn li { rare 

ie gram peptone in 25 c.'¢c. solution B_-.....-....._-.----------- i —_— top. 

\ gram peptone and yj, gram NaNO; in solution B.__....__.._- mee 

Ite gram casein in 25 c. c. solution B_._.....-.-...---------------- wa’ peers 


{Fair at top. 
\Slight at top. 


\ is gram casein in 25 c. c. solution B.__..-.--..-..-.------------- co 
hh PCE Tal DumMoe (eee) cnn ae ee ee ee ome ome top. 
joa REST ITO Wy LUCE re es oe ee ee ees aga gipaeeienes 
jMers small area, but copi 
| white TREC re ee ME a Sep el See eee Oe. 
} : oO. 
white pine shavings with bean decoction.-..-.....-.---------- { Da 


oO. 
yRenpeed to nocules. 
Do, 
(Very good. 
Do. 


30 MUSHROOM GROWING AND SPAWN MAKING. 


It is not possible here to enter into a detailed discussion of the 
results, but attention is directed to the fact that under ordinary con- 
ditions Agaricus campestris does not give a copious growth when 
nitrogen is furnished from an inorganic salt and carbon in the form 
of the well-known sugars. Calcium hippurate in a solution of the 
necessary salts has almost invariably given better growth than other 
organic salts and carbohydrates. In general, casein has been a better 
source of carbon, or of carbon and nitrogen, than other proteids. 

When the manure is of good quality it furnishes, in pure cultures, 
a source of necessary nutrients, whether fresh or fermented, whether 
as a decoction or an infusion (a cold aqueous extract). 

Acid and alkaline media.—Manuure which has undergone fermenta- 
tion for a few weeks is usually slightly acid in reaction. Under 
certain conditions of fermentation the acidity is increased, and this 
is probably an important factor in making the manure from animals 
fed with green foods less valuable for mushroom work. Some acid 
tests were made of beds which had failed to yield satisfactory results, 
and in many instances it was found that the acid content was much 
above the normal. A small series of experiments was therefore 
instituted to determine the relative amount of acidity or of alkalinity 
most favorable for the growth of the spawn under pure-culture con- 
ditions. In this test there were also included several other edible 
fungi, the results of all of which are included in the table below. 
These experiments were made in large test tubes, and in such a test 
it was impracticable to determine absolute acidity or alkalinity, and 
from the results only a rough qualitative comparison could be antici- 
pated. Potassium hydrate and lactic acid were used as reagents. 
The duration of the experiments was one month, and duplicate cul- 
tures were used in every instance. 

Although the results are not wholly uniform, it may be inferred 
that in the case of Agaricus campestris a marked acidity of the 
medium would be unfortunate; Calvatia cyathiforme, on the other 
hand, seems to have grown somewhat better, in general, in the more 
acid media; Coprinus comatus grows under a wider range of condi- 
tions; and Coprinus atramentarius, in this instance, thrives in an 
alkaline medium. Further tests on a quantitative basis are required 
before definite conclusions may be drawn. This matter will also 
receive further attention when facilities are at hand for undertaking 
to better advantage than has yet been possible the practical growing 
of the other species, besides Agaricus campestris, included in this test. 


TEMPERATURE AND MOISTURE, ae 


TABLE VII.—Results of tests of acidity and alkalinity. 


Extent : 
Nature of xtent of growth | 
Medium. chain Agaricus | Calvatia cya-| Coprinus co- |Coprinusatra- 
compost. | campestris.| thiforme. matus. mentarius. 
Fresh ....| Very slight | Very slight --| Slight---...--| 1 See very 
slight. 
4 drops KHO -.......-.-..- Rotted -._| Slight ------|--.-- (3G tlelas Pyet Very slight -- i amina- 
ed. 
Fresh _...|1 good, 1|}1 none, 1| lverygood,1| Very slight. 
2 drops FAO yr Sye5-20 25 3- fair. slight. excellent. 
Bones oon eee good - aoe aa bs I ce ues ea = 
resh __..| Good-_------ ery slight __| Excellent --..| Very good. 
1 drop KHO -.----.-.-....- Rotted _-.| Very good -| None-_.-------}----- ages. Excellent. 
Fresh -...| lvery good,| 1 good,1none |-_---- iYo eo ere Very slight. 
Control | : 
<7 33550 te re Rotted _._| Veryslight Heat na- | None-_-_....._-| None. 
| ea. 
Fresh __._| 1 contami- 1 slight, 1} Excellent ---_| Very slight. 
Edrop acio..:-.-.----.--:. | 2 2 a4 Sele 
. Rotted .._| Very slight | Wee sligittues|=7-2> dove Do. 
Maroosacid oer ay | eso goes 23 SRA OE sibs oe Gg ts. Do. 
Ls: lta |(Rotted ---]_---- ado'=.-- Very good -._| None_-------- Do. 


TEMPERATURE AND MOISTURE. 


The temperature factor is, next to that of good spawn, perhaps the 
most important in mushroom growing. It has been frequently 
stated that mushroom growing is not profitable when the temperature 
may not be maintained more or less continuously at from 50° to 60° F. 
It is very probable that the exact temperature which may be con- 
sidered an optimum will vary somewhat in different sections of 
the country. It will be noted later in detail that the temperature 
factor acts not so directly upon the growth of the spawn or the produc- 
tion of mushrooms as indirectly to render some other conditions of the 
environment injurious. It is best to consider that in practice the op- 
timum temperature for mushroom growing varies from 53° to 58° F. 

When the matter of temperature was first under consideration, 
a series of pure cultures of Agaricus campestris was placed at 
different temperatures in the laboratory in order to determine the 
rapidity of growth. It was soon found that a temperature above 
60° F. and, indeed, as high as from 80° to 85° F., was much more 
favorable to rapid growth than a lower temperature, provided, 
of course, that the higher temperature did not encourage a too 
rapid drying out of the culture. It was soon definitely ascer- 
tained that the conditions of pure-culture growth are essentially 
different from those attending the growth of mushroom spawn in 
the bed. This was perhaps best indicated by comparing spawn 
grown in pots at 85° F. under impure conditions with similar 
spawn grown at 50° F. At the former temperature, even though 
the conditions of moisture were properly maintained, there was 
little or no growth. Foreign fungi, molds, and bacteria, as well 
as insects, were, however, abundant. At the lower temperature there 
was little or no evident appearance of other fungi, molds, er insects; 

23994—No. 85—06 m——3 


32 MUSHROOM GROWING AND SPAWN MAKING. 


yet the mushroom spawn grows slowly and continuously so long as 
other conditions are maintained. From numerous experiments of 
this nature it is apparent that the temperature relation is one which 
is governed by the competition to which the mushroom spawn is sub- 
ject in the bed. ‘This is, of course, wholly in accord with the results 
obtained from the study of the relative growth made by mushroom 
spawn in fresh and composted manures. 

The statement previously made, therefore, that the optimum tem- 
perature may vary slightly in different localities is true on account of 
the fact that the mites, insects, and other animal pests of mushroom 
growing may vary considerably in different localities, or under dif- 
ferent conditions, even though there may not be a great variation, 
perhaps, in the bacterial and fungus flora of the, compost upon 
which the mushrooms are grown. Certain insects, for example, are 
more abundant in a moist climate, but if special precautions can be 
taken to eliminate all such pests, the growth problem is confined to 
the interrelation existing between the mushroom spawn and the 
microscopic flora of the compost. Mushrooms grown in the open 
will probably show greater variation with reference to the tem- 
perature factor than those grown in caves or cellars. 

While a number of interesting problems would be presented by a 
study of the interrelation of the mushroom mycelium with that of 
other microscopic fungi present in the compost, these are matters of 
detail; and it has been wholly impossible thus far to give any atten- 
tion to suggestions which have been furnished by the experimental 
data. It may be possible that other species of mushrooms are more 
independent of insects and other microscopic fungi, and such fungi 
may therefore be more suitable for cultivation at high temperatures 
than is Agaricus campestris or any of its close allies. A considerable 
effort is being made to obtain spawn of certain species of Agaricus, 
and also of other edible mushrooms which make their appearance 
during the warm weather. At this time, however, it is not possible 
to say what results of value may be anticipated from this line of 
work. 

The direct effect of a temperature above the optimum upon the spo- 
rophores is manifest through lengthening of the stipes and rapid 
expansion of the caps, ordinarily accompanied by toughness and 
decreased size. In other words, the lower grade market product is 
produced at the higher temperature. 

The moisture factor is also one of importance. It is undesirable 
that the place in which mushrooms are grown should be very damp, 
or dripping with water. Nevertheless, a fairly moist condition of 
the atmosphere should be maintained throughout the growing and 
productive period. There should be a gradual but slight evaporation 
from the surface of the beds, and sufficient ventilation to insure this 


PREPARATION OF THE COMPOST. 83 


is believed to be essential. It is certain that in poorly ventilated 
caves mushrooms do not succeed. On the other hand, in a dry 
atmosphere, or exposed to drying winds, mushroom beds soon cease 
to bear, while such sporophores as are developing may have their 
caps cracked and torn. 

Mushrooms are grown in cellars, caves, or specially constructed 
houses largely on account of the fact that temperature and moisture 
are then practically under control. The nature of the structure or 
cellar which is constructed for mushroom growing must be deter- 
mined, therefore, not merely by its expense, but by the effectiveness 
of the structure in regulating the factors indicated under the par- 
ticular climatic conditions. 

It is not possible at this time to discuss cellar or house construc- 
tion, and the accompanying illustration of mushroom houses (Plate 
VI, fig. 1) must suffice to give an idea of the types which are in use. 


PREPARATION OF THE COMPOST. 


It is not to be understood that there is one and only one method of 
preparing compost for mushroom growing. Nor is it always .neces- 
sary that the compost shall be in one particular stage of fermenta- 
tion or decay. In fact, every change of condition elsewhere may 
necessitate a similar change in the amount of fermentation which 
may be most desirable. At the outset it should be understood that 
it is not the “ fermentation” which is absolutely essential. The 


a@Répin, 1. c. (See translation in The Garden (London), February 5, 1898. 
Special reprint, pp. 10-16.) Heré it is stated that “manure is rendered capa- 
ble of supplying nutriment suitable for mushrooms only by means of fermenta- 
tion;”’ further, that “all the higher orders of mushrooms, the spores of which 
I have succeeded in causing to germinate, have a sterile spawn of a similar 
nature.” Again, the conclusion is expressed somewhat indefinitely that manure 
is “rendered suitable” by means of chemical combustion, which is said to 
proceed rapidly only at a temperature above 178° F.; that it is not the soluble 
substances in the manure which are valuable, but rather the cellulose matter, 
together with the necessary salts. 

In this connection it is of interest to note that the material constituting many 
of the beds in the experimental cellar at Columbia, Mo., were fermented at 
comparatively low temperatures. A complete temperature record was kept of 
18 small compost piles in which special kinds of manure were prepared, 
and in only one instance was the temperature in any pile more than 140° F. 
In some cases 120° F. was the maximum attained, 

Répin implies that mushrooms will not grow in manure until there has been 
effected “the destruction of all the soluble organic matters, which disappear 
through the agency of bacteria or are consumed in the process of oxidation.” 
Very simple nutrition experiments clearly demonstrate that these conclusions 
are erroneous. 

It may be stated, however, that peculiarities appear when the fresh manure 
contains certain compounds which render it injurious; for example, the my- 
celium does not grow readily in pure culture upon fresh manure from animals 
fed almost wholly on green forage. Such manure is improved by fermentation. 


34 MUSHROOM GROWING AND SPAWN MAKING. 


“ fermentation ” is of itself a minor matter. In pure cultures, where 
sterile media are employed, mushroom spawn starts slowly, but 
finally grows best, in general, upon fresh (wholly unfermented) 
manure. It grows least well, or, rather, less densely, so far as tested, 
on very well fermented manure. This certainly indicates that it is 
not fermentation which is ordinarily advantageous. In practical 
mushroom growing, however, it is not possible to deal with pure cul- 
tures; and, therefore, other conditions of the environment must be 
correspondingly changed. The rapid oxidation action of bacteria, 
and perhaps of independent ferments, upon manure causes a consid- 
erable rise of temperature. At the higher temperatures (which 
may be maintained as long as there are present rapidly oxidizable 
food products) bacterial action is vigorous, and is unquestionably 
injurious to mycelial development. Wholly aside from the rise of 
temperature accompanying their activities, bacteria are otherwise 
injurious. In fact, manure which is put to ferment in a small test 
tube shows little or no rise of temperature above that of the place in 
which it is incubated. Nevertheless, the mycelium of the mush- 
room will not grow under such conditions. Rapid bacterial action 
is therefore prejudicial. Under those conditions where bacterial 
action is not rapid, fresh manure might be used to advantage; in 
other words, if the beds are so constructed that’ the manure ferments 
very gradually, without either excessive bacterial action or rise of tem- 
perature, then spawning might be made in fresh manure. 

The old belief that rotten manure does not have the necessary 
strength—that is, does not produce so .vigorous a mushroom growth 
as that which has been less transformed by bacterial action—has been 
confirmed by practical experiments. This loss of effectiveness is 
probably due, in part, to a change in texture or to other physical 
changes. In well-rotted manure there is ample food material to sup- 
port a very good growth of mycelium in pure cultures. This has 
been chemically proved by sterilizing such manure and growing mush- 
room spawn upon it in pure culture. Nevertheless, by comparing 
(in Table VIII) No. 12 with Nos. 13, 14, and 15, it will be seen that 
beds prepared with well-fermented manure and left for some time 
before spawning do not yield so well, It is believed that here the 
physical condition has much to do with the result. 

The latter does not by any means invalidate the following practice, 
which has commended itself to some very successful growers: The 
manure is piled in very large compost heaps, where it is kept moist 
and is turned only once or twice. It ferments very slowly. Then it 
is carted into the cave, or mushroom house, long before it could be 
considered in proper condition to be spawned. The beds (usually 
flat when this is the procedure) are made immediately. These are 
fairly well moistened and compressed, then left to undergo a gradual 


PREPARATION OF THE COMPOST. 35 


fermentation, which may require a month. When the manure shows 
a tendency to fall to the temperature of the room it is spawned. 
Meanwhile, it will doubtless be found that a heavy crop of some 
small species of Coprinus will have appeared. The presence of this 
fungus is not injurious, but rather it may be taken as an indication 
that the conditions are favorable. 

Ordinarily the manure is obtained as fresh as possible. It should 
include the straw used in bedding the animals, and the quality of the 
straw will determine-to some extent the value of the manure. The 
straw of cereals is far better than that of most other grasses. The 
more resistant straws seem greatly to improve the texture of the com- 
post for mushroom purposes. Commercially it is a mistake’ to 
attempt to get the manure free from straw. If fresh manure is not 
obtainable, that which has been trampled by the animals is ordinarily 
rich, well preserved, and desirable. It ferments best in large piles, 
and these may be of considerable extent, about 3 or 4 feet deep 
throughout. If not uniformly moist the material should be sprink- 
led. At no time is a very heavy watering desirable. In from four 
days to a week or more the compost should be turned, or forked over, 
and a second turning will be required a week or ten days later. 
Water should be added only when necessary to maintain a moist (but 
not a wet) condition. With this amount of moisture, and with the 
piles deep enough to become fairly compact as a result of their own 
weight, there will be little danger of any injurious fermentation. 
During the normal fermentation the temperature may rise higher 
than 150° F. In from fifteen to twenty-one days or more, depending 
upon the conditions, the temperature will begin to fall, and the com- 
post may be used in the construction of the beds. When used in the 
beds, it has ordinarily lost all objectionable odor, and the color of the 
straw has changed from yellow to brown. In figure 2 on Plate V 
is shown a shed in which the manure is composted during the summer. 

As stated in Farmers’ Bulletin No. 204: 

It is the custom with some growers to mix a small quantity of loam, about 
one-fourth, with the manure. This enables one to use the manure earlier; and, 
indeed, under such circumstances it may sometimes be used with but little or no 
composting. Nevertheless, the majority of growers have obtained greater suc- 
cess by the use of the manure alone, and this is also the writer’s experience. 
Very well-rotted compost should not be used in mushroom growing if large and 


solid mushrooms are desired. When sawdust or shavings are employed for 
bedding the animals, the composting may require a somewhat longer period. 


It has been the experience of some of the most successful growers 
that the use of shavings for bedding material in the stables does not 
injure the value of the product for mushroom work. The presence 
of a large amount of sawdust is, however, objectionable so far as the 
writer’s experience goes. Compost containing much sawdust is 


36 MUSHROOM GROWING AND SPAWN MAKING. 


necessarily very “short,” and therefore the physical condition is not 
the most favorable for Agaricus campestris. 

In another chapter attention is called to the fact that the value of 
the manure depends to a considerable extent upon the feed given the 
animals. It would not be wise to depend upon that obtained from 
stables in which hay and green foods are used to too great an extent. 
Moreover, it is not believed that compost made from the manure of 
cattle barns is in mushroom growing as desirable as stable manure. 

In some cities the municipal ordinances require that the manure 
shall be promptly removed from the feeding stables or that it shall 
be disinfected. In the latter case crude carbolic acid, or even corro- 
sive sublimate, may be used to secure this end. Manure thus disin- 
fected is, of course, undesirable for mushroom work. For the same 
reason the manure of veterinary hospitals is-of questionable value. 

It is not wholly improbable that some other waste products of the 
farm, field, and forest may be utilized in mushroom growing; never- 
theless, no such product has yet been found which, under the condi- 
tions of the experiment, has yielded sufficiently to make it of special 
interest in growing Agaricus campestris. Among the products which 
have been tested, either alone or in conjunction with some commer- 
cial fertilizer, are the following :. Leaves of deciduous trees, needles of 
conifers, sawdust, cotton-seed hulls, cotton seed, corn stover, sorghum 
stover (or bagasse), rotten hay, sphagnum, and yeddo fiber. The 
writer is convinced that greater profit may be anticipated, for the 
present, at least, if the culture of Agaricus campestris is confined to 
imanure; and if other edible forms which grow in the woods are used 
in beds of leaves, etc., as indicated elsewhere in these pages, it is quite 
possible that such a fungus as Coprinus comatus may be grown suc- 
cessfully in this latter way. It may, however, be too much to hope 
that the morel may also be thus made amenable to culture, although 
leaf mold is in nature the favorite habitat of this fungus. 

From the prompt and abundant growth of Agaricus campestris on 
half-rotted leaf mold in pure cultures, it was thought that mushrooms 
might be grown to advantage upon this product. The practical 
experiments made to test this point are distinctly discouraging, as 
shown by reference to No. 17, Table VIII; Nos. 3 and 4, Table IX, 
and No. 11, Table X. 

For the most part manure may be composted in the open air. It 
may, however, be prepared with greater uniformity under cover. 
During midsummer, protection may be desirable on account of dry- 
ing out, while in the winter it is more important in case of excessive 
cold. If it is necessary to compost manure during the winter, more- 
over, the piles should be of considerable depth. 


INSTALLATION OF BEDS. 37 


INSTALLATION OF BEDS. 


In making the beds, as well as in other phases of mushroom work, 
regard must be had for all environmental conditions. The type of 
bed should be determined by convenience, and the size, to a cer- 
tain extent, by the temperature to which the beds may be exposed. 
The flat bed, frequently referred to as the English type, is more 
commonly employed in the indoor work in England and America. 
With this type merely the entire floor space may be utilized, as illus- 
trated in the frontispiece, Plate I, or the beds may be arranged in 
tiers of shelves. In figure 1 on Plate V a view may be had of the 
supports for shelf beds in a large commercial house. In this house 
there is the greatest economy of space. The shelf system gives the 
greatest amount of bed space and is certainly most economical where 
the floor space is an important factor. Such beds do not require 
great depth, but merely sufficient to insure an ample development of 
spawn. They should be from 8 to 10 inches deep after being firmed 
or compressed. 

The ridge-bed system is employed almost exclusively in the caves 
about Paris. This system is also in use in open-air culture. It may 
be used to advantage in low cellars, caves, or houses when labor is 
not too expensive. Ridge beds increase slightly the surface area and 
permit of easy passage from one part of the cave to another. The 
size of such beds in caves, or under other conditions where the tem- 
perature remains practically uniform, should be not more than 2 feet 
wide at the base and 15 inches high, tapering gradually to the top 
when compressed. Slanting beds are commonly employed next to 
the walls. Large beds are desirable under changeable open-air 
conditions. 

The prevalent opinion among amateurs that the bed should always 
be deep enough to maintain a considerable heat is believed to be erro- 
neous. Grown under more or less uniform conditions, mushrooms 
seem to require no bottom heat, and the bed should fall to the tem- 
perature of the room some time after spawning. Bottom heat, and 
hence large beds, are, however, desirable when sudden changes of 
weather would so reduce the temperature of the bed as to delay 
growth. Under similar conditions, as well as in dry air, mulching 
may be required. 

As previously stated by the writer in Farmers’ Bulletin No. 204 
of the Department of Agriculture— 

In any case, the manure is made up in the form of the bed desired and should 
be firmed, or compressed, to some extent immediately, in order to prevent dry- 
ing out and burning when the secondary fermentation takes place. At this time 
the manure should be neither wet nor dry, but merely moist. The only prac- 


tical test of the proper moisture content of the manure which can be relied upon 
is when, upon compression, water can not readily be squeezed out of it. 


38 MUSHROOM GROWING AND SPAWN MAKING. 


SPAWNING AND CASING THE BEDS. 


From what has been said concerning the temperature requirements, 
it will be evident that spawn should not be inserted in the beds until 
the temperature has fallen low enough to insure successful competi- 
tion on the part of the mycelium with other organisms. In many 
urticles on mushroom growing it has been suggested that beds may be 
spawned when the temperature has fallen to about 90° F. From 
experience and observation, the writer can only conclude that such a 
temperature is frequently fatal, and it is believed that the tempera- 
ture of the beds should be permitted to fall to 70° F. before being 
spawned. In fact, the most successful results have been obtained at 
temperatures from 65° to 70° F. It was formerly believed that if 
the spawn were inserted at 90° F. this higher temperature incited the 
rather dormant mycelium to rapid and vigorous growth. It is clear, 
however, that the rapid development of new mycelium from the pieces 
of spawn brick inserted is not so important a factor as suitable 
conditions for continued growth. If the temperature falls rapidly 
from 90° F. after spawning, however, no injury may result. Never- 
theless, it is to be considered an unfortunate condition. 

The bricks of spawn may be broken into from ten to twelve pieces, 
from 14 to 2 inches square. These pieces may be inserted about 1 
inch beneath the surface of the manure. In flat beds they may be 
placed from 10 to 12 inches apart throughout the bed, and in ridge 
beds the pieces should be inserted on each side alternately, one near the 
top and the next near the bottom. It is well to insert the pieces 
vertically, as the mycelium does not then seem so readily to suffer 
damping off. After spawning, the beds should again be firmed, and 
they are then ready to be cased or loamed whenever this process may 
seem most desirable. At the time of spawning the beds should be in 
the best condition possible for the growth of the mycelium. Delay 
in growth at this time is one of the surest indications of a light yield. 
If the bed contains the proper amount of moisture, and if the walls 
and floors of the house or cellar are sprinkled occasionally, so as to 
maintain a moist condition of the atmosphere, it is possible to avoid 
wholly the use of water upon the beds immediately after spawning. 
In no case should a bed recently spawned be heavily watered. ‘The 
surface may be sprinkled, if there is a tendency toward drying out. 
The same test for moisture content as has been outlined previously 
in these pages in the chapter on preparing the manure should be 
followed. The beds should become gradually somewhat drier, how- 
ever, during the growth of the spawn. 

The absolute water content for the bed at the time of spawning 
should be about 40 per cent, although this will vary considerably, 
according to the conditions, and especially with relation to the 
quantity of straw in the manure. 


MUSHROOM GROWING. 39 


If the spawn grows rapidly at first and spreads throughout the bed, 
it will not be injured by a slight drying out, or by a temperature even 
as low as 32° F. On the other hand, a continuous high temperature 
for several days, or excessive watering, is sure to result in an irrep- 
arable injury. In several instances where the experimental beds of 
the writer have been made during the late autumn, and where a 
vigorous growth of spawn has been secured before the advent of the 
coldest weather, the beds have remained unproductive throughout the 
winter months, or so long as the temperature remained intermittently 
below 40° or 50° F. With warmer weather, these beds have come 
into bearing several months later, and where the temperature has then 
remained favorable for some time a good yield has been obtained. In 
this case, moreover, the bed will bear much longer at a temperature 
of 60° F., or above, than if the temperature has been constantly in the 
neighborhood of 60° F. throughout the growing season of the spawn. 
As a rule, beds thus filled with spawn and then subjected for a time 
to cold conditions yield at the outset much larger mushrooms than 
beds exposed to a more constant temperature, even if this constant 
temperature may be the optimum. 

At any rate, the beds must be “ cased ” as soon as convenient after 
the spawn is inserted. As a rule, one should wait from one to two 
weeks in order to be sure that the spawn is growing. Casing consists 
in applying to the bed a layer of loam from 1 to 14 inches deep. In 
France the casing soil consists usually of calcareous earth, sometimes 
mixed with loam. Ordinary loam of almost any quality will suffice. 
This should be secured in advance, and it is well to protect it from the 
weather, so that at a convenient time it may be worked over and, if 
necessary, screened, in order to free it from large pebbles or trash. 
When the loam is applied, it should, on ridge beds, be carefully 
firmed. When cased a bed should require watering for the most part 
merely to maintain a moist surface. 


MUSHROOM GROWING. 
EXPERIMENTS AT COLUMBIA, MO. 


The practical experiments in mushroom growing which have been 
undertaken at Columbia, Mo., were designed, in the first place, to 
determine the exact effect of conditions upon the growth of mush- 
rooms, and in the second place to test or immediately apply the 
results obtained or suggested by the laboratory work. The effects 
of temperature, moisture, etc., have already been discussed, and the 
conclusions drawn have been based upon the most careful observa- 
tions of the experimental beds, as well as upon the evidence which 
has been obtained by a personal study of the conditions in commer- 
cial mushroom houses and caves both at home and abroad. It is 
needless to give in detail the record of all failures or of poor yields 


“40 MUSHROOM GROWING AND SPAWN MAKING. — 


invariably obtained when the conditions were unfavorable—that is, 
when they were beyond the limits which have been more or less 
definitely stated as requisite. On the other hand, the results which 
are given do not represent the best yields obtained; they are those 
which seem to be most instructive. 

The experimental work has been seriously handicapped in one par- 
ticular. With only one set of experiments (those recorded in Table 
VIII) has it been possible to maintain a temperature constantly 
between 50° and 60° F. Unfortunately a north basement room 
which gave those results during the winter of 1903-4 has not since 
been available for the work. The results are, however, comparative 
when not absolute. 

The results given in Table VIII are referred to in various parts 
of this bulletin. Attention should be directed to the fact that many 
of these beds were yielding well when the experiment was neces- 
sarily closed to make room for a second series of experiments 
planned during the same winter. Beds Nos. 6, 9, 13, 25, and 40, for 
instance, each yielded between 8 and 15 ounces the day the experi- 
ment was closed, while beds Nos. 2, 10, 14, 23, 26, 30, and 37 each 
yielded 1 pound or more on the same day. 

It is to be noted that a considerable number of beds in this series 
produced more than 1 pound per square foot, and some nearly 
2 pounds for a similar area. It is certain that some beds would 
have yielded more than 2 pounds if they could have been per- 
mitted to produce longer. 


TABLE VIII.—Yields of experimental mushroom beds. 


| 


i , = cs) = ae He 
5 B./4 |8 |es|® 8 
we SA fi g 3 o|> Nn. 
Org a&|@ aH/2 | 8s 
23 ge | ae8| 22/28 | 2. )as 
, j o.. 
we sae ic Be Source of the spawn. Ty 5g ag i. iy $3 op 
® 
he 8 % A ak “5 Das a & 5 
58 gg | See ee 
Fi gaiz |= |g8le iz 
5 a iy Be s a 
Z Z a val is 4 nH 
1.) Feri eft ef dy| Alaska, Old . ssis cel cosas segs 27 53 54 | 107 6| 18.0 
horse manure. 
Bl aes Go - Asceiti. 2s Old American made --.-.....-------- 104 20 |} 20 6 8.6 
SD) eee 3 fn ee nelly + 9 ER Ys English, current year market 51 TAL Yocoe ‘ 6 1.0 
product. 
Dl hans OOAs cewtucad Him lish, 2 years Old. 6. .ccccacucccscs |e occlu cee le eeeee 0 6 0.0 
Bil aed. r: > i ae oe Higlish, 1 year Old i .~ wet «snp econ. dl cease. cele eee 0 6 0.0 
| Mi ~caceuwesee ra U.S. Department of Agri- 51 47 68 | 115 6} 18.8 
culture. 
ssh Ehre UO fees i teese Bohemia, U. 8S. Department of 53 48 ng 5| 13.0 
Agriculture. 
Ol lwown's Ot caademeeres Mixed varieties, U.S. Department 51 78 34| 112 6| 18.6 
of Agriculture. 
w lcteen MO ifeeasacesen Bohemia, U. 8S. Department of 1 at 1 gf ee 102 6} 17.0 
Agriculture, light spawning. 
bl pan GU sacakatwese. Bohemia, U. S. Department of 46 7 65 | 136 6} 22.6 
Agriculture, heavy spawning. 
cy bene OG wa cwewonvene Agaricus amygdalinus, old......--..|.-.-.-|.-----|------ 0 6 0.0 
12| Fermented) Bohemia 8. Department of 61 Ble ee 5 6 0.8 


U. 
horse manure Agriculture. 
(bed left for 2 
months before 
being spawned) 


peri- 


Number of the ex 
mental bed. 


5 


m& 8 


MUSHROOM GROWING. 41 


TaBLE VIII.— Yields of experimental mushroom beds—Continued. 


ERO Sy 6 SB I BB GS 
Bala |S |28)e | 
og 3 we | a 
ia |O7 z ag} 2 oe 
Wot eS .. | mi. tae | eRe) 
; nba oe ty cay Sus | Ae 
2-9 te a Source of the spawn. = 69) 83/22) 83 Bo 
+i Od] o/s |So] 3 
a8 |S), |/2 | as 
m [->) _ - —_ fo) = — Pp 
33 lex Paz al a FA 
Pig = =2 oa 
foals — ee fina fe 
a(S |3 |$2/2 [3 
a |b lh lac ld | 
Fermented/| Bohemia, U. 8S. Department of | 49; 110 50/ 160) 6) 27.7 
horse manure. Agriculture. | / 
a eee eee mole tee nh weit? Bae. | 61} 241) 40) 281) 12) 23.4 
Leaf mold ----... Calvatia cyathiforme __....--------- Paes ioe jeres.... 6 0.0 
seri do __..-.----..| Bohemia, U. S. Department of |------|---.--| 1 46iP? 4010 
Agriculture. 
Fermented sta- | Alaska, U.S.Departmentof Agri-| 48 | 9| 21.0 
ble manure;{| culture. | | 
bed fairlycom- — | 
pact. 
Fermented sta- ----- (1 ee ee eee | 53 93} 30| 123 6) 20.5 
ble manure. | 
cae do ......------, Bohemia, U. S. Department of | 48} 101} 39} 140| 6] 2.3 
Agriculture. | 
aoe ‘ig: SS Eee | Var.?, U.S. Department of Agri-| 53 96 | 37) 133 6 |. 22.2 
culture. 
a yf do __...--.----| American commercial more than §---.--|--.--.|------ 0 6 0.0 
1 year old. | 
1 =, do __...--.----| American commercial, Bohemia..-| 53) 111 53 | 164 | 8 | 20.5 
it Figen ts: 23 NP Aye Ue eee 2 eet.) | 46 67 | 113 | 9|/ b2 
ee do _...-..-----| Bohemia, U. S. Department of | 46 22 50 72) 6) 12.0 
Agriculture. Loose cakes; dried.) 
[S3ur3 3th) 2 eres Bohemia, U. S. Department of 49 | T4 75 | 159 6 | 26.6 
te ool Watered freely | 
/ ate. | / | 
285. do a2. 2-5--4 | Bohemia, U. S. Department of | 49; 42) 51, 93) 6) 15.5 
Agriculture. Wateredfreely. | / | | 
B28" do ._....------| Bohemia, U. S. Department of | 46; 89/} 30) 119) 6/ 19.8 
Agriculture. 
}3-28- Git 7 Saal eae i fines: dnralenspute aii ete, “eS ceael tes a 5d 90| 55) 145 8} 18.1 
embmaod) Gia-.| 52 0O-.. 0. - topos. sede ate eas 51 | 129| 146| 2% 9| 30.5 
blemanureand | 
5 pounds cot- | | 
ton-seed meal. | 
Fermented sta- | English commercial, St. Louis --_..|------|------ | 0 6; 0.0 
ble manure. | | 
ae “it hous 5 See | English commercial, New York ---|------|------ | ee pee | 6) 0.0 
se do _......-....| Bohemia, Americal commercial ___| 42 7 32 | 102 6| 17.0 
ae ot) ee ee Alaska, American commercial----- 46 7 31 | 101 6) 16.7 
ae ava | Pronch. commercial Hake 6) | 222|--- 2-2. 0 8} 0.0 
Fermented sta-| Bohemia, U. S. Department of | 53, 47); 96 143 9} 1.9 
blemanureand| Agriculture. | 
cotton-seed 
hulls. 
| Fermented sta-|__-_- G Tk See We Aes ee Per a Rae ae 104 6| 17.3 
ble manure; | 
bed heavily 
compressed. 
aS. do __.---------| Var.?, U.S. Department of Agri- 46 58: 46] 104! 6) 17.3 
culture. 
Fermented sta-|-__--- On ee ee ee 46; ll; uu 22 6 3.7 
ble manureand | 
sphagnum. 
Fermented|-...- oD SSR TA eee 5O.|,,, 44 |2-222. | 44 a i 
sheep manure. } | 
Fermented sta-|----- Age. eA bo ee ae 46 18 39 57 8 7.1 
ble manure, } 
cotton-seed 
hulls, and cot- 
ton-seed meal. 
Fermented cot-| Bohemia, U. S. Department of TY] eS eee 5 9 0.6 
ton-seed hulls Agriculture. 
and cotton- 
seed meal. | 
Manure mold-_-__|..--- 6 oR a A a REA ET oS TTS ae ol RN eee 2 6 | B same 
5:77 eS ee beter gt cyathiforme. Pure cul- |; ~---.|------|c<<<s6 0 6) 0.0 
ures. 
Old compost, left | Bohemia, U. S. Department of Me ees | ee 5 9 0.6 
2 months. be- Agriculture. 
fore spawning. 


42 MUSHROOM GROWING AND SPAWN MAKING. 


The series of experiments outlined in Table IX followed directly 
upon the series given in Table VIII. The beds in the first series 
were made in midwinter, and as the manure had been well fermented 
there was little or no rise of temperature after the beds were made. 
The spawn was therefore inserted at an unusually low temperature. 
During thaws in the late winter there was considerable seepage 
through the walls of the room. Some of the wall beds—Nos. 14 to 
21—-were seriously damaged, but although beds Nos. 7 to 13 were 
also wall beds seepage was not evident in this region. Within about 
thirty days after vigorous mushroom production began in this series 
the basement was flooded, and the work was therefore brought to an 
abrupt close. The yield up to that time is given, however, since in 
this series there are included many fertilizer tests. 


TaBLE IX.—Yields of experimental mushroom beds in a north basement room, 


1904. 
eg £ bah 
2A go|\8|%% 
ue CH | oF | an 
°= | Bedding material and fertilizer. Spawn used. oR BS Le 
og B e|"s|? § 
25 2 D | pm 
Ez Bale |8* 
A A al <q 
1| Stable manure and cotton-seed | Bohemia, U. S. Department of 48 33 6 
hulls. Agriculture. 
a ee 1G teas2 ks eae. 5 See Se Se Go fe.) 20s cate OR 1-25 48 94 6 
3 | Leaf mold and stable manure ------ eeeee do . .. =>. -2 Ree SS ee 48 38 6 
| ee (Eee oes ae ee Jazese GO = =scsssescsssseese es heceeerssese 48 36 6 
5 | Stable manure and sphagnum..-..--- jas=s GO 22222222225 sezte2 Cee 61 4 6 
6 | Stable manure and cotton-seed |__.-- GO 2 Soin Sass So a 61 64 6 
meal. : 
Alb Se bt ee Sena Sere Sas See pare GO smo ecna manned aha ee ee 73 6 
Stable manure, timothy fed_--.----|---.- do 22S i gh0o Dee 73 2 6 
ey Ee ih, Se SE ae EG So eee ee do... cuss. f2.0e. Bop oa de 0 6 
10 | Stable manure, clover fed --.-----..|----- Go 222 ELGG Bs ee se es 1 6 
Jt -12.2: = dg tS Sk. A et Go =. Leni. AU OOS. 22 dent 3 6 
12 | Stable manure, bran fed_-_.----.----|----- 46 = 2 Ose EtG et, RS. Sse 66 84 6 
13 eee Oe ca ashens a aeeon'snas 5 ee G6): [fob ee SOE fees 54 | 109 6 
14a| Stable manure, corn fed_----------- fcwaad GOSS . SS ee 71 12 6 
15a|__._.. Pt oe ey eee. > aes Ss le ieee. do ....2.. -24 22322. 2 68 8 6 
16a! Stable manure, oats fed__-..--..----|----- do .....4 22 eee See pie 2 ee 8 6 
d do 80 14 6 
71 24 6 
66 40 6 
vel 17 6 
55 6 
61 6 
55 6 
tilizer: KCl, 1 ounce; KNOs, 1 
| ounce; bone meal, 7 ounces. ) 
24 | Stable manure and incomplete fer- |----- GO . cists Sek 2.88 ecto | 64] 380 6 
| tilizer: NaNO;, 1 ounce; bone 
| meal, 7 ounces. 
25 | Stable manure and NaCl, 2 ounces.|....-. GO s c2scccscas secacs Seka wees 66 41 6 
26 | Stable manure and NaN Os, 2ounces.}|-..--- dot ..tgcek. c . Ai ee 48 42 6 
27 | Stable manureand MgSO,,2ounces. .-.-- a ee 66 39 6 
28 | Stable manure and K,SO,,2ounces.|-...- OO suse. «ok 2... ee 64 46 6 
29 | Stable manure and kainit, 4 ounces.|----. Ce oe 64 62 6 
30 | Stable manure and CaCh, 2 ounces.|----- GUL o2c5.0:5. REPO eee 64 48 6 
31 | Stable manure and Na,.HPO,, 2 |--..- Go . .-.00- es AA. Seen 64 65 6 
ounces. 
82 | Stable manure and (NHy,).SO,4, 2 |----- Oe SS ikfe stein = a nim oa ace a 54 41 6 
ounces. 
33 Stable manureand NaNOg, lounce; |----.. Oy ee a ery ee he 68 80 6 


kainit, 2 ounces. 


aSome of the beds in this block—Nos. 14-21—were seriously injured by seepage water, and the 
results are untrustworthy. 


MUSHROOM GROWING. 43 


TasLe 1X.—Yields of experimental mushroom beds in a north basement room, 
1904—Continued. 


as S a L 
4 ro ee z | _ 
38 Pele lye 
air On| og | 2g 
°e | Bedding material and fertilizer. Spawn used. On| oe | 2 
$8 Oo, | "ao | 3 
SE - > : os teal Yat 
& Aa ie | Som 
Bo BY |e | 2° 
4 Aa |p < 
PEP NORMADNT Os 3-2 2oser_ screens Hugi commercial (ordered as 68 34 6 
resh). 
ot) |g (oS SS / a aes Spawn from bed in full bearing.._. 66 12 6 
36 | Stable manure, lime dressing ------ Bohemia, U. S. Department of 68 8 6 
¢ | Agriculture. 
37 | Stable manure,ammonium molyb- ----.- 6 pp as Nemetlge ten AT thes Callie aM plea (?) 6 
date, } ounce. 
38 | Stable manure, ZnNOs, 1 gram_---- ----- ee ee ee tee eee Onan (?) 6 
39 ... Agaricus amygdalinus -__---------- 68 7 6 
tl) eae SUC ASE EE ie aha Bohemia, U. S. Department of 64 77 6 
Agriculture. 
Al if ceepree: 2 = nenenn See 7. eke English commercial (New York). 77 4 6 
Ce ee 2 eee ees eee Pee fe ET Bohemia, U. 8. Department of 64 33 6 
| Agriculture. 
3 LD Boe 2 eee ee Spawn from old bearing bed _--..-|__---- 0 6 
| BASES hee AAs oS Sat So SL SEAS 2S Plewr opis ostreatus. 22.22 2--2 522524 ERE 0 6 
Dal hese a Co Speee sod $22 as Se ee ee eae English commercial (Philadelphia) -___--- 0 6 
46 | Stable manure and sawdust -_-_--__- Bohemia, U. S. Department of |__---- 0 6 
Agriculture. 
47 | Stable manure -_____._.-_-.-.-------- Var.?, American commercial___..-| 48 60 6 
[O02 aC he ee ee ee Alaska, American commercial _.._. 64 22 6 


From the experiments given in the foregoing table further proof 
is furnished of the fact that stable manure alone, when of good 
quality, is sufficient for the growth of mushrooms. The addition 
of nutrient salts as fertilizers has not, on an average, given any 
marked increase in yield, but rather the contrary. It is hardly 
possible that the quantity of salts used on the beds was too little to 
make the effect felt. On the other hand, it was not sufficient to be 
injurious. It is evident from the experiment in bed No. 29, for in- 
stance, that the addition of 4 ounces of kainit could not have been 
injurious. In some instances the results obtained by the use of fer- 
tilizers were poorer than where the manure alone was used. This, 
however, the writer believes to be due largely to differences in the 
spawn used, or the differences in condition owing to the location of 
the bed, for subsequent experiments with some of the salts which 
seemed to be either injurious or beneficial have not wholly confirmed 
these results. It is to be noted, however, from the experiment in bed 
No.6 of this series and also from bed No. 30,in Table VIII, that the 
beds treated with cotton-seed meal have invariably yielded somewhat 
above the average. These beds do not come into bearing quite so rap- 
idly as those in which manure alone is used. It is thought that this is 
due to the fact that bacterial action is at the beginning more rapid in 
beds containing cotton-seed meal, and that, consequently, when this 
wave of bacterial growth has passed the nutrition of the spawn is 
favorably affected. Experiments had already indicated that manure 


44 MUSHROOM GROWING AND SPAWN MAKING. 


from animals which were fed a poor diet, such, for instance, as 
grass or hay alone, is much less valuable than where the animals are 
well fed. The experiments in beds Nos. 10 to 22 were designed to 
test the value of some different feeds. The writer was fortunate in 
being able to secure manure from work animals which were being 
used in feeding tests where very different foods were employed. 
Unfortunately, however, the mushroom beds were located next to a 
basement wall, and in beds Nos. 14 to 21 the results were vitiated by 
the fact that there was considerable seepage water in that region 
during the thaws and heavy rains of the spring. Nevertheless, it is 
believed that the experiments in beds Nos. 8 to 13 are trustworthy. 
An attempt was made to check these results by using some of this 
manure in tube cultures, and it was found that the manure used in 
beds Nos. 8, 9, 10, and 11 particularly was unfavorable for the 
growth of the mycelium even in the pure cultures. 

On account of its stimulating action upon the spores of Agaricus 
campestris a small quantity of ammonium molybdate was applied 
to one bed, No. 37, in order to test its effect upon the growing 
mycelium. Moreover, since certain salts of zinc at considerable dilu- 
tion have been found to increase greatly the quantity of mycelium 
produced by other fungi, zinc nitrate was employed in an adjacent 
experiment. The results of these two tests were the same. There 
was a profuse mycelial development and an abundant production of 
small deformed sporophores. 

Table X also summarizes a series of some interest. These beds were 
spawned early in November, 1904. Soon after the spawn began to 
spread throughout the beds—about December 15—the temperature 
of the room fell to 40° F. From that time on until Mareh 1, 1905, 
the temperature was constantly below 52°, and on several occasions 
as low as 32° F. After two or three weeks of warmer weather the 
beds began to bear vigorously, and the mushrooms, particularly the 
first ones, were of unusual size and of excellent flavor. Numerous 
individuals weighed from 6 to 8 ounces immediately after the sepa- 
ration of the ring, and a few mature specimens ranged from 10 to 14 
ounces. 


MUSHROOM GROWING. 45 


TABLE X.— Yields of experimental mushroom beds—Third series. 


Compar- 
Bed z ative 
N Material constituting bed. Spawn used. yield per 
2 bed, in 
ounces. 
aiiesitable manure —s ....--- 225. 222L2.225_- English commercial, 2 years old__-_----- 0 
(Se Sa it cote ee eee Columbia, “green”? spawn, U.S. De- 7. 
partment of Agriculture. 
See ae din: 2.3 eS ee ea eee ae Poor grade English commercial, re- 16 
cent importation. 
‘ipo A 2 Re eee Ps Good grade English commercial, re- 49 
cent importation. 
ee fy 2 ee See ee eee Good grade English commercial, 6 40 
; months old. 
(>| ee Gg Rs 2228S Se caer eae American commercial _....- 22-2..-as--< 57 
i ON IRE EASE SES) CEU a tee eel ee Ome aia. MOLES IS £3 4) 0-e eee 34 
* UO S68 Se Bae re ee Oe i a ce oe a 54 
Otrets. ps PS. oS iis hehe lee U.S. Department of Agriculture, Co- 56 
lumbia. 
10 | Rotted sawdust and stable manure ____|____- opens 2. sas ejes 85 ALES 1 31 
11 | Leaves and stable manure -___- eee eee 30 
Weiisaw Oust 2.225.242. 22 fib sss yo? At: 3 
PeameaVes. 2... 8. =.- == Bee: (ee oe is ee. ein = 6 
14 | Stable manure --.-....-...--. American commercial, probably A. ar- 60 
vensis, Var. 
ABI 22% 5 Riep ee tee) OPE e vis! ett ee eee y - American, A. villaticus -_..........-.--- 68 


In some publications on mushroom growing the claim is made that 
old or practically exhausted beds may be brought into bearing again 
by heavy fertilization with hquid manure or with a weak solution of 
potassium nitrate. From a commercial point of view, no measurable 
success has resulted from any trials of this nature made by the writer ; 
consequently, it is believed that exhausted beds should be immediately 
discarded. From the standpoint of mushroom sanitation, this is 
also particularly desirable. 


VARIABILITY IN MUSHROOMS GROWN UNDER DIFFERENT CONDITIONS, 


The writer does not intend to discuss even in a general way the 
relationships of the various forms of Agaricus—that is, those that 
may be considered allies of A. campestris—which he has cultivated or 
studied in the field. Some reference to the variability of common 
forms should, however, be made. For a comprehensive study of spe- 
cies and varieties, a knowledge of European forms as well as of those 
found in America is essential. Authors differ so widely in their 
descriptions of species, as well as in their conceptions of them, per- 
haps, that in the absence of unlimited material nothing short of con- 
fusion results from any attempt to harmonize opinions. It is sufli- 
ciently difficult to separate what many would regard as varieties of 
A. campestris from those of A. arvensis. When specific rank is 
bestowed also upon such forms as A. pratensis, A. villaticus, A. mag- 
nificus, A. rodmani, etc., the difficulties are greatly increased. The 
writer has grown many forms of Agaricus, and, as might be expected, 
there seems to be no form which will remain practically constant 
under variable conditions. Besides general size, size of spores, etc., 


46 MUSHROOM GROWING AND SPAWN MAKING. 


some of the characters used in separating the common forms are color 
of gills; character of ring, particularly as to whether single or double; 
shape of stipe; color and markings of pileus; color of flesh, ete. In 
following the development of these characters in different forms, 
many variations will be found. Agaricus campestris grown on com- 
posted leaves shows very little pink in the gills. The color changes 
rapidly from dull pinkish-brown, or almost white, to a leaden hue. 
Several brown-capped forms, usually considered varieties of A. cam- 
pestris, never show a bright-pink surface unless produced under 
exceptionally favorable conditions, moist air being a sine qua non. 
The ring is naturally variable. In any variety of A. campestris it is ~ 
not uncommon for an edge of the partial veil to remain attached to 
the base of the stem as a volvate line, or this line may be left at any 
stage during the elongation of the stem. Again, if the lower margin 
of the partial veil on the stipe separates slightly from the stipe, and 
upon drying curves slightly upward, there is an indication of a double 
ring. A very good double ring appeared on a number of very vig- 
orous specimens of an undoubted variety of A. campestris during the 
present season. It is possible that there is a greater tendency to pro- 
duce a double ring when conditions are favorable for the production 
of the most vigorous mushrooms. Agaricus arvensis is also very 
variable with respect to the formation of a double ring, as also in the 
persistence of the partial veil. 

The shape of the stipe is in many forms dependent upon the con- 
ditions. Under favorable conditions a brown variety of A. cam- 
pestris may have a very short, thickened, equal stem, when grown on 
manure, and practically uniform at maturity, while the same form 
grown on decayed leaves may show in the main a stipe with thick- 
ened base, gradually tapering to the top. The color of the cap is of 
undoubted value as a varietal or specific character, yet it must be 
remembered that whether the surface be smooth or rough, merely 
fibrillose, or broken into scales of definite form, may depend entirely ~ 
upon whether produced in moist air or in dry air, subjected to 
drying after being wet, etc. The color of the flesh is also dependent, 
to a considerable extent, upon the conditions. A specimen grown in 
even fairly unfavorable conditions will show the flesh somewhat 
darkened, and on exposure the characteristic pink tint will not be 
even momentarily visible. In other words, a considerable range of 
variation must be anticipated, and in comparisons there should be 
stated very clearly the conditions under which the particular forms 
are produced. . 


THE CULTIVATION OF VARIOUS SPECIES OF MUSHROOMS, 


In Table X are given the results of a single test with Agaricus 
arvensis, or What is supposedly a brown variety of this species, and 


MUSHROOM GROWING. 47 


also of a single experiment with A. villaticus. In both cases the 
yield was excellent. It is not well to draw definite conclusions 
from individual tests, but it is believed that both of these forms will 
yield profitably in general culture under conditions similar to those 
required for A. campestris. Plate III, figure 2, indicates the size 
and compactness of the mature sporophore of A. villaticus. More- 
over, both of the species above referred to are to be recommended 
for texture and flavor. Two forms of Agaricus fabaceus (see Pl. 
III, fig. 1), both with amygdaline odor and flavor, have been tried 
in relatively few experiments. In no case has the yield been very 
good, and further experiments will be required before it will be pos- 
sible to state under what conditions these forms may be most suc- 
cessfully grown. At the Missouri Botanical Garden Prof. Wiliam 
Trelease has for some time grown successfully one of these varieties. 

Owing to the profuse and rapid growth of the mycelium of Copri- 
nus comatus in pure cultures, it was anticipated that it might easily 
be grown in beds. The few experiments thus far made indicate that 
in impure cultures (beds) of leaf mold the mycelium grows and 
spreads very slowly. Hot weather prevented the maturity of the 
tests, but no sporophores were produced during a considerable pe- 
riod. In similar experiments Lepiota rhacodes and Tricholoma per- 
sonatum were used. The former has given unsatisfactory results thus 
far, but the latter is promising. 

It is not yet time to report on the possibility of growing the better 
and larger species of puffballs and the morel. It has already been 
indicated that the mycelium of these fungi grows well in pure cul- 
tures. From the pure cultures it has also been demonstrated that 
spawn may be made, but it has not been determined under what con- 
ditions the fruit may be produced. Figure 1 on Plate IV shows a 
young specimen of one of the puffballs, Calvatia craniiformis, the 
spawn of which is produced with the least difficulty. 


COOPERATIVE EXPERIMENTS. 

During the winter of 1902-3 a small quantity of experimental 
spawn made by the writer was sent out to mushroom growers for 
trial; in 1903-4 this spawn was made in large quantity, and trial 
packages were sent to more than 100 growers or interested persons. 
At that time Farmers’ Bulletin No. 204 had not been issued, and the 
instructions which could be furnished inexperienced growers were 
inadequate. Nevertheless, an attempt was made to obtain reports 
from all persons receiving the experimental spawn, even from those 
who had applied for and received spawn when the season was too far 
advanced for successful work except in caves and cool cellars. A 
number of reports were received, but, as might be expected, fully 50 
per cent of these indicated that the conditions under which the experi- 

23994—No. 85—06 m4 


48 MUSHROOM GROWING AND SPAWN MAKING. 


ments were made were wholly unsatisfactory, and that, therefore, no 
favorable results could be anticipated. Among those whose reports 
indicated that the conditions were favorable, or fairly favorable, only 
a small percentage reported failures, while four-fifths of those claim- 
ing success secured yields of more than one-half pound per square 
foot of bed space, many obtaining more than 1 pound per square 
foot. In two instances a yield of nearly 2 pounds to the square 
foot was reported. The frontispiece, Plate I, a bed in full bearing, 
and Plate VII, figure 1, showing the mushrooms as prepared for mar- 
ket, are photographs furnished by cooperating growers who are now 
also making spawn of pure-culture origin. It was suggested to 
growers who received the experimental spawn that a comparative test 
of the English or other commercial spawns with that received from 
the Department of Agriculture would be of interest. Comparative 
tests were made and reported by 10 growers. In most cases the 
English spawn, obtained at random on the market, failed to grow. 
In only one case did the English spawn prove better than the pure- 
culture product, and in this instance the spawn furnished by the 
Department when used was nearly one year old. 

Failures may always be anticipated when attempts are made to 
grow mushrooms under adverse conditions, and it must be said that 
greater success was obtained from the cooperative work than could 
have been hoped for, considering the fact that many of the persons 
who sent in reports were wholly inexperienced and were practically 
unguided. 

During the present year experimental mushroom spawn has been 
sent to more than 200 interested persons, and this will doubtless be 
the last general distribution of this product by the Department of 
Agriculture. Representing the varieties of Agaricus campestris 
commonly grown, mushroom spawn of pure-culture origin is now an 
established market. product. In order that the standard of the 
American spawn may be maintained, spawn makers, dealers, and 
growers should see to it that only the fresh, recently dried product 
is used. 

Nevertheless, it is hoped that this cooperative work may be carried 
forward, looking toward the development of better varieties or the 
bringing into culture and the testing of new species. 


CAVE FACILITIES IN THE UNITED STATES. 


Cave facilities in the United States are by no means so meager as 
has been supposed. There are in some sections caves from which 
rock for Portland cement has been mined. Some of these have been 
utilized for mushroom growing. There are also natural caves of 
great extent in many of the States of the Central West—especially 


OPEN-AIR CULTURE. 49 


in Indiana, Missouri, Kentucky, and Arkansas—as well as in Vir- 
ginia.t’ The difficulty is to obtain caves within a convenient distance 
from cities, for stable manure becomes expensive if it must be hauled 
many miles or transported long distances by the carload. Again, 
caves should be easy of access, since after each crop every vestige of 
soil, manure, etc., of the preceding crop must be removed as a sani- 
tary precaution. This is especially necessary since there is much 
waste space in most natural caves, and it becomes a very difficult or 
expensive matter to fumigate. If the cave system is extensive, it 
must also be possible to give it thorough ventilation. Many natural 
caves are the courses of subterranean streams. The latter are by no 
means objectionable if there is no danger from overflow. In many 
caves the stream has long since found a new channel and the cave 
is dry. Seepage water, usually accompanied by continuous stalactite 
and stalagmite formation, is undesirable. In some of the Eastern 
States coalpits or coal mines may be important for mushroom pur- 
poses. Where the coal mine is not too deep, or where perfect venti- 
lation may be given, there is no reason why it is not entirely suitable 
for mushroom growing. 


OPEN-AIR CULTURE. 


In some sections of England and France open-air culture of mush- 
rooms in beds is practicable during the late autumn and winter 
months, in which case the productive period may extend into the 
spring. The difficulties in the way of open-air culture are not merely 
those of maintaining a more or less uniform temperature, but also 
of maintaining practically constant conditions of moisture. For 
these reasons it is necessary to mulch the beds heavily with clean 
straw. In some instances a light mulch of straw is permitted to 
remain even during the period of production, for a rapid drying out 
of the surface would be hazardous or fatal. It is better, perhaps, 
to put the beds under some form of protection, such as an improvised 
cold frame. 

In regions where the climatic changes are marked, open-air cul- 
ture is probably not to be recommended during any season for com- 
mercial purposes. It is probable that there are some areas in the 
United States in which open-air culture might be practiced with 
profit. It has seemed that certain sections of California might be 
favorable for this phase of the work. In the interest of experiments 


«The writer is indebted to Prof. C. F. Marbut for the information that caves 
are to be expected in the Silurian limestone, which occurs particularly in the 
extension of the Shenandoah Valley, in the bluegrass region of Kentucky, and 
in the Ozark region of Missouri and Arkansas; also in the Lower Carboniferous 
limestone, which extends into Indiana, Kentucky, Tennessee, and Missouri. 


50 MUSHROOM GROWING AND SPAWN MAKING. 


along this line the writer has made a special attempt to acquaint 
himself with the conditions in that section of the country. This has 
seemed particularly desirable, inasmuch as fresh mushrooms could 
not be shipped to the far West from sections in which they are at 
present grown in quantity. From the information obtained it is 
thought that successful open-air mushroom growing might be antici- 
pated in those sections where the average temperature is between 
48° and 55° F., provided there are relatively few days when the 
temperature falls as low as 32° F. At the same time, open-air cul- 
ture can not be recommended for those sections in which dry winds 
are prevalent. As a rule, during the wet or winter season the rain- 
fall is so light that heavy mulching would probably suffice to prevent 
injury from excessive wetting. Nevertheless, it seems apparent that 
even in regions most favorable for open-air culture some inexpensive 
partial protection against the changes of temperature due to direct 
sunlight, or against heavy rainfall, would be desirable. 

It was also ascertained that Agaricus campestris appears naturally 
in some quantity during the months of January and February, or 
longer, during the rainy season. This, however, is also true of other 
species of fleshy fungi. The large size of some of the specimens of 
Agaricus campestris and A. arvensis found would seem to suggest 
that they were produced from an unusually vigorous mycelium. 
This may be the result of a condition analogous to that previously 
mentioned, where, on account of the low temperature of the atmos- 
phere, the spawn may develop slowly through a considerable period, 
and finally, under favorable conditions, sporophores of unusual size 
are produced. 

In the following table are given the monthly mean temperatures 
from several representative stations in California during the years 
1899 and 1900. From this table it will be seen that so far as the 
mean temperature is concerned Eureka and San Francisco would be 
especially favorable during a large portion of the year. Independ- 
ence and Red Bluff are likewise satisfactory, while San Luis Obispo, 
Santa Barbara, Los Angeles, and San Diego show a mean which is 
perhaps rather too high.. The moisture of the atmosphere, the 
prevalence of hot winds, the variation in the daily temperature, and 
the number of hot or cold days must all be considered. From the 
data obtained, the general conclusion seems to be that the most favor- 
able regions are those where conditions correspond closely to those of 
Eureka and San Francisco. This, however, represents a large region, 
including a considerable portion of the San Joaquin and of the 
Sacramento valleys. In a few places experiments have already béen 
undertaken to determine the possibilities for the development of this 
work, but no definite recommendations can be made until the experi- 


OPEN-AIR CULTURE. 51 


mental evidence is at hand. It may be said, moreover, that some of 
the regions which seem to be too warm for open-air culture may be 
especially favorable during several months at a time for mushroom 
growing in ordinary cellars, or in very simply constructed mushroom 
houses. In those sections the winter and early spring months would 
doubtless give the most satisfactory conditions; and this period, 
fortunately, corresponds with the tourist season—a season when the 
market demands are greatest. It is also possible that with mulching 
and with simple protection, mushroom growing may be successful in 
some of the Eastern States. 


TasLeE XI.—Mean monthly temperatures at points in California, in degrees 


Fahrenheit. 
j 
Eureka. |San Francisco. [872 Uuis |santaBarbara. 
Obispo 
Month. Reolg.s Gaeta 
: 1899. 1900. 1899. 1900. 1899, 1900. | City. F.H.S.4 
} 
; 
JUDE 22 a ee ieee 47.5) 50.4 53.0} 50.7) 54.2] 56.2 | 53.0 55.4 
LE Le Bees Sie Bes ae ee 44.4 48.6) 51.6) 53.6) 54.4 56.2} 54.6) 58.0 
EP ahs See sap oe Bab gl ate 48.0 50.5 52.2 55.2 54.0 58.2 55.3 | 57.4 
TC ECR 8 ERUPT ERS BSE ae eee 48.2) 50.0) 54.6) 54.0) 56.4] 54.2) 57.9) 59.3 
eh eee. 49.6 54.4 52.6 57.0 54.0 61.6 59.4 59.4 
te ee eee ee 2155 52.09 56.2 56.9 57.6 62.4 63.9 62.6 64.4 
(To 2: ee ae ee ae 54.8 56.4 55.9 58.2 64.4 64.2 65.5 68.1 
emmeeeieiia ni oso 55.9| 57.0] 58.3| 59.7] 640] 649] 66.9| 68.9 
Pepe 5 ss 54.8 56.6] 58.2 63.3) 65.5 64.4 66.1 69.9 
5 1 CES ORL ESS EES APOE oe 52.0 53.8 59.3 58.8 59.6 62.8 62.6 | 64.8 
November .........-..... ae 55.9| 53.3] 56.8| 56.3] 57.4| 59.8] 591| 647 
December 120.22 2212022222222] 48.0|. 50.8| 49.6] 50.2| 543] 55.6| 55.6| 58.4 
ToT ae £8 ees Seer eee 50.9 53.2 54.9 56.2 58.4 60.2 59 62.3 
| Los Angeles. San Diego. |Independence. Red Bluff. 
Month. erckter ote ot = | 
; 1899. 1900. 1899. | 1900. 1899. 1900. | 1899. 1900. 
| | Pt Pa oe 
DEUMEBEP OA at. 31 soars. 1 56 68} 56.0| 57.1| 40.2) 466] 48.8| 48.8 
USED es le 54 58 | 53.4 57.2 46.5 48.1 51.6 | oat 
LT e Date eee eh Rehr ae ere 57 60 56.4 59.1 50.5 54.9 | 52.2 58.6 
april er ee 60 57 58.0 57.1 59.4 52.0| 60.8 57.6 
“+3 fi Teg or ot bere ey eae ees eee 60 64 58.0 60.6 60.0 65.8 63.2 | 67.0 
Dh ree adele! Salle el ie 65 67 61.4 63.9 74.2 75.4 | 76.8 
LST Ee ia et ee ee ae 70 yal 65.6 67.1 80.4 79.4 82.0 82.6 
Au he es phe ee He ee 69 68 65.8 65.7 72.6 72.4 73.8 77.0 
vO PTAD Loy Se eS ee epee eae 70 67 | 65.5 65.3 74.6 63.5 78.0 69.9 
_ 5. Bly tee ele sat ot She 63 64 62.7 62.8 5d. 4 58.8 61.0 60.0 
Mevemiber =. 220 saris" 62 66 61.0 63.7 49.4 50.4 54.4 54.8 
UL: id) sivo sect ei ah aaa 58 60 58.7 59.7 43.1 43.4 45.5 45.4 
Salo ay lai @) 64) o.2) @6) 58.9) 59.2 | 62.4) 62.5 
} } 


* Foothills or suburbs of Santa Barbara, at an elevation of 750 feet above the city. 


Occasionally one reads of successful natural cultures of mush- 
rooms; that is, the production of this plant in pastures, lawns, etc., 
under more or less natural conditions. At Columbia, Mo., the writer 
has made numerous attempts to spawn plats in pastures and lawns; 
but thus far failure has attended every attempt. The spawning has, 
moreover, been tried at every season of the year. It is believed that 
in the section of the country mentioned only exceptionally favorable 
seasons will permit any success in this phase of open-air culture. 


52 MUSHROOM GROWING AND SPAWN MAKING. 


MUSHROOM SPAWN MAKING. 


The mycelium of the cultivated mushroom has long been known 
commercially as “ spawn.” From early times it has been recognized 
that mushrooms may be grown from spawn, and it is quite certain 
that in all attempts to propagate mushrooms spawn has been used 
for the purpose. 

In France, in England, and in other countries in which the mush- 
room has long been grown it is recognized that it is not profitable 
continually to take growing spawn from one bed to be preserved as 
“seedage ” for the next crop. The common expression is that the 
spawn “runs out” in about three years. There seem to be few or 
no definite experiments indicating the exact conditions under which 
the spawn in two or three years loses the power of vigorous mush- 
room production. Nevertheless, it is the almost unanimous opinion 
of all extensive growers that there is a marked diminution in the 
yield after several successive propagations from the spawn in the 
mushroom bed. This has seemed to be true in the writer’s experi- 
ments, although it must be said that accidents to experiments under- 
taken have made it impossible to report at this time upon the nature 
of this running out. That deterioration does result is apparently a 
fact accepted by all scientific men who lave given attention to mush- 
room growing. It is possible, however, that under certain conditions 
the spawn might be repeatedly propagated without loss of prolific- 
ness. It is not necessary to enter here into a discussion of possibili- 
ties or to attempt to explain why weakening might be evident under 
ordinary conditions. 

A “chance” method—F¥or practical purposes it is necessary to 
renew the spawn and to secure, if possible, spawn which has not pre- 
viously weakened itself by the production of mushrooms—known as 
virgin spawn. Natural virgin spawn may be found wherever “in 
nature ” it has been possible for the spores to germinate and to pro- 
duce a mycelium. Ordinarily such so-called “ spontaneous ” appear- 
ances of spawn may be anticipated in compost heaps, rich garden 
beds, pastures near the feeding places of animals, ete. 

Many attempts have been made by practical growers to develop 
spawn from spores, sowing the gill portions of mature mushrooms in 
specially constructed beds; but the results, so far as the writer is 
aware, have not been satisfactory. As a rule, therefore, growers 
have been compelled to rely wholly upon a virgin spawn which has 
been obtained by chance. It is said that in the vicinity of Paris 
some persons make a business of searching for this virgin spawn, 
which they sell to the growers at a high figure. It is claimed that 
they become so adept in detecting the differences in the character of 
growth, the quality of odors, etc., that they can distinguish not only 


MUSHROOM SPAWN MAKING. 53 


Agaricus campestris, but also some of its varieties. In England 
much of the virgin spawn has been obtained from pastures. Where a 
“spontaneous ” growth of spawn is observed, trenches are dug, and 
these are filled with good stable manure. The latter in time becomes 
penetrated, and it is highly prized for cultural purposes. As a rule, 
the virgin spawn is used in spawning beds, which, when well pene- 
trated, are torn down, and the whole bed used as flake spawn in 
spawning the general crop. Again, the virgin spawn may be used 
in spawning the brick, or cakes, this. being the form in which English 
spawn is usually made. However adept persons may become in the 
identification of various varieties of spawn by odor, etc., this must 
be considered essentially a chance method. 

A “ selective” method—From what has been said it will be per- 
ceived that very little advancement could be made in the selection of 
desirable varieties of mushrooms, in varietal improvement and the 
like, so long as the chance method of securing spawn should prevail. 
The studies in the germination of mushroom spores previously re- 
ferred to were encouraged by the apparent necessity of beginning 
with spores from mushrooms of known qualities in order to effect 
improvement. In recent years the investigations of Costantin * upon 
spore germination have found application in a department of the 
Pasteur Institute. By a secret method, mycelium is grown from the 
spores in pure cultures. These cultures, which are, of course, pure 
virgin spawn, are then offered for sale to the growers. This spawn 
does not seem to have received deserved consideration on the part of 
the growers. The secret method of effecting spore germination re- 
ferred to by Répin® has also been practically applied by one of the 
largest seed firms in Paris. In general, however, French growers 
have not profited so much by the new methods, perhaps partially on 
account of the fact that these methods are not known and partially 
because of the expense of the new virgin spawn. It is to be noted 
that these methods imply pure cultures to begin with. 

The successful germination studies with chemical stimulation men- 
tioned in this paper were soon overshadowed by the discovery of the 
ease of making tissue cultures. The use of the latter method has 
been the means of a sudden advancement in spawn making in this 
country during the past two years, for many practical men have 
been quick to see the advantages which it offers. 

Pure-culture precautions——It has already been stated that the 
pure-culture method of making virgin spawn is not one which will 
prove successful in the hands of wholly inexperienced persons, or of 
those who are unwilling to spend time and use the utmost care in the 
manipulation of the cultures and the culture material. The use of 


¢Costantin, J., loc. cit. b Répin, C., loc. cit. 


54 MUSHROOM GROWING AND SPAWN MAKING. 


pure-culture methods necessitates to a considerable extent a knowl- 
edge of the bacteria and molds which are everywhere present in the 
air and which are especially abundant wherever there are dusty or 
damp, moldy conditions. The principle of making pure cultures is 
briefly this: The materials, or media, and all the vessels employed 
must be sterilized, which implies being heated at a temperature suffi- 
cient to kill all germs present in the vessels or materials used. If 
the vessels used are test tubes or other pieces of glassware with small 
mouths, they should, previous to sterilization, be plugged with cotton 
batting. This cotton batting prevents, when carefully manipulated, 
the entrance of germs from the air, and therefore keeps the vessel 
or medium in a pure or sterile condition. If such a vessel is opened, 
this should be done in a room free from currents of air or falling 
dust particles; and, while open, tubes and other apparatus should be 
held in a more or less horizontal position, so that they will be less 
liable to contamination. It follows, of course, that the cotton plug, 
if removed, should not come in contact with any unsterilized sub- 
stances. If, now, a small quantity of the growing mycelium of a 
mushroom from a pure culture is transferred to such a sterilized 
tube, using for this transfer sterile needles, or scalpels, there will be 
little danger from foreign organisms, and the piece of mycelium 
inserted will therefore grow as a puie culture free from all other 
fungi or bacteria. 

The tissue-culture method.—In making pure cultures of mushrooms, 
large test tubes or wide-mouthed bottles may be used. These should 
be carefully cleaned, and, if possible, a sterilization should be given 
by means of dry heat as a preliminary precaution. In this event the 
tubes are plugged with cotton plugs and placed in a dry oven made 
for the purpose. They are heated to a temperature of about 150° C., 
and this temperature should be maintained for nearly an hour. 
Ordinarily, however, in rough work it is not essential to employ this 
preliminary sterilization. In either case the tubes are next partially 
fled (about two-thirds) with the manure, or half-decayed leaves, 
upon which it is desired to grow the virgin spawn. A plug is 
inserted in each tube, and the tubes are then sterilized in a steam 
boiler or under pressure. If sterilized under steam pressure, as in an 
autoclave, it is necessary to use about 15 pounds pressure and to allow 
the tubes to remain at this pressure for from fifteen minutes to half 
an hour. If the sterilization must be effected in a boiler or in an open 
water bath, it can only be done at 100° C., of course; and it is then 
desirable to boil the tubes for at least one hour on each of two or three 
successive days. 

With the tubes thoroughly sterile, the next step is to make the 
cultures or inoculations. By the tissue-culture method it is implied 


MUSHROOM SPAWN MAKING. 55 


that the inoculations are made from pieces of the tissue of a living 
mushroom. It is at this stage that selection may be made. One 
should procure from a bed of mushrooms in full bearing a mushroom 
which represents the most desirable qualities that are to be found. 
Size, quality, and general prolificness must all be considered, as well, 
also, as other characteristics in any special selections. One may de- 
sire, for instance, to select from a variety which yields throughout 
a long period—one which is resistant to higher temperatures, ete. 
Having found the mushroom from which it is desired to propagate, 
plants as young as possible may be used, and those which show the 
veil still intact are especially desirable. With a scalpel, or a pair 
of forceps, which has been sterilized by passing the blade through a 
gas flame, or even the flame from an alcohol or ordinary lamp, small 
pieces of the internal tissue may be removed, and these pieces trans- 
ferred to the tubes, without, of course, coming in contact with any 
object whatever which has not previously been sterilized. It is a 
good idea to wash the mushroom first, so that no dust will be made. 
The plant may then be broken open longitudinally and bits of the 
internal tissue readily removed without fear of contamination when 
one becomes adept in this kind of manipulation. Immediately upon 
inoculation the cotton plug is replaced in the tube, and after all the 
tubes are inoculated they should be put out of the dust, preferably in 
a situation where the temperature is about that of an ordinary living 
room. In the course of several days a slight growth may be evident 
from the tissue if the conditions have been perfectly sterile. In the 
course of a week or more the growth should become very evident, and 
in three weeks the moldlike development of mycelium should spread 
to practically all parts of the medium in the tube. The method of 
making pure cultures and the laboratory apparatus usually involved 
are shown in Plate VI, figure 2. 

When the tubes are thoroughly “ run ” the contents may be removed 
and used in spawning brick. The contents of a single tube may 
spawn several bricks when carefully employed. If no transfers are 
made of the growing mycelium from one lot of tubes to another, the 
writer has not found it at all impracticable or unfavorable to utilize 
this first lot of bricks later in spawning others. No further trans- 
fers, however, should be made from these bricks to others under any 
circumstances in spawn making. As elsewhere indicated, such a con- 
tinuous transference is injurious to the vigor of the spawn and 
diminishes the quantity of mushrooms produced. 

The commercial process —The essentials in spawn making are (1) 
a uniform, compact manure brick; (2) vigorous and well-selected 
virgin spawn to be used in inoculating the bricks, and (3) favorable 
conditions for the storage of the bricks during the growth of the 
spawn. 


56 MUSHROOM GROWING AND SPAWN MAKING. 


It should be indicated that there is no one method of making brick 
spawn. The process may and will be varied by each spawn maker. 
Any skill or mechanical devices which will simplify or improve the 
process in any particular are to be recommended. 

The materials entering into the composition of the brick are fer- 
mented stable manure, cow manure, and sometimes a small quantity 
of well-selected loam. Perhaps the chief value of these different con- 
stituents is as follows: 

In the horse manure the mycelium grows most readily. The cow 
manure binds the materials together into compact brick. The loam, 
which is perhaps least essential, is supposed to prevent cracking or 
hardening of the surface, and therefore contributes to the appearance 
of the finished brick, at the same time tending to prevent rapid fer- 
mentation during growth. It also in some cases facilitates the uni- 
form spread of the mycelium. If fresh manure is used, the necessity 
of using loam is perhaps to be emphasized. 

In the experiments which have been made under the auspices of 
the Department of Agriculture these materials have been used singly 
and in various combinations, and it is beyond doubt that the relative 
proportions of these should be determined by the special conditions 
under which the spawn is made. Excellent results have been ob- 
tained by using a mixture of from two-thirds to three-fourths stable 
manure and the remainder cow manure. In this case the compost 
for the brick is subjected to fermentation previous to its use. When 
loam is employed it may be used in more or less equal proportion to 
the cow manure; and the quantity of stable manure should about 
equal that of the other two ingredients. If the straw present does 
not become sufficiently disintegrated during the preparation of the 
manure to enable one to make a smooth brick, it should be removed, 
in part at least. 

The dry bricks podinanily measure about 53 by 8} by 14 (to 1}) 
inches. They should therefore be molded of somewhat larger size, 
perhaps 6 by 9 by 2 inches, since there is considerable contraction 
during drying. The mold consists merely of an oak frame of four 
pieces strongly riveted together. It may also be profitably lined 
with thin steel plates. In molding the brick one of two methods 
may be followed: (1) The compost may be thoroughly wet or pud- 
dled; then, with the mold upon a board of suitable width, the manure 
is compressed into it, the mold removed from the brick then formed, 
and the board pushed along for a succession of such impressions. 
The boards supporting the bricks are then disposed in racks and the 
bricks dried for a few days, or until they may be turned on edge for 
further drying out. (2) The compost may be used in a condition 
which is merely moist. It is compressed into the brick with some 
force, a mallet being often employed. The brick thus obtained is 


MUSHROOM SPAWN MAKING. 57 


sufficiently rigid to be immediately handled if necessary. By this 
method, unless the compost has been in excellent condition, the bricks 
are not so smooth as might be desired for commercial purposes. In 
some instances they have then been subjected to a repress process, 
an old repress brick machine being adapted for the purpose. In 
such cases the bricks are made thicker to begin with. The second 
method has been discontinued by some who at first employed it. 

Two methods are also employed in spawning: (1) The more com- 
mon method is to insert into the brick near both ends a piece of the 
virgin spawn obtained for the purpose. A cut is made with the 
knife, the spawn inserted, and a stroke of the knife effectively closes 
the surface. This must be done as soon as the brick can be readily 
handled. (2) The bricks are dried until merely moist throughout; 
then, on being piled, nocules of spawn are placed between successive 
bricks, a piece at each end. In either case the bricks are not piled 
for the growth of the spawn until in good condition as to moisture 
content. This should be determined not by the surface, but by the 
interior of the brick. In the pile the surface will soon become moist. 
When the first method is employed it is sometimes customary to 
spread between the layers of brick in the pile a little moist manure 
or sawdust. It has been determined, also, that the absolute mois- 
ture content of the brick should be about 40 per cent, which is the 
same as for the mushroom bed. Tests of the moisture content of 
bricks growing well have varied from 35 to 474 per cent. 

Occasional examination should be made to determine the tempera- 
ture and the extent of growth. In order that the bricks may become 
thoroughly penetrated, more than a month will usually be required. 

The most favorable conditions for the growth of the spawn are 
practically the same as for mushroom growing. A fairly moist 
atmosphere, maintained, if necessary, by spraying, and a more or 
less uniform temperature (55° to 60° F.) are to be preferred. The 
size of the piles will depend upon the other conditions; but if there 
is any danger of considerable fermentative activity the bricks should 
be so disposed as to permit perfect ventilation between two or more 
adjacent rows. 

When the bricks are thoroughly “ run ” they are dried under cover 
before being shipped or stored in bulk, since in a moist brick the 
spawn would continue to grow and would soon produce small mush- 
rooms or else would become moldy. Well-penetrated bricks of spawn 
‘are shown in Plate VII, figure 2. The areas of mycelial growth 
should be evident to the eye. The growth should be moldlike, how- 
ever, rather than composed of very large threads or fibers. 

The suggestion made in a previous publication that mushroom 
spawn should be sold by the brick (with a uniform standard of size) 
seems to have been adopted by American makers. The trade names 


58 MUSHROOM GROWING AND SPAWN MAKING, 


suggested for the common types of Agaricus campestris in culture 
have also come into use. It is certain that these names, Alaska, 
Bohemia, and Columbia, designating respectively a white, a brown, 
and a more or less cream-gray form, do not include all forms in 
cultivation. Until a careful study has been made of varieties, how- 
ever, this nomenclature will enable spawn makers to keep in mind 
certain types, and will make it possible for growers to ask for a 
spawn yielding a color demanded by their special markets. 


THE VITALITY OF MUSHROOM SPAWN. 


Many of the early experiments in mushroom growing undertaken 
by the writer were made in the hope of being able to ascertain the 
more frequent causes of failure and some of the chief difficulties 
encountered by American mushroom growers. The ordinary com- 
mercial spawn used by amateurs, that is, such as is obtainable upon 
the market during the winter months, was purchased wherever possi- 
ble. Samples of this spawn were placed under conditions which were 
supposed to be most favorable for growth. Nevertheless, in the 
majority of cases there was no indication of the development of a 
new mycelium from the bricks of spawn thus obtained. From these 
results it was suspected that much of the spawn which reaches the 
amateur grower may be considerably injured, or even killed, by 
transportation or improper conditions of storage; for it must be sup- 
posed that most of this spawn is in good or at least fair condition 
when exported from Europe. 

Subsequently the writer was able to look into the matter of spawn 
making in Europe and France, and he was convinced that the diffi- 
culty of securing good spawn in England is not a very serious factor. 
The same is true with reference to the material which is obtained by 
both extensive and small growers in France. 

Special importations of some of the commercial English and 
French spawns were made, and this was packed, shipped, and stored 
under conditions as favorable as may ordinarily obtain. This spawn 
was imported during midwinter and stored until March or early 
April, when it was used in spawning some experimental beds. The 
conditions of the experiments were practically the same throughout, 
yet in not more than half the beds was there a favorable development 
of mushroom spawn. A distribution of the French spawn, both the 
commercial flake and the improved cake spawn, was made to several. 
prominent American growers. Some of these growers experienced 
entire failure, while others reported that, after a slow beginning, beds 
spawned with this material made a good yield. The general conclu- 
sion, reenforced by observation and by the experience of practical 
growers, could only be that a large percentage of loss in mushroom 


VITALITY OF MUSHROOM SPAWN. ; 59 


growing is attributable to the injury suffered by the spawn after its 
preparation. This conclusion has been further strengthened by the 
experience of the past three years. From Table VIII, beds Nos. 1, 
2, 4, 5, and 30, it will be seen that, under conditions where fresh spawn 
has invariably made a good yield, the spawn which is more than a 
year old is, for the most part, seriously injured or killed. - To be 
exact, in only one case was there any production of mushrooms by 
spawn which had been kept for a year or longer. It must be said that 
no attempt was made to keep these spawns under similar conditions or 
under the most favorable conditions. For the most part the spawn 
was stored in the dry laboratory room, in which the temperature was 
more or less variable, but never extreme. The old American spawn 
which was used in experimental bed No. 1, in Table VIII, was stored 
in a basement room where the average temperature was undoubtedly 
cooler than that of the laboratory room. 

From experimental beds Nos. 1, 5, 4, and 5, in Table X, it is again 
seen that old spawn is unreliable. In this particular case the mate- 
rial was furnished by a prominent mushroom grower—an English 
spawn importer. This spawn had been stored in a dry house and 
was therefore subject to similar conditions. In Table VIII, beds 
Nos. 31, 32, 35, and in Table IX, Nos. 34, 41, and 45, there is further 
proof of the loss of vitality in the imported spawn ordinarily offered 
for sale in many of our cities. In these cases spawn was bought on 
the market just as offered for sale to the amateur buyer; “ best on 
hand ” was asked for, but no stipulation was made that is should be 
of recent importation, and no guaranty was asked. The tests were 
not, therefore, to compare the very best English with the best Amer- 
ican spawn, but merely to secure an indication of some of the causes 
of failure by the purchase at random of English and French spawn 
on the market. Even in times past the extensive mushroom growers 
have either imported their spawn direct, or made sure that they 
were obtaining the best product that the market could furnish. 
Unfortunately, it has not been possible to compare, in any experi- 
ments thus far concluded, the best English with the best American 
spawn. 

The results seem also to indicate that brick spawn maintains its 
vitality longer than the flake material, and that brick spawn made 
of loose, light material is less retentive of vitality than that made 
after the formula commonly followed in England. This proves to 
be an unfortunate factor to be dealt with in the attempt to reduce 
by all means the weight of the brick. The reduction in weight 
would be most desirable, since freight upon this material adds con- 
siderably to the price of market spawn. To the poor keeping quali- 
ties of loose spawn is perhaps due the large number of failures with 
French flake spawn, and perhaps also some of the failures with the 


60 . MUSHROOM GROWING AND SPAWN MAKING. 


newer form of French brick spawn. The latter is made in the form 
of very small, thin bricks, which are unquestionably more afiedted 
by weather conditions than the larger English bricks. 

These results have seemed to demand that special attention should 
be given to methods of spawn making in the United States in order 
that growers might be able to secure this product as fresh as -pos- 
sible. Moreover, it was desirable, as previously indicated, to at- 
tempt work leading to the selection and improvement of varieties. 
The success of the work in spawn making has been almost all that 
could have been anticipated. By the pure-culture methods described, 
several firms are now making grades of brick spawn which have 
yielded remarkably well. This fact is now thoroughly recognized 
by a large number of the best growers throughout the country. 
Probably as many as 50,000 bricks were sold during 1904, and it is 
perhaps to be expected that several hundred thousand will be sold 
during the present year. 

It is to be regretted that it has not yet been possible to abandon 
the pure-culture process by means of which the virgin spawn is made 
while retaining the advantages of selection. Nevertheless, it should 
be remembered that the very difficulties of this process insure its use 
only by those who are able to give it their best attention and who will 
doubtless develop it to the fullest commercial extent. It has not been 
supposed by the writer that the work thus far accomplished will en- 
able all mushroom growers to manufacture their own spawn with 
comparative ease. In other phases of horticultural work it is not so 
much to individual growers as to progressive seedsmen that we look 
for the best seed of improved varieties. The same thing apparently 
must be anticipated in the development of the mushroom industry. 
The growing of selected spawn may, in general, become a specialized 
process. ° 

Nevertheless, it is believed that in time a method of spawn pro- 
duction from spores without pure-culture precautions will be devel- 
oped. The necessity of developing immediately, or placing on a 
practical basis, the pure-culture process has temporarily directed the 
experimental work along other lines. 


O 


Bul. 85, Bureau of Plant Industry, U. S. Dept. of Agriculture. PEATEs. 


Fig. 1.—A FINE CLUSTER OF AGARICUS CAMPESTRIS, THE HORTICULTURAL VARIETY 
COLUMBIA. 


Fic. 2.—MorRELS (MORCHELLA ESCULENTA), ONE OF THE FINEST EDIBLE FUNGI. 


Bul. 85, Bureau of Plant Industry, U. S. Dept. of Agriculture. PLATE III. 


Fic. 1.—AGARICUS FABACEUS, THE ALMOND-FLAVORED 
MUSHROOM. 


FiG. 2.—AGARICUS VILLATICUS, A PROMISING SPECIES, FLESHY AND PROLIFIC. 


Bul. 85, Bureau of Plant !ndustry, U. S. Dept. of Agriculture. PLATE IV. 


Fig. 1.—A YOUNG SPECIMEN OF THE COMMON PUFFBALL 
(CALVATIA CRANIIFORMIS). 


Fic. 2.—THE OysTER MUSHROOM (PLEUROTUS OSTREATUS), GROWING ON DECAYED 
WILLOW Loa. 


Bul. 85, Bureau of Plant Industry, U. S. Dept. of Agriculture. PLATE V. 


Fic. 1.—A MUSHROOM HOUSE PROVIDED WITH GAS-PIPING FRAMEWORK 
FOR SHELF BEDS. : 


Fic. 2.—THE PREPARATION OF COMPOST. 


Bul. 85, Bureau of Plant Industry, U. S. Dept. of Agriculture. PLATE VI. 


; 


Fic. 2.—THE METHOD OF MAKING PURE CULTURES, SHOWING THE APPARATUS AND 
MATERIALS. 


Bu!. 85, Bureau of Plant Industry, U. S. Dept. of Agriculture. PLATE VII. 


FiG. 1.—MUSHROOMS PREPARED FOR THE AMERICAN MARKET. 


Fia. 2.—Goop (‘‘WELL-RUN’’) MUSHROOM SPAWN, BRICK FORM. 


‘SATV¥q 3LVd 4O SN3GY¥V5 NAYNNS GNV SANNQ GNVS ONIMOHS ‘GaNO 14 JO NMOL 3H1L WOYNS NOIDZY ANOS GINO 3H1L 4O MBIA WHAN3H 


= 


PLATE I. 


oO 


) 


‘U. S. DEPARTMENT OF AGRICULTURE. 
BUREAU OF PLANT INDUSTRY—-BULLETIN NO. 86. 


B. T. GALLOWAY, Chief of Bureau. 


AGRICULTURE WITHOUT IRRIGATION IN THE 
SAHARA DESERT. 


BY 


THOMAS H. KEARNEY, 
PHYSIOLOGIST. 


VEGETABLE PATHOLOGICAL AND PHYSIOLOGICAL 
INVESTIGATIONS. 


IssuED NOVEMBER 16, 1905. 


WASHINGTON: 
GOVERNMENT PRINTING OFFICE. 
1905. 


BUREAU OF PLANT INDUSTRY. 


B. T. GALLOWAY, 
Pathologist and Physiologist, and Chief of Bureau. 

VEGETABLE PATHOLOGICAL AND PHYSIOLOGICAL INVESTIGATIONS. 
ALBERT F. Woops, Pathologist and Physiologist in Charge, Acting Chief of Bureau in 
Avsence of Chief. 

BOTANICAL INVESTIGATIONS. 

FREDERICK V. COVILLE, Botanist in Charge. 


FARM MANAGEMENT. 
W. J. SPILLMAN, Agriculturist in Charge. 


POMOLOGICAw INVESTIGATIONS. 
G. B. BrRacKEetT?r, Pomologist in Charge. 


SEED AND PLANT INTRODUCTION AND DISTRIBUTION. 
A. J. PIETERS, Botanist in Charge. 


ARLINGTON EXPERIMENTAL FARM. 
L. C. Corspett, Horticulturist in Charge. 


INVESTIGATIONS IN THE AGRICULTURAL ECONOMY OF TROPICAL AND SUB- 
TROPICAL PLANTS. 


O. F. CooK, Bionomist in Charge. 


DRUG AND POISONOUS PLANT INVESTIGATIONS, AND TEA CULTURE 
INVESTIGATIONS. 


Ropney H. True, Physiologist in Charge. 
DRY LAND AGRICULTURE AND WESTERN AGRICULTURAL EXTENSION. 
Cart S. SCOFIELD, Agriculturist in Charge. 


EXPERIMENTAL GARDENS AND GROUNDS. 
E. M. Byrnes, Superintendent. 


SEED LABORATORY. 
Evcar Brown, Botanist in Charge. 


J. E. ROCKWELL, Editor. 
JAMES E. JONES, Chief Clerk. 


VEGETABLE PATHOLOGICAL AND PHYSIOLOGICAL INVESTIGATIONS. 
SCIENTIFIC STAFF. 
ALBERT F. Woops, Pathologist and Physiologist in Charge. 


Erwin F. Smitu, Pathologist in Charge of Laboratory of Plant Pathology. 

HERBERT J. WEBBER, Physiologist in Charge of Laboratory of Plant Breeding. 

WALTER T. SWINGLE, Physiologist in Charge of Laboratory of Plant Life History. 

NEWTON B. Pierce, Pathologist in Charge of Pacific Coast Laboratory. 

M. B. Waite, Pathologist in Charge of Investigations of Diseases of Orchard Fruits. 

MARK ALFRED CARLETON, Cerealist in Charge of Cereal Laboratory. 

{[ERMANN VON SCHRENK, in Charge of Mississippi Valley Laboratory. 

Vv. H. Rours, Pathologist in Charge of Subtropical Laboratory. 

C. O. TOWNSEND, Pathologist in Charge of Sugar Beet Investigations. 

T. H. Kearney, A. D. SHAMEL, Physiologists, Plant Breeding. 

P. H. Dorsett,* CorNeELIus L. SHEAR, WILLIAM A. ORTON, W. M. Scott, ERNsT A. 
Bessey, BE. M. FREEMAN, Pathologists. 

Kk. C. Cuiitcotrr, Papert in Cultivating Methods, Cereal Laboratory. 

C. R. BALL, Assistant Agrostologist, Cereal Laboratory. 

JoseruH S. CHAMBERLAIN,” J. ARTHUR LE CLERC,° Physiological Chemists. 

FLora W. PATTERSON, Mycologist. 

CHARLES P. HARTLEY, KAR” F. KELLERMAN, JESSE B. Norton, CHARLES J. BRAND, 
T. RALPH ROBINSON, Assistants in Physiology. 

DeaNe B. SwWINGLe, GeorGe G. Hepecock, Assistants in Pathology. 

ERLEY SPAULDING, P. J. O'Gara, FLORENCE HepGES, Henry A. MILLER, ERNEST B. 
Brown, Lesuigz A. Firz, LEonaRD L. HarTER, JOHN O. MeRWIN, A. H. LEIpIGH, H. F. 
BLANCHARD, Scientific Assistants. 

W. W. Copsey, Tobacco Papert. 

JOHN VAN LEENHOFF, Jr., T. D. BECKWITH, Esrperts. 


“Detailed to Seed and Plant Introduction and Distribution. 
» Detailed to Bureau of Chemistry. 
© Detailed from Bureau of Chemistry. 


to 


LETTER OF TRANSMITTAL. 


U. S. DEPARTMENT oF AGRICULTURE, 
Bureau or Pirant Ixpustry, 
OFFICE OF THE CHIEF, 
Washington, D. C., September 6, 1905. 

Sir: I have the honor to transmit herewith and to recommend for 
publication as Bulletin No. 86 of the series of this Bureau the accom- 
panying paper, entitled “Agriculture without Irrigation in the Sahara 
Desert.” 

This paper was prepared by Mr. Thomas H. Kearney, one of the 
physiologists of this Bureau, and the data for it were obtained on a 
trip which he made to northern Africa for the Office of Seed and 
Plant Introduction and Distribution, primarily for the importation 
into the United States of offshoots of valuable Tunisian date varieties. 
It is believed that the methods described may be useful in some cases 
in our southwestern desert regions where date culture is being intro- 
duced. 

The accompanying illustrations are necessary to a clear understand- 
ing of the text of this bulletin. 

Respectfully, B. T. Gattoway, 
Chief of Bureau. 
Hon. James WIson, 
Secretary of Agriculture. 


PREPAC E. 


In view of the interest in farming without irrigation that is now 
being manifested in the arid portion of the United States, an account 
of a region where agriculture is carried on under extremely adverse 
natural conditions is particularly timely. The present paper deals 
with a highly developed system of date-palm culture in the Oued 
Souf, a remarkable and little-known part of the Sahara Desert in 
northern Africa. Strictly speaking, it is not dry-land agriculture 
with which we have to do in the Souf region, for while the rainfall 
is practically nothing and irrigation is impracticable, the roots of the 
trees quickly find their way to ground water. However, it is quite 
possible that similar conditions may be found to exist in this country 
in some parts of the desert region of the Southwest, and that the Souf 
system, with or without irrigation, can be utilized there on a small 
scale in growing certain orchard crops with a view to forcing fruit 
to early maturity, so that it can be put upon the market much in ad- 
vance of the bulk of the crop. 

The Oued Souf was visited by Mr. Kearney at the end of Novem- 
ber, 1904, the journey having been made from Nefta, in southwestern 
Tunis, where he had spent several weeks in a study of the date palm. 
This expedition to northern Africa was made under the auspices of 
the Office of Seed and Plant Introduction and Distribution of the 
Bureau of Plant Industry. 

Acknowledgment is here made to Captain Bussy, Chef du Bureau 
Arabe at El Oued, for the cordial assistance rendered by him to Mr. 
Kearney during the latter’s stay in the Souf region. 

A. F. Woops, 
Pathologist and Physiologist. 
OFFIcEe OF VEGETABLE PATHOLOGICAL 
AND PrystoLocicaAL INVESTIGATIONS, 
Washington, D. C., August 22, 1905. 


oO 


. ay 
rh 
ae 


COS EEN TS: 


PREM ROREMIg GT OLIMEES Revenant ne ae ek EO ah eee ok Cate 


Population 


Climate __- 


OOO RGSS. TLIC EY 2 Sane a ss en a a AS ee Par 


Sols. 2 


Pitrem ceca iOel Smears O54 ie. Bee A ney (an Ss oe AD a Be 
TEV amici ae Se a oes 28d OA ee TOR SE Ret aed pe ee em 
CHire® Ont JORIS Shee ae eRe A ee Se 5: eek eee aS Eee ee Bee 
Pa ORME TS ATIC es xia) oe eee Or Rereant Ia wk Rot Vy SN StL ees EEE 
2 ETE ETI AS pe = a eae oy 52 ee ge ae eee ee ee 
{EIST GLa SE Se pee ene te Oe ae Pe ee ape (pe Be ee ee ema ea Semen Sc 


Yields - 


Rermeite cn cmie nyo ONIN. 68 - rin ti) ee ee ee eee tans eA ye ty Ue Soe sens Dea 


Conclusion 


DEA MRED S es os oo NN Grey OS te oe 


7017—No. 86—05 M 


2 7 


w 2 


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~F Ot Ct H © 4 


Co W wW 


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PuatTE I. 


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III. 


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Ve 


ILLUSTRATIONS. 


PLATES. 


General view of the Oued Souf region from the town of El Oued, 


Page. 


showing sand dunes and sunken gardens of date palms__Frontispiece. 


Fig. 1.—High sand dunes east of El Oued. Fig. 2.—A typical 
dwelling house of the Oued Souf, showing its cubical form and 
roof composed of flattened cupolas. Fig. 3.—General view of 
the Oued Souf region, showing sunken date gardens and sand 
Gunes. ooo cn sodas 2222 Seo eee soe 

Fig. 1.—Near view of sunken palm garden and surrounding dunes. 
Fig. 2.—Gradual extension of a palm garden by cutting down bor- 
dering sand hills. Fig. 3.—Vegetable garden irrigated by well 
near bottom of basin in which date palms are grown ---------- 

Fig. 1.—Hole on slope of dune near bottom of basin in which a 
young palm is planted. Fig. 2.—Camel manure ready for appli- 
cation in a date garden --. .. 2.225. 22... 52222502 =ee= eee 

Fig. 1.—‘t Dokana,”’ or mound of earth and plaster for strengthen- 
ing the baseof apalm. Fig. 2.—Rhars palm, showing thickness 
of trunk . 22.2. -.-22--2 2 eee 


TEXT FIGURE. 


Fic. 1. Map showing location of the Oued Souf with respect to other locali- 


ties in Algeria and Tunis 
5 


30 


30 
30 


30 


10 


B. P. I.—185. V. P. P. I.—144. 


AGRICULTURE WITHOUT IRRIGATION IN 
THE SAHARA DESERT. 


INTRODUCTION. 


In the great desert of northern Africa, stretching across in a belt 
from southeastern Algeria to the borders of Tripoli, is the region 
known as the “ Erg.” It is a land of enormous sand hills, some of 
which reach a height of 500 feet. Chain after chain of these great 
dunes, with knife-edge summits and steep slopes and trough-like 
valleys between, extend diagonally northeast and southwest across 
this part of the Sahara. (PI. II, fig. 1.) It is like an ocean caught 
in a raging storm, with its huge billows rising skyward and held fixed 
and motionless. Not a Jeaf nor a blade of grass, not a bowlder nor 
a pebble mars the smoothness of the sand. Never is the least trace 
of water to be seen on its surface. The few drops of rain that fall 
at rare intervals are drunk up as soon as they touch the thirsty 
ground. Pure quartz sand it is, ight yellow in color and so fine of 
grain that the least breath of air sends a little cloud of it curling off 
the sharp crests of the ridges. When a hard wind blows the air is 
filled with it, the sun is blotted out at noonday, and the traveler can 
hardly see his horse’s head in front of him. The sharp-cornered 
particles of sand sting his face and blind and bewilder him. The 
vague tracks of camels and donkeys, the only roads through this 
wilderness, are quickly covered up, all landmarks disappear, and 
without an experienced guide one is sure to be hopelessly lost. 

It is a desolate and unfriendly landscape, yet at times not without 
a weird beauty of its own. When the sun is high the glare is blind- 
ing and there is little to attract one in the scene. But in the early 
morning and the late evening the sand assumes a golden color, and 
the dense black shadows cast by the dunes bring out their contours 
in sharp relief. Then their surface is seen to be modeled by series 
of delicate ripple marks left by the wind, and one finds it hard to 
believe that when he climbs the next high ridge he will not see the 
ocean at his feet. 


10 AGRICULTURE IN THE SAHARA DESERT. 


Who would suspect that amid these mountains of bare sand, where 
even the hardy shrubs and grasses of the desert find no foothold,* 
men can live by the products of the soil? Yet in the very heart of the 
Erg, two long days’ ride east or west from the nearest habitations, 
there exists one of the most highly developed agricultural communi- 
ties in the world. This is in the country known as the Oued Souf, 
situated in extreme southeastern Algeria (see fig. 1), about midway 
between the oases of southwestern Tunis and the Algerian oases 
known as the Oued Rirh, in which latter the date palm is grown by 


SOUF 


e 
Ourlana “1 Oued 


e7ougourts 


R A| 


Fic. 1.—Map showing location of the Oued Souf with respect to other localities in Algeria 
and Tunis. 


Europeans upon a commercial scale.’ From El Oued, the capital of 
the Souf, it is about 70 miles southwest to Tougourt, the chief town 
of the Oued Rirh, and about the same distance northeast of El Oued 
is Nefta, the nearest oasis in Tunis. The elevation of El Oued is 
ubout 257 feet above sea level. 

aQOnly eight species of flowering plants were found growing wild in the Souf 
region by Massart. See “ Un voyage botanique au Sahara,” p, 249 (1898). 

4 See Plate II (map) in Bul. 58, Bureau of Plant Industry, “ The Date Palm,” 
by W. T. Swingle. 


INTRODUCTION. 11 


From the tops of the lofty sand hills that surround El Oued an 
excellent view can be had, and there one can form a clear idea of the 
character of this remarkable country. (See Pl. I, frontispiece, and 
Pl. II, fig. 3.) Assuredly there are few regions where any sort of 
agriculture is carried on under more extraordinary conditions. As 
Tar as the eye can reach it rests upon an expanse of pure sand, heaved 
up into range after range of dunes.“ In the hollows among these 
dunes are the gardens of date palms, sometimes mere pockets contain- 
ing 10 or 20 trees, sometimes larger basins in which are groves of 50 
to 100 palms. 

Often the bordering sand hills are much higher than the tallest 
of the palms, so that many of the gardens can not be seen until one is 
on the very edge of the basin. In other places, however, the ridges 
are lower or gaps occur, allowing a cluster of feathery crowns to 
peep through. These are of such a dark green as to look almost black 
against the pale sand. <A more striking contrast of colors could not be 
imagined. The trunks are rarely seen until one reaches the brink of 
the basin or pocket in which the palms are growing. 

We have before us, in short, a network” of basins or hollows con- 
taining small groves of date palms and separated by great hills and 
ridges of sand. The aspect of the country is wholly different from 
the Oued Rirh in Algeria and the Djérid in Tunis, where each oasis 
is a dense continuous forest of date palms, containing often several 
hundred thousand trees,’ and situated upon Gomparatively level land. 

Such, then, is the country of the Souf, a land where there is prac- 
tically no rainfall, where there are no streams nor springs nor flowing 
wells to furnish water for irrigation, where the soil is a pure hard 
sand devoid of organic matter and blown about in clouds by every 
wind, so that unceasing vigilance is needed to keep the gardens free 
from it, and where the summer heat is almost as great as anywhere 
in the world. Yet here the date palm grows to perfection, yielding 
fruit of better quality and in larger quantity than elsewhere in the 
Sahara. How this has been brought about we shall presently see in 
these pages; but first we should know what manner of men are they 
who have developed a flourishing agriculture in a land where Nature 
seems to frown most severely upon all efforts to win a living from the 
soil. The race that has succeeded so well in the face of such tremen- 
dous obstacles must needs be an interesting one. 


aThe surface of these dunes is easily moved, even by a light breeze, but the 
core is said to be stationary and composed of stratified materials. 

b The Arab word “erg” (piural areg) means “a vein.” 

¢In 1899 there were only 192,000 date palms in bearing in the entire Souf 
region. 


12 AGRICULTURE IN THE SAHARA DESERT. 
POPULATION. 


There are about 25,000 people in the Souf region, 5,500 of whom 
inhabit the capital town, El Oued, and its immediate neighborhood. 
Several distinct tribes are included in this population, some being 
chiefly nomadic shepherds, and others more sedentary, devoting most 
of their time to the care of their gardens. They are for the most part 
a healthy and strong looking race, and are much more energetic than 
the inhabitants of other north African oases. This is doubtless 
partly due to the unceasing labor demanded by the conditions under 
which they live and partly to the fact that their climate is a healthful 
one, despite the intense heat of summer. There is no standing water 
nor even moist surface soil, and mosquitoes are said to be unknown. 
The dry air and the hot sand are not friendly to the germs of con- 
tagious diseases. The conditions are, therefore, very different from 
those in other oases of the Sahara, which are so overirrigated as to be 
mere swamps 1n summer, scourged with malarial fevers. 

The inhabitants of the Souf region, who are called “ Souafas,” 
depend for a livelihood largely upon the products of their gardens, 
but they have other resources as well. The more nomadic tribes 
possess flocks of sheep and goats. They have almost a monopoly of 
the trade of camel drivers in a large part of the Sahara, guiding 
caravans eastward into Tunis, westward to Biskra, and far south 
into the heart of the great desert. Their camels are considered the 
largest and finest of the Sahara. The men of the Souf are indefati- 
gable walkers, thinking nothing of traveling 20 or 25 miles a day 
through the loose sand. Their camel’s-hair shoes, tightly bound 
around the ankles, are much better adapted to this sort of travel than 
the loose-fitting, heelless slippers generally worn by the Arabs. 

In building their houses, as in cultivating their palms, the Souafas 
have many difficulties to contend with that are not experienced by 
the dwellers in other oases. Elsewhere in the Sahara, sun-dried 
brick, like the Mexican adobe, is the universal building material. 
But in the Oued Souf there is no clay to be had. Consequently the 
town of El Oued and all the villages of the region are constructed 
with irregular masses of grayish, crystalline, gypseous rock, cemented 
by plaster made from the same material, which thus furnishes both 
stone and mortar. Wood being very scarce, the roofs are not flat, 
wooden ones, but consist of rows of small, flattened cupolas, not 
unlike old-fashioned beehives, which give a very odd look to the cube- 
shaped houses. (PI. II, fig. 2.) In its architecture, as in everything 
else, the Souf is unique. The immaculate cleanliness of the villages 
is surprising to the traveler who is familiar with the filthy streets of 
most Arab towns. The pure, dry sand is constantly drifting among 
the houses, and quickly buries all refuse. 


CLIMATE, 13 


CLIMATE. 


Exact data in regard to the climate of the Oued Souf are not easily 
obtainable. The observations here given were made chiefly by a 
medical officer of the French army during the summer of 1884." 

‘Unfortunately records covering a period of several years: are not 
available.” 

The summer temperatures are very high, few hotter localities being 
known in the Sahara. The monthly maximum shade temperatures 
observed by Escard in the summer of 1884 are as follows (in degrees 
Fahrenheit) : 


evita 2 ee eons Gael eA OS bese se er nen 116.5 
COLEUS oa 2 oe Na ea ae LOOT SCP tempers rao ee ee Lee 115 
SI 2 ae ee ee ee ee OGA | AO CLO DETR eas 2522 a eT 91.5 
LGN? 2 SS 122 


In June, 1904, a maximum of 127.5° F. is said to have been reached. 
The sum total of temperature during the summer, a factor of the 
greatest importance in the ripening of the finer varieties of dates, is 
said to be greater in the Oued Souf than in the Oued Rirh and the 
Djérid. At the time of the writer’s visit (November 22-26, 1904) 
cool, cloudy weather prevailed, and the nights were decidedly cold. 


a@Wscard, Etude médicale et climatologique sur le pays de l’Oued Souf. 
Archives de Méd. et de Pharm. Milit., 7: 33 (1886). 

b Since the above chapter on climate was written the records of observations 
made at El Oued during the whole of 1904 and parts of 1908 and 1905 have been 
received, through the courtesy of the Director of the Meteorological Service of 
Algeria. These observations necessitate some modifications in the previously 
written discussion of the climate of the Souf. The absolute maximum temper:- 
ture at El Oued in 1904 was 121.8° F. in May instead of 127.5° F. in June. The 
absolute minimum in 1904 was 32° F. The mean relative humidity during 190+ 
at El Oued was 58.8, which is lower than the normal at Tozer, in Tunis (60.8), 
but higher than the normal at Ouargla and Biskra, Algeria (47.2 and 48.4, re- 
spectively), and at Yuma, Ariz. (42.9). The sum of the monthly means of 
evaporation at El Oued in 1904 is 156 inches, while the normal at Tozer is 94 to 
98.5 inches. The total precipitation at El] Oued in 1904 was 3.25 inches, while 
the normal yearly total is 3.61 inches at Ouargla, 6.75 inches at Biskra, 5.1 
inches at Tozer, and 2.83 inches at Yuma. 

The observations on the prevailing direction of the wind during 1904 and the 
first half of 1905 do not agree with those made by Escard and quoted in the 
text, but the data are insufficient for an adequate discussion of this factor. 

As the climatic factor which is most important in date growing is probably 
the effective temperature during the ripening season, a calculation has been made 
of the sum totals of daily mean and of daily maximum temperatures above 
64.4° F. (18° C.) during the months of May to October, 1904, at El Oued, and, 
for the sake of comparison, of the sums for the same months of the same year 
at Ouargla and Biskra, Algeria; Tozer, in Tunis, and Mecca, in the Salton 
Basin, California. Records at Mecca for May and June, 1904, not being avail- 
able, the records for those months at Imperial, Cal., were substituted in making 


14 AGRICULTURE IN THE SAHARA DESERT. 


Almost every winter freezing temperatures are reached, although 
probably the minima are higher and frosts are less frequent in the 
Oued Souf than at Tougourt and at Tozer. In the winter of 1903-4 
the absolute minimum was 32° F., and for several preceding winters 
Nah es 

In respect to atmospheric humidity, the absence of surface water 
probably tends to keep the air drier than is generally the case in the 
oases of the Sahara. 

The rainfall is said to be even less than in the Oued Rirh, where 
the average yearly precipitation (at Tougourt) is only 5.3 inches. 
Most of the rain is divided between two periods—October to Novem- 
ber, and February to, March. The rains are generally torrential in 
character and fall during several successive days, with intervals of 
sunshine. 

The sky is nearly always clear in summer. Toward the end of 
August light clouds appear in the morning and evening, but no rain 
falls until October. During the writer’s four days’ visit to the Souf 


the calculations for Mecca, experience having shown that the temperatures 
recorded at Mecca and at Imperial are very nearly identical. 


TABLE I.—Sum of daily mean temperatures above 64.4° F. (18° C.) from May 1 to October 


31, 1904. 
| Degrees | Degrees 

Locality. Fahren- centi- 

heit. grade. 
Wl Oned, -Aleerig 23. en oo cate ee ace 3, 672.6 2,040.3 
Ouargia, Algeria. 2t f= See ern PND Tae EN Soot nis every 3, 961.3 2,200.7 
Biskra,. Algeria. 2. 5222 hw a eee 3,140.4 1,744.6 
Tozer, "Cunis. 2.2 -:. .- 2. te eens he olen oe ea A 3,831.3 2,128.5 
Meeca, Cal o.- faeces BS eee 3,591.5 1,995.3 


The sum of daily means at Biskra for 1904 is nearly 200° F. lower than the 
normal, as based upon observations covering a period of 10 years (see W. T. 
Swingle, Bul. No. 53, Bureau of Plant Industry, p. 66), and the sum at Mecca 
for that year is about 400° F. lower than the normal for Indio, Cal., a few miles 
distant. 


TABLE II.—Sum of daily marimum temperatures above 64.4° F. (18° ©.) from May 1 to 
October 31, 1904. 


| Degrees Degrees 


Locality. | Fahren- centi- 

heit. grade. 
Ml Oued, Algerians 225 20 dBRe acoso dete een ec oss Jo eth ee andes ee ee 6,325.7 8,514.3 
Ouargla, Algeria: > <5. HS ons. c a cset wucem ote eee oe eee 6,501.6 3,612.0 
Biskra, Algeria . US GAS he ae 2c Ded 26550 Veale eect oivinncire i ales ieee 5, 219.1 2,899.5 
Pozer, "TUNG 25 an ae. ew als we ook een Sac e en ee 5,908.6 3,279.8 
Macen, Cal oso. oa coe nc peers pandddobceve Wade Onno eee eee oe 6,370.1 3,539.0 


In calculating the sums of daily maximum temperatures allowance was made 
for mean monthly minima falling below 64.4° F., in accordance with the prac- 
tice suggested by Swingle (ibid., p..67). The sum of daily maximum tempera- 
tures at Biskra during May to October, 1904, is about 250° F. lower than the 
normal for 12 years (Swingle, ibid., p. 68). The sum for Mecca is about 700° 
Flower than that for Imperial, Cal., in 1902. ; 


CLIMATE. £5 


at the end of November the sky was overcast about half of the time, 
and there were occasional gusts of cold, drizzling rain. 

Winds are probably more frequent and more violent in the Oued 
Souf than in the other groups of oases mentioned. It would appear, 
in fact, that windiness is the ordinary condition there. During the 
winter, northwest and northeast winds prevail. From April to 
October, however, the wind is generally from the south (the sirocco) 
or the southeast (the simoom). The sirocco is the hottest wind, but 
is less frequent than the simoom, which is generally more violent and 
transports more sand. All these are winds that blow more or less 
steadily for several hours and often days at a time. Cyclonic sand 
storms also occur, arising suddenly and lasting but a short time. 
Such storms are never accompanied by rain. 

Owing to the lack of natural vegetation (see Pl. IT), and the fine- 
ness of the sand with which the country is covered, strong winds 
carry with them a great deal of material, so that the face of the land 
is being constantly altered. This is shown by the fact that an 
apparently fresh wagon track noticed by the writer on his journey 
from Nefta was completely buried in many places by large heaps of 
sand. During his four days’ stay in the Souf country, a strong wind 
blew constantly, often making travel difficult, as the particles of sand 
stung the face and made it hard at times to keep the eyes open. The 
air was frequently so full of sand that one could see but a few rods 
ahead. 

The slopes of the dunes that border the gardens are very steep, so 
that when a heavy wind is blowing much sand rolls down upon the 
floor of the garden. Generally there is a fence or palisade along the 
crest of the dunes, made by sticking palm leaves or pieces of gypsum 
rock close together (PI. I, fig. 3; Pl. III, fig. 1; Pl. IV, fig. 1), but 
this only partly arrests the blowing and drifting sand, and it is nec- 
essary to remove it frequently from the gardens. The task is a labo- 
‘rious one, as the sand must be carried up the steep hillside in baskets 
and dumped on the outer slope of the dunes. But if it were neglected, 
in a few years the trees would be buried, especially in smaller gardens. 
The writer saw several little gardens that had been abandoned by 
their owners where the basin was almost filled and only the crowns 
and a small part of the trunks of the trees still projected above the 
soil. 

Another injurious effect of the sand-carrying winds is that when 
harvested the dates always have more or less sand adhering to the 
skin, and this must be brushed or washed off before they are fit for 
export. Dates that had been kept for some weeks in the houses of 
natives, and even those freshly gathered from the trees, were very 
unpalatable to the writer on this account; although the Souafas them- 
selves do not seem to mind eating a good deai of sand with their dates. 


16 AGRICULTURE IN THE SAHARA DESERT. 
WATER SUPPLY. 


There is no surface water in the whole Souf country, excepting, 
possibly, a small sebka, or salt pond, of which the writer was told, 
but which he did not see. There are no natural springs, although 
eround water is everywhere very near the surface in the hollows 
among the dunes. It is said to occur sometimes in strata of pure 
quartz sand, sometimes in gypseous sand. The distance to standing 
water is said to reach as much as 40 feet in different parts of the 
region, although averaging considerably less; but in the bottoms 
of the basins in which date palms are grown it is encountered often 
at a depth of only 2 or 3 feet below the surface of the soil, thanks to 
the extensive excavation that has been done. In one garden, near 
the town of E! Oued, the writer saw water standing at a depth of 6 
feet in a large hole that had been dug to receive manure. The Souf 
oases are believed to mark the course of a buried Quaternary stream ; 
Oued Souf means “ murmuring river.” ¢ 

As we shall presently see, the date palm is not irrigated in the Souf 
country, receiving at most a few waterings by hand during the first 
summer after planting. In almost all the gardens, however, shallow 
wells occur,’ the water of which is used for household purposes and 
for irrigating small plats of garden vegetables. (PI. III, fig. 3.) 
These are generally situated on the slope of the bordering dunes, 10 
feet or less above the bottom of the basin, and water stands in them 
at a depth of 10 to 16 feet. In the town of El Oued the wells are 
much deeper than in the gardens, water standing in them at 30 to 40 
feet. All this water is under a slight pressure, rising in the wells 
about 1.5 feet higher than the general water table. Small gardens of 
vegetables and tobacco,’ irrigated from deeper wells, are also located 
in some parts of the region far above the bottoms of the basins. Prac- 
tically no grain is raised, wheat and barley being brought by caravan 
from other parts of Algeria and from Tunis, to be exchanged for 
dates. 


a{t is difficult to obtain a very satisfactory idea as to the distribution of the 
ground water in the Souf region, its depth at various points, and the amount of 
excavation necessary to enable the roots of the palms to reach it easily. The 
natives themselves give the most conflicting answers to questions upon this 
subject, and there are serious discrepancies in the accounts that have been 
published by French authorities upon irrigation. The whole matter evidently 
needs to be carefully studied by competent hydrographers, a study which is cer- 
tainly warranted by the rarity of this type of agriculture. 

>In 1888 Rolland estimated that there were 4,431 wells in the Souf region. 

¢ Tobacco growing, which is unrestricted in Algeria, is a profitable industry in 
the Souf country on account of that region’s nearness to the frontier of Tunis, 
where the growing of this crop is forbidden by law and where the selling of 
tobacco is controlled by the government. Agents of the Tunisian tobacco 
monopoly frequently visit the Oued Souf to purchase supplies. 


WATER SUPPLY. 17 


The plats of vegetables that are irrigated from wells in the date 
gardens are situated on terraces constructed in the side of the sand 
hills, usually 10 feet or less above the floor of the basin. (PI. IIT, 
figs. 1 and 3.) The well water is raised by hand in a shallow bucket, 
generally made of basketware covered outside with pitch, but some- 
times of goatskin, which is hung on the small end of a slender palm 
trunk and counterpoised by a piece of rock fastened to the large end. 
The pole is fastened by its center to a crosspiece that is supported by 
two vertical posts made of stouter palm logs or of cemented rock. The 
bucket is emptied into a little cement basin adjoining the well curb, 
whence the water flows through a system of small conduits into the 
plats that are to be irrigated. Flood irrigation alone is practiced. As 
there is no soil in the region from which ditch banks and ridges that 
will stand up when wet can be made, the conduits and ridges of the 
plats, as well as the lining and curb of the well, are made of the 
same dark-gray plaster with which the walls of the houses are 
cemented. Plugs of wool are used for stopping the conduits at places 
where water is to be diverted into the plats. Among the vegetables 
most commonly grown are cabbages, turnips, radishes, carrots, pump- 
kins, melons, watermelons, onions, tomatoes (a small-fruited sort), 
and peppers. 

In parts of the Souf region, especially east of the capital town, 
El Oued, the water of the wells is said often to contain enough mag- 
nesium and other salts to make it disagreeable for drinking.*. West 
of the town, on the other hand, the water is said to be generally very 
pure. The difference is thought to be sufficiently great to have a 
marked effect upon the quality of the dates, the most renowned 
Deglet Noors of the Souf region being produced near the village of 
El Amiche, where the water is purest. The peculiar character of 
the water supply of the Oued Souf is not without advantages. Emi- 
nent authorities are of the opinion that the underground sheet is 
abundant and that it is much less lable to exhaustion than in the 
Oued Rirh, where numerous flowing artesian wells exist. 


a Well water in the Souf, according to-an analysis cited by Jus (Les oasis 
du Souf du Département de Constantine, Bul. Acad. d’Hippone, No. 22, p. 67 
(1886), has the following contents of solid matter in grams per liter of 
water : 


SUE SVGREITREYS) SSE TEE BS ee ee er ye ee ee eh eee oe be pk A Oe ae en gee ees 1.99938 
CIA GTS EGIS Sg CSS tS ee ee Ree eee ey ee Se eee ee . TT69 
“Tai OLIGO Sh if = oS See aa ee eee pers te A See eee ee . 2999 
MABE EeSe ang, CISSOlved: Organic Matters. = oe Se Bee ee . 0690 
Silicates, etc., in suspension______. Ns Ss es st eee . 0335 

I) Sa) seg et 8 Ne > Se ee ee A ke ee eg 3. 1786 


Schirmer (Le Sahara, p. 261) states that the mean sult content of well water 
at El] Oued is 2.77 grams per liter. 


18 AGRICULTURE IN THE SAHARA DESERT. 
SOILS. 


The soil of the whole Oued Souf region is a fine-grained, light- 
yellow quartz sand, which is practically uniform in character to a con- 
siderable depth.*| Here and there beds of a coarse, rather soft, gyp- 
seous rock occur at a depth, it is said, of 10 to 20 feet*and in strata 
1.5 to 10 feet thick. The crystals of which this rocks composed 
are very large, often 1 foot long. They are often aggregated into 
masses which, on account of their shape, are known as “ Souf roses.” ” 
It is therefore a fair inference that the Souf soils are sufficiently 
rich in ime. They are very poor in organic matter and doubtless in 
nitrogen. Other data as to their composition are wanting. 

In the eastern part of the region the soil of all the gardens is said 
to be somewhat saline, and the writer was told that there is even a 
small sebka (salt pond) in that section, although he saw nothing of 
these conditions. There is said to be nowhere enough salt to injure 
seriously the palms themselves, but the yields of fruit are diminished 
by this cause, and the dates are somewhat smaller and of slightly 
inferior quality. Consequently, palms in full bearing in the gardens 
west of El] Oued are worth from two to ten times as much as those 
in the gardens east of that town. The Souafas do not pretend to 
distinguish some varieties of the date palm as being more resistant 
to salt than others, as do the inhabitants of the Djérid oases, where 
the salinity of the soil is often very pronounced. Neither have they 
adopted any special methods of preparing and handling salt land 
by drainage, flooding, or otherwise, as is the case in the Tunisian 
oases. It is fortunate for the Souafas that their soils are not saline, 
or but very slightly so, as it is hard to see how they could possibly 
reclaim strongly saline lands in view of the conditions of water sup- 
ply in their country. 


THE DATE GARDENS. 
Let us now have a closer look at the gardens. (PI. III, figs. 1 
and 2.) The craterlike basins which they occupy are generally cir- 


cular or nearly so, and from 35 to 50 feet deep. The bottom is 
entirely given up to the palms. Descending to the floor of the basin, 


aAccording to Jus this hard quartz sand extends to a depth of 3.5 to 4.5 feet; 
next there are from 7 to 8.5 feet of a “ reddish gypseous sand;” and then 3 to 
5.5 feet of cither “a fine quartz sand” or “a yellow gypseous sand.” 

+The composition of this rock, as given by Jus (ibid., p. 69), is: 


Per cent. 
Quartz sand___ <a te oo ee eo a 37. 00 
Clay — os a i wa a wo el 
Gypsum ae t : ie ee a tn we nd 41. 40 
Carbonate of lime_ Pee mate ty. i an ee oe ae en a os ee 3. 20 
Carbonate of magnesia pi cn ll a lc Nc 1. 50 
UC i a a car ee rk wl meee drs sc 


THE DATE GARDENS. 19 


or “ ghitan,” as the natives term it, we find it to be a practically level 
expanse of clean, bare sand, checkered with the bright sunlight and 
the singularly black shadows that are cast by the trunks and leaves. 
(Pl. V, fig. 1.) The palms stand farther apart than in the gardens 
owned by natives in the Djérid and the Oued Rirh, but are not 
planted in rows and at equal intervals, as in the French plantations 
in the latter region. While native gardens elsewhere in the Sahara 
are a perfect jungle of various fruit trees, besides garden vegetables, 
barley, and alfalfa underneath the palms, in the Oued Souf one sees 
only scattered pomegranate and fig trees, and the groves have an un- 
familiarly open and bare look. While in other oases the soil is 
often rich and black and is almost always moist, here it is quite dry 
on the surface. One misses, too, the irrigation and drainage ditches 
by which the gardens of the Djérid and the Oued Rirh are cut up into 
small plats. 

Another feature of the Souf date orchards that immediately at- 
tracts attention is the enormous thickness of the trunks of the trees. 
They sometimes attain 3 feet in diameter. (PI. V, fig. 2.) This is 
probably due to the trees being comparatively far apart, thus receiv- 
ing plenty of light and air from every side, and is, perhaps, also to 
some extent a reaction to the buffeting of the sand-laden winds. At 
any rate, it is a useful character, giving the trees power to withstand 
the winds that prevail here to a greater extent than in the other 
oases of the northern Sahara. The relatively small height of the 
palms, which rarely exceed 30 feet in the Oued Souf, gives them a 
further advantage in this respect. Frequently, when the base of the 
trunk has become weakened and there is danger of the tree blowing 
down, the natives make a “ dokana,” or low, circular mound of soil, 
plastered on the outside, to strengthen it. (See Pl. V, fig. 1.) 

The palms are almost invariably strong and healthy looking. The 
foliage is extraordinarily well developed, and the leaves commonly 
measure 15 to 20 feet long. The yields of fruit, as stated by the 
natives, are very heavy. 

So unusual are the conditions under which date palms are grown 
in the Souf country that further details as to the methods used by 
the natives can not fail to be interesting. 


PLANTING. 


As the date palm is a tree that requires a great deal of water, it 
can evidently be grown in a dry country without surface irrigation 
only in places where its roots can quickly make their way to ground 
water. This is exactly the condition obtaining in the Oued Souf, 
where the palms are artificially watered only during the first summer 
after the offshoots are planted, and are then left to shift for theim- 
selves, so far as water supply is concerned. 


20 AGRICULTURE IN THE SAHARA DESERT. 


As we have seen, the bottoms of the basins where the palms are 
grown are not only far below the summits of the surrounding sand 
hills, the height of which is increased by the sand removed in exca- 
vating the gardens, but are even considerably lower than the mean 
surface of the country. It is said that in starting a new garden the 
practice is first to sink a well in the bottom of the basin in order 
to find out the depth at which water stands. The floor of the basin - 
is then scooped out until it is so near ground water that when a hole 
i} to 4 feet deep is made to receive the young palm its roots will 
have to descend only about 1 or 14 feet to reach standing water. It 
is said that to attain the desired depth it is generally necessary to 
remove 10 to 20 feet of sand. 

The date palm is always artificially planted in the Oued Souf, 
never springing up spontaneously from seed, as in other oases. It 
is never planted elsewhere than on the floor of the basins among the 
dunes, or at most a very few feet above the bottom. These basins 
are probably in all cases natural depressions, but are artificially 
deepened to facilitate the roots reaching ground water. New gar- 
dens are frequently started in unoccupied basins, and old ones belong- 
ing to enterprising owners are being constantly extended by cutting 
down the slopes of the bordering sand hills and planting a few palms 
every year or so. (See Pl. III, fig. 2.) The larger and better-situ- 
ated basins are now all occupied by gardens, and for the newest 
plantations it is often necessary to use small, shallow depressions, 
where there is frequently room for but half a dozen trees. Some- 
times the slope is not cut down quite to the level of the older part of 
the garden, the new palms being set out slightly above the level, on a 
terrace made in the side of the sand hills. (Pl. IV, fig. 1.) When 
planted on the slope or near the foot of it, sections of palm log or a 
number of palm leaves are placed on the uphill edge of the hole to 
check the drifting of sand into it. 

Owing to the scarcity of offshoots in the Souf region, the work of 
extending the gardens does not proceed as rapidly as the energetic 
population could wish. The French attribute the nonproduction of 
offshoots in the Souf to the fact that the palms are so valuable there 
that it does not pay to let the offshoots develop, absorbing a part 
of the energy that would otherwise go to fruit production. They 
believe that the Souafas find it actually cheaper to send to the Oued 
tirh for suckers, paying 40 to 60 cents apiece for them in addition 
to the cost of transportation, than to let them grow on their own trees. 
Sconomic considerations aside, however, it is probable that the date 
palm does not sucker as freely in the Oued Souf as in other oases, 
because of the dry condition of the surface soil, never wet by irriga- 
tion, and because the blowing sand tends to bury the young offshoots 
and to lacerate their tender buds. The natives, when questioned 


THE DATE GARDENS. 21 


about the comparative rarity of offshoots at the base of their palms, 
reply simply that it is due to the absence of irrigation, without going 
into details. Whatever may be the cause of the deficiency, there is a 
great demand for offshoots, and to supply this demand caravans are 
sent to procure suckers, especially of the Deglet Noor variety, to Tou- 
gourt, or even as far as Ouargla, 135 miles away. In those oases they 
are produced more freely, the palms being irrigated. 

Offshoots for planting are generally taken from the mother palm 
about the end of February. The natives say that they could be 
planted even earlier, but in that case the parent tree is likely to suffer 
from the access of cold air to the wound made in cutting off the sucker. 
Tn case the offshoots are removed in midwinter, their bases are slightly 
charred before planting, and this is thought to protect them from the 
cold. 

The hole made to receive the young palm is sometimes as much as 
6 feet in diameter, but probably in most cases less. Its depth, as we 
have seen, depends largely upon the distance to ground water, being 
generally 14 to 3 feet in the bottom of the gardens near the town of El 
Oued. A young palm was seen near El Oued that had been set out 
near a well about 6 feet above the bottom of a garden, in a hole 33 
feet deep. (See Pl. IV, fig. 1.) 

The palms are not set out in straight lines. They stand much far- 
ther apart than in gardens belonging to natives in other oases, 20 
feet being the average distance. This wide planting is probably 
necessitated by the poverty of the soil, which is practically a pure 
sand, while the almost entire absence of subsidiary cultures makes the 
shade afforded by close planting less valuable than in other oases. 

It is estimated that the planting and care of a young Deglet Noor 
palm up to the time it begins to yield costs $25 in the Oued Souf, as 
against $5 to $10 in the Djérid oases of Tunis. 


CARE OF PALMS. 


During the first summer after it is planted the palm may receive a 
few irrigations by hand with water from the well that is situated on 
the hillside in nearly every garden, although it is said that frequently 
no irrigation whatever is given. While still very small, before the 
leaves have grown out enough to project far above the mouth of the 
hole in which it is planted, the tree is protected from drought and 
from the cold of winter by covering the hole with palm leaves, dead 
pumpkin vines, ete. . 

When the palms are manured the sand that has piled up into a 
low mound around the base of the tree is removed and the soil beneath 
is thoroughly worked. This is apparently the only cultivation the 
trees receive. 


22 AGRICULTURE IN THE SAHARA DESERT. 
FIGHTING THE SAND. 


While elsewhere in the Sahara irrigation entails the heaviest labor 
connected with palm culture, in the Oued Souf it is the struggle that 
must be waged with the constantly encroaching sand that demands 
the tireless efforts of the gardeners. Every strong wind carries great 
volumes of sand. The shght fences of palm leaves and the low walls 
of gypseous rock that are constructed along the crests of the bordering 
dunes are only a partial protection against this invasion. Once over 
these weak barriers, the sand rolls down the steep slopes almost like 
water. The danger is always present, but is most pressing when the 
dates are ripening. Then bunches of fruit that hang close to the 
ground can be half buried by a few hours of high wind, and only the 
most strenuous efforts can save them. Ifa second storm occurs before 
the bulk of the sand is removed, the crop is hopelessly lost. 

The work of cleaning out the basins is very laborious, being done, 
like the original excavation, almost entirely by hand. Travelers in 
this region have compared this work to the activities of ants, rather 
than of men. Laborers shovel the sand into baskets and carry them 
in a ceaseless procession to the top of the slope, their feet sinking 
deep into the flowing sand at every step. After a heavy sand storm 
the work must be continued from dawn to dark. In summer, during 
the blazing midday hours, the heat is too great for such heavy labor, 
and the removal of the sand goes on at night and in the early morn- 
ing hours. At times a large part of the population of the region is 
engaged in this heavy task. It is paid for at the rate of 1 cent for 
every 5 baskets of sand, and the laborer has, in addition, the privilege 
of eating as many dates as he desires in the garden in which he is 
working. Only rich proprietors use the sturdy little gray donkeys 
of the Souf for transporting the sand from their gardens. 


MANURING. 


The soil of the Oued Souf is practically nothing but pure sand, 
containing even in the older palm gardens very little organic matter. 
Manuring is consequently essential not only to the production of good 
yields but even to the well-being of the palm itself.” 

It is not uncommon in the Souf country to see palms that have 
thick trunks up to a certain point, above which they contract more or 
less abruptly to a much smaller diameter. In many cases, at a still 
greater height, the trunk again becomes thicker. This state of things 
is explained by the natives as due to a partial starvation of the tree at 

«For that matter, manuring is generally practiced by good farmers in the 
oases of the Oued Rirh and the Djérid, although there the growing of legumi- 
nous food and forage crops (broad beans and alfalfa) helps to restore to the soil 
the nitrogen that is taken up by the palins. 


THE DATE GARDENS. 93 


the time when the trunk began to diminish in size. If manure is 
subsequently supplied to it, the palm is soon able to return to its 
normal rate of growth and the trunk again becomes larger. 

Palms are not manured until they are 10 or 12 years old. At that 
age each tree usually receives 10 sacks (5 camel loads) of manure, 
half of which is applied on one side of the trunk the first year and 
the other half on the other side the following year. Thereafter, in 
order to obtain the highest yields, the trees should be manured every 
twelve or fifteen years, although sometimes thirty years are allowed 
to elapse between two manurings. Older palms receive as much as 
14 sacks of manure (7 at each application). Camels’ dung (see 
Pl. IV, fig. 2) only is used for date palms in the Oued Souf, although 
in the oases of Tunis that of donkeys is preferred, the natives there 
considering camel manure injurious where irrigation is practiced. 
The cost of a sackful of camel manure in the Oued Souf was stated 
by one informant to be 25 to 30 cents, while another placed it at 40 
to 45 cents. In either case it is evidently an expensive article. 

Manure is never used until it is thoroughly rotted, and even then it 
is not allowed to come into direct contact with the base of the tree. 
It is placed in a hole that is dug to a depth of 3 to 6 feet below the 
general level of the floor of the basin and at a distance of 5 or 6 feet 
from the foot of the palm. When several neighboring palms are to 
be manured at the same time, the hole is dug in the center of the space 
among them, and is made so large that none of the palms is more than 
6 feet distant from its edge.t| The hole is then filled with a mixture 
of one part manure and one part of a bright yellow sand called 
“ baker,” which is somewhat more loamy and probably contains more 
gypsum than the surface sand of the region, and is obtained at a 
greater depth. Unmixed manure is never used, even though thor- 
oughly rotted, being considered injurious to the palm roots. The soil 
removed in digging the hole is never put back, being “ dirty,” as the 
natives express it. 

October is considered the best season for applying manure, al- 
though March is also a good time. Unskillful growers sometimes 
manure their palms at other seasons, but this is thought to do more 
harm than good. Sometimes, unless the hole is opened and _ the 
manure removed as soon as the tree shows signs of injury, it is said 
to die from the effects of manuring at the wrong period. 

The effect of manuring upon the yield is large and almost imme- 
diate. It is said that a tree which bears 200 pounds of dates one year 
will often give 400 pounds the season following if meanwhile manure 


@QOne such hole, freshly excavated at the time of the writer’s visit, occupied 
much the greater part of the area among four palms, being 12 feet long and 5 
feet wide. It was divided unequally by a narrow ridge of soil left in place. 
The object of this division could not be learned, 


24 AGRICULTURE IN THE SAHARA DESERT. 


has been applied. No distinction between varieties is made in the 
Oued Souf in manuring, nor, so far as could be ascertained, in regard 
to other cultural practices. 


HARVEST. 


At the time of the writer’s visit (November 22-26, 1904) the date 
harvest had been completed in all the gardens of the Souf country. 
The Deglet Noor harvest is said generally to begin about October 25. 
In the Tunisian oases, on the other hand, the harvest of Deglet Noor 
and Fteemy dates—the two most important varieties—was at its 
height in November, and continued throughout December and even 
the earlier part of January. Of course, in the latter case many of 
the dates were ripe long before they were gathered, and the long 
duration of the harvest was largely due to the relative scarcity of the 
expert labor required, the crop being many times as large as in the 
Souf. Yet it seems certain that in the Oued Souf, dates, especially 
the Deglet Noor variety, ripen earlier than in the Oued Rirh oases 
of Algeria or the Djérid oases of Tunis. This would be expected 
from the fact that the summer is drier and likewise hotter in the 
Oued Souf than in the oases of the Djérid.. Furthermore, the situ- 
ation of the gardens, in hollows bordered by hills of light-colored 
sand that are generally higher than the tallest palms, is favorable to 
an early ripening of the dates, as they must receive a great amount of 
additional heat by reflection from the soil. More perfect natural 
conditions for forcing fruit to early maturity could probably not be 
found in the world. 

This greater heat and dryness of the Souf climate affect the fruit 
in other ways than merely by hastening its ripening. The dates pro- 
duced are reputed to be the best grown in the Sahara. They seem, 
as a matter of fact, to be sweeter and at the same time drier and more 
solid than in the Djérid. This is especially true of the Deglet Noor, 
which is of decidedly firmer texture, containing less water. The 
Souf dates are said to keep better and to be more adapted to export 
than those of the other oases, showing less tendency to blacken and 
become moldy. 

It was a matter of regret to the writer that the harvest was not 
witnessed in the Souf, although it could not be learned that the meth- 
ods followed there differ from those practiced in other oases. So far 


«Rolland (iHydrologie du Sahara Algérien, Paris, 1894, p. 222) describes the 
basins as “a sort of fiery furnace, under the influence of solar radiation. ‘The 
dates here attain perfect maturity. Tere are realized, better than anywhere 
else in the Sahara, the conditions assigned by the Arab proverb to the prosperity 
of the palm and the excellence of its fruits: ‘Its feet in the water and its head 
in the fire of heaven.’ ” 


‘THE DATE GARDENS. 25 


as is known, the pollination of the female flower clusters in the spring 
is also effected in the same way as in the Oued Rirh.* 


YIELDS. 


It was impossible to secure very reliable statements of yields from 
the natives, but from all that could be learned these must be unusu- 
ally large in the Souf country. The clusters of the Deglet Noor are 
said frequently to weigh over 55 pounds each, and to attain some- 
times 90 pounds. Single trees of this variety, which is one of the 
lightest bearing kinds, sometimes yield as much as 330 pounds in 
the Oued Souf. It was estimated in 1883 that the date crop from the 
175,000 palms (of all varieties) in full bearing then existing in the 
Souf region was 7,000,000 pounds. This would mean an average 
yield of 40 pounds per tree, as against an average yield of 28 pounds 
estimated to have been produced in the Oued Rirh the same year.” 
A good palm in full bearing is valued at from $50 to $130, according 
to the variety to which it belongs. 

The practice of planting the trees farther apart than in other oases 
is perhaps one reason for the large yields. By wide planting, not 
only do the roots of each palm have a larger feeding area, but the 
trees do not shade each other so much and more of the fruit can 
develop and ripen. Moreover, the climatic and topographical condi- 
tions, as we have seen, are exceptionally favorable to the ripening of 
dates. 

VARIETIES CHIEFLY GROWN. 


As has already been indicated in this paper, date palms in the 
Oued Souf rarely, if ever, spring up from seed, as they do in other 
oases where the conditions are more favorable to the spontaneous 
development of the palm. They are propagated only by offshoots 
that are taken from the parent tree and planted by the grower. 
Consequently we do not see a multitude of seedlings, generally of 
very inferior quality and of almost endless diversity of characters. 
filling every waste corner and roadside and even crowding out good 
trees in gardens that are not well cared for. Practically every palm 
grown in the Souf belongs to some well-known and well-liked variety. 

The number of varieties found in this region is considerable. 
Most of the gardens contain a mixture of several kinds, although in 
some of the recently created ones the tendency is to plant only one 


“Described in Bul, 53, Bureau of Plant Industry, “The Date Palm,” by W. T. 
Swingle, 1904, pp. 26-29. 

b These estimates are quoted from Rolland (ibid., p. 324). The overwhelming 
importance of the date crop in the Souf region is shown by the fact that the 
same author states the value of the 1883 crop of dates to have been $301,730, 
while that of all other crops combined was only $20,900 in that year. 


26 AGRICULTURE IN THE SAHARA‘ DESERT. 


sort, most often the Deglet Noor. Nearly all the popular varieties 
of the Souf are also common in the Oued Rirh.* 

On the other hand, some of the most characteristic Souf types are 
very rare in the Djérid cases of Tunis, only 70 miles away. Such 
individuals as occur there are mostly grown from offshoots that have 
been brought directly from the Souf. The principal variety common 
to the two regions is the Deglet Noor, which is now abundant in 
Tunis, but is said to have been first introduced into that country from 
Algeria about two hundred and fifty years ago. The Souafas still go 
to the Oued Rirh oases to procure offshoots, and they very likely 
brought thence those with which the first gardens were started among 
the sand hills of the Souf. 

The most important of the numerous varieties of the Souf, in point 
of abundance as well as of quality, are, in about the order named: 
Deglet Noor, Rhars, Tafazween, Massowa, Deglet Beida, and Taker- 
mest. Of these Rhars is the earliest and Deglet Noor the latest to 
ripen. After the Deglet Noor, Tafazween is the best sort that is 
widely grown. It is a large, reddish-bay-colored, translucent date, 
very sweet and rich in flavor.’ A highly esteemed but very rare 
variety 1s the Fezzani, which is said to be superior even to the Deglet 
Noor when dried, and to keep well for two years. Rhars, a variety 
that is celebrated for its heavy yields, is extensively planted. 

In the Oued Souf, as in other oases of Algeria and Tunis, the 
Deglet Noor is the only variety that has any importance as an article 
of export to Europe. It is consequently the most valuable, the more 
so because the natives themselves generally esteem it above all others. 
Deglet Noor dates are carried from the Souf by caravan to Biskra, 
whence they are shipped by railway to the seaports. Souf Deglets 
are said to be about the earhest to reach the Biskra market. Their 
good keeping and shipping qualities have already been discussed as 
probably due to the peculiar climatic conditions, which give them an 
advantage over dates from oases where the palms are lavishly irri- 
gated and the air is moister. On the other hand, they appear to be 
smaller than those in the Djérid and to be inferior in color and gen- 
eral appearance. The latter disadvantage is very likely due in great 
part to the sand-laden winds to which they are exposed. The Deglet 
Noor palm is said to be hardier in the Souf region than elsewhere, 
showing greater resistance to disease and to unfavorable climatic 
conditions. The foliage of the date palm appears to be less subject 
to the attacks of seale insects than in other oases, which is perhaps 
attributable to the extreme dryness of the atmosphere. 


a })xceptions are said to be the Fezzani, Massowa, Ali Rashid, and Guettara 
varieties. 

» Twenty offshoots of this variety were obtained from the Oued Souf for trial 
in the United States through the kindness of the French commandant at El 
Oued, Captain Bussy. 


bo 
“J 


CONCLUSION. 
CONCLUSION. 


The type of agriculture practiced in the Oued Souf is not dry- 
land farming, for it depends upon the ground water, which in the 
gardens is everywhere near the surface of the soil. It affords us, 
however, an excellent object lesson of what can be done under the 
most adverse natural conditions in producing a valuable crop, for 
throughout northern Africa the Oued Souf is renowned for the large 
yields of its date orchards and the high quality of their fruit. 

It may be that nowhere in the United States are the conditions with 
respect to ground water such as to allow of a close imitation of 
agricultural methods used in the Souf country. One lesson is, how- 
ever, to be drawn from them. The sand hills concentrate and reflect 
so much heat that the hollows among them are veritable forcing 
houses, where dates ripen considerably earlier than elsewhere. Have 
we not here a hint of what may be done in the Salton Basin and 
perhaps in other hot, arid regions in the Southwest where large sand 
dunes exist, and where artesian or other sources of water supply 
for irrigation are available? It seems certain that in pockets of 
this character excavated among the dunes the Deglet Noor and other 
valuable varieties of dates could be forced to early maturity. 

Dates ripened in this way a few weeks ahead of the bulk of the 
crop would command a fancy price, especially as the quality of the 
fruit produced under these conditions would in all probability be 
exceptionally fine. Nor are the possibilities limited to the date 
palm. Other fruits, such as figs, pomegranates, and grapes, could 
perhaps thus be put upon the market in advance of those from any 
other locality in the United States. The experiment is certainly 
worth trying. The American fruit grower, awake as he is to every 
new idea, may find something worthy of imitation in the example of 
these sturdy inhabitants of a remote corner of the Sahara. 


DESCRIPTION OF PLATES. 


Puate I. (Frontispiece.) General view of the Oued Souf region from the town of 
El] Oued, showing sand dunes and sunken gardens of date palms. 


PuaTeE II. Fig. 1—High sand dunes east of El Oued. -The group is standing on 
a ridge separated by a ravine from the very high dune in the background. 
Fig. 2.—A typical dwelling house of the Oued Souf, showing its cubical 
form and roof composed of flattened cupolas. Fig. 3.—General view of the 
Oued Souf region, showing sunken date gardens and sand dunes. Fence of 
dead palm leaves along crest of dune in foreground. 


PLATE III. Fig. 1—Near view of sunken palm garden and surrounding dunes. 
Vegetable garden in left foreground, showing small size of checks. Near it 
a young palm, planted ina hole. Fig. 2—Gradual extension of a palm garden 
by cutting down bordering sand hills. Oldest palms in background, youngest 
in foreground. Tig. 3.—Vegetable garden irrigated by well near bottom of 
basin in which date palms are grown. 


PLATE LY. Fig. 1.—Ilole on slope of dune near bottom of basin in which a young 
palm is planted. Fig. 2.—Camel manure ready for application in a date 
garden. 


Puare V. Fig. 1.—* Dokana,” or mound of earth and plaster for strengthening 
the base of a palm. Shows also distance between trees, absence of other 
cultures, and play of light and shadow on floor of basin. Fig. 2.—Rhars 
palm, showing thickness of trunk. 


50 


O 


Bul. 86, Bureau of Plant Industry, U. S. Dept. of Agriculture. PLATE Il. 


Fic. 1.—HIGH SAND DUNES EAST OF EL QUED. 


Fia. 2.—A TyPICAL DWELLING HOUSE OF THE OUED SOUF, SHOWING ITS CUBICAL 
FORM AND ROOF COMPOSED OF FLATTENED CUPOLAS. 


Fig. 3.—GENERAL VIEW OF THE OQUED SOUF REGION, SHOWING SUNKEN DATE 
GARDENS AND SAND DUNES. 


REG Ate. Prireeqererire” 


oh’ . 
Ay 
> -, 
? 
Fy : . 
ht 
=<. . 
4 7 sty 


Bul. 86, Bureau of Plant Industry, U. S. Dept. of Agriculture. PLATE Ill. 


FiG. 1.—NEAR VIEW OF SUNKEN PALM GARDEN AND SURROUNDING DUNES. 


Fig. 2.—GRADUAL EXTENSION OF A PALM GARDEN BY CUTTING DOWN BORDERING 
SAND HILLs. 


FiG. 3.—VEGETABLE GARDEN IRRIGATED BY WELL NEAR BOTTOM OF BASIN IN WHICH 
DATE PALMS ARE GROWN. 


putes, ativap poe 


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- 
. 
oy 
7 
- 
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+ ~ 
** 
« 
6 
e 


~ 


Pe 
- 


- ~ me 


Bul. 86, Bureau of Plant [ndustry, U. S. Dept. of Agriculture. PLATE IV. 


Fig. 1.—HOLE ON SLOPE OF DUNE NEAR BOTTOM OF BASIN IN WHICH A 
YOUNG PALM IS PLANTED. 


Fig. 2.—CAMEL MANURE READY FOR APPLICATION IN A DATE GARDEN. 


ee, — ie ; hs a 
ie: Last z= a 
Pg papanmwapee te 


7 ae 


"W1Wd VW 4O 3SVQ 3HL ONINSHLONSYLS HOS 
YaLSV1d GNV HLYVy 4O GNNOW YO «'VWNVHOG,,—"} “DIF 


“MNNY | 4O SSANMOIH L SNIMOHS ‘'W1Vd SYVHY—'S ‘DI4 


Bul. 86, Bureau of Plant Industry, U. S. Dept. of Agriculture. 


eet ae a 
ee 


PLATE. V. 


U. S. DEPARTMENT OF AGRICULTURE. 
BUREAU OF PLANT INDUSTRY—BULLETIN NO. 87. 


B. T. GALLOWAY, Chief of Bureau. 


DISEASE RESISTANCE OF POTATOES. 


BY 


L. R. JONES, 


BorTANIST OF THE VERMONT AGRICULTURAL EXPERIMENT 
STATION AND COLLABORATOR OF THE BUREAU 
oF PLANT INDUSTRY. 


VEGETABLE PATHOLOGICAL AND PHYSIOLOGICAL 
INVESTIGATIONS. 


IssuED DECEMBER 5, 1905. 


~ WASHINGTON: 
GOVERNMENT PRINTING OFFICE. 
1905. 


BUREAU OF PLANT INDUSTRY. 


B. T. GALLOWAY, 
Pathologist and Physiologist, and Chief of Bureau. 
VEGETABLE PATHOLOGICAL AND PHYSIOLOGICAL INVESTIGATIONS. ‘ 
ALBERT F. Woops, Pathologist and Physiologist in Charge, Acting Chief of Bureau in Absence of Chief. 
BOTANICAL INVESTIGATIONS. 
FREDERICK V. COVILLE, Botanist in Charge. 
FARM MANAGEMENT. 
W. J. SPILLMAN, Agriculturist in Charge. 
POMOLOGICAL INVESTIGATIONS. 
G. B. BRACKETT, Pomologist in Charge. 
SEED AND PLANT INTRODUCTION AND DISTRIBUTION. 
A. J. Prerers, Botanist in Charge. 
ARLINGTON EXPERIMENTAL FARM. 
L. C. CorBeEtTT, Horticulturist in Charge. 


INVESTIGATIONS IN THE AGRICUL Se cere OF TROPICAL AND SUBTROPICAL 
Ss. : 


O. F. Cook, Bionomist in Charge. 
DRUG AND POISONOUS PLANT INVESTIGATIONS, AND TEA CULTURE INVESTIGATIONS. 
RODNEY H. TRUE, Physiologist in Charge. 
DRY LAND AGRICULTURE AND WESTERN AGRICULTURAL EXTENSION. 
CARL S, SCOFIELD, Agriculturist in Charge. 
EXPERIMENTAL GARDENS AND GROUNDS. 
E. M. BYRNES, Superintendent. 


SEED LABORATORY. 
EDGAR Brown, Botanist in Charge. 


J. E. ROCKWELL, Editor. 
JAMES E. JONES, Chief Clerk. 


VEGETABLE PATHOLOGICAL AND PHYSIOLOGICAL INVESTIGATIONS. 
SCIENTIFIC STAFF. 
ALBERT F. Woops, Pathologist and Physiologist in Charge. 


ERWIN F. SMITH, Pathologist in Charge of Laboratory of Plant Pathology. 

HERBERT J. WEBBER, Physiologist in Charge of Laboratory of Plant Breeding. 

WALTER T. SWINGLE, Physiologist in Charge of Laboratory of Plant Life History. 

NEWTON B. PIERCE, Pathologist in Charge of Pacific Coast Laboratory. 

M. B. WAITE, Pathologist in Charge of Investigations of Diseases of Orchard Fruits. 

MARK ALFRED CARLETON, Cerealist in Charge of Cereal Laboratory. 

HERMANN VON SCHRENK, in Charge of Mississippi Valley Laboratory. 

P. H. RourFs, Pathologist in Charge of Subtropical Laboratory. 

C. O. TOWNSEND, Pathologist in Charge of Sugar Beet Investigations. 

T. H. KEARNEY, A. D. SHAMEL, Physiologists, Plant. Breeding. 

P. H. Dorsett,a CORNELIUS L. SHEAR, WILLIAM A. ORTON, W.M. Scott, ERNsT A. BESSEY, E. M. FREE- 
MAN, Pathologists. 

E. C, CuHiLcott, Expert in Cultivating Methods, Cereal Laboratory. 

C. R. BALL, Assistant Agrostologist, Cereal Laboratory. 

JOSEPH S. CHAMBERLAIN,» J. ARTHUR LE CLERC,¢ Physiological Chemists. 

FLORA W. PATTERSON, Mycologist. 

CHARLES P. HARTLEY, KARL F. KELLERMAN, JESSE B. NORTON, CHARLES J. BRAND, T. RALPH ROB- 
INSON, Assistants in Physiology. 

DEANE B. SWINGLE, GEORGE G. HEDGCOCK, Assistants in Pathology. 

PERLEY SPAULDING, P.J.O'GARA, FLORENCE HEDGES, HENRY A. MILLER, ERNEST B. BROWN, LESLIE 
A. Fitz, LEONARD L. HARTER, JOHN O. MERWIN, A. H. LEIDIGH, H. F. BLANCHARD, Scientific 
Assistants. 

W. W. Copsey, Tobacco Expert. 

JOHN VAN LEENHOFF, Jr., T. D. BECKWITH, Experts. 


a Detailed to Seed and Plant Introduction and Distribution. 
» Detailed to Bureau of Chemistry. 
¢ Detailed from Bureau of Chemistry. 


LETTER OF TRANSMITTAL. 


U. S. DEPARTMENT OF AGRICULTURE, 
Bureau oF Puiant INDUSTRY, 
OFFICE OF THE CHIEF, 
Washington, D. C., October 7, 1905. 

Str: I have the honor to transmit herewith the manuscript of a 
technical paper entitled ‘‘ Disease Resistance of Potatoes,” which 
embodies a report upon investigations conducted in cooperation with 
the Vermont Agricultural Experiment Station. 

This paper is a valuable contribution to our knowledge of disease 
resistance in European and American varieties of potatoes, and I 
respectfully recommend that it be issued as Bulletin No. 87 of the 
series of this Bureau. 

Respectfully, B. T. GaLitoway, 
Chief of Bureau. 
Hon. JAMES WILSON, 
Secretary of Agriculture. 


~ 


J 


PREFACE. 


The potato is one of the most important food crops of the United 
States. It is, moreover, one which is subject to a number of serious 
plant diseases. Some of these, notably the late-blight, can be con- 
trolled by spraying. Yet this remedy is not applied by all farmers, 
and the annual loss amounts to many millions of dollars. Other dis- 
eases, like dry-rot and the bacterial blight, are not controlled by spray- 
ing and require a different line of treatment. The subject of disease 
resistance in plants has received increased attention of late, and it is 
likely that the introduction of disease-resistant varieties of potatoes, 
by supplementing spraying and special cultural practices, will be 
of great practical value in lessening the waste caused by disease. 
Although but little has been done in the United States toward secur- 
ing varieties resistant to disease, the attention of potato specialists in 
other countries has already been directed toward this aim. 

As a preliminary step in our work, Dr. L. R. Jones, botanist of the 
Vermont Agricultural Experiment Station, was commissioned to 
inquire into the occurrence of potato diseases abroad and the methods 
employed for their control, particularly with reference to the produc- 
tion of disease-resistant varieties. Doctor Jones spent six months— 
from April to September, 1904—in this work. 

In the course of Doctor Jones’s European itinerary, information of 
more or less value was secured at the following places, successively, 
and this was supplemented by a considerable correspondence covering 
a somewhat wider area: Marseille, Naples, Florence, Munich, Halle, 
Berlin and vicinity, Dresden and vicinity, Bonn and vicinity, Wagen- 
ingen (Holland), Amsterdam and vicinity, Groningen, Delft, Gembloux 
(Belgium), Paris and vicinity, London and vicinity, Reading, Cam- 
bridge, and Edinburgh and vicinity. The thanks of the Department 
are accorded to the various officials and other botanists in the coun- 
tries visited, whose uniform courtesies made possible the success of 
the mission. 

As shown in this report, a considerable number of varieties are 
reputed to be disease resistant. The best of these were selected by 
Doctor Jones, and limited quantities of seed tubers were imported by 
the Office of Seed and Plant Introduction and Distribution of the 


- 


o 


6 PREFACE. 


Bureau of Plant Industry. These are now being tested in trials con- 
ducted in cooperation by the Bureau of Plant Industry and the State 
experiment stations in Vermont, Florida, Colorado, and Oregon, and 
by the Bureau at the Arlington Experimental Farm, near Washing- 
ton. In order fully to acclimatize the foreign varieties, these trials 
will be continued for three years before a final report is made on their 
adaptation to American conditions. This test, together with the 
review of our present knowledge contained in this bulletin, will estab- 
lish a proper foundation for future efforts in breeding better and more 
disease-resistant varieties. The field for such work is very large. 
Up to the present time most of the breeding has been for resistance 
to the late-blight (Phytophthora infestans), and this will continue to be 
the principal problem in the northern tier of States; but there is also 
much promise of success in securing new varieties to resist scab, dry- 
rot, bacterial blight, and other troubles, which in the Southern and 
Western States are more injurious than late-blight. It is hoped that 
potato specialists will give increasingly careful attention to this fea- 
ture in their breeding and testing of varieties, for it is only by sucha 
general interest and effort that the desired information can quickly be 
secured. 


ALBERT F. Woops, 
Pathologist and Physiologist. 
OFFICE OF VEGETABLE PATHOLOGICAL AND 
PHYSIOLOGICAL INVESTIGATIONS, 
Washington, D. C., October 5, 1905. 


CONE ENTS: 


Page. 

REDNESS ee pb nee Pak peo Pein ew olde Palees aac abe a 9 
Per meN COMIN T OCs * serree et tie Se ee ee LoS a Se asiele oc s's Kes sasec 9 
Observations on potato diseases and disease resistance in Europe....--------- 12 
DEPETEMUNSINERSE CLIREMBCH 2-922 5) ee eo 2 ae Sa oot. 12 
ioe DT OWARSUOU 22 Wraeee atte oo Aeon | ee eee eee See Boot 12 
OE CLOWN -OU bas so Ss hake oe we Hse aa eat ay Sats 13 
ee Se Se tae ee on IE a ee Shek ab Seon sane 13 
MIRE TPEISCIDPISES a2 1 os eee 5 a ote ee NE seen are 14 

LP DERI PG) (STC) Oa oa a teas Pe a ee ee PR ee nea a ey I a Oe, aE 14 
Meerioict remisithec Lo Stal 2) 2 2 he ee ete as. seed 2 15 

Mine msGaAD-like: (ISCASCS =" c0 Sin 2 os yooh ee Buk eee eae cae Be ee 16 

CUCINA LCTE CO ISCHSCR OM yeh cele Sn oases iene Sk a ihn ea ae 17 

Re ER res Se oS Nhat tren a ie ia ee anya cis owas fae DE SE Sa Soe 17 
PieTEsbeMUCISCASCS- eons ere eee ak aoe ete o Sib eee a ade 18 
Late-blight and rot due to Phytophthora infestans.........--------------- 19 
Resistance as shown toward late-blight and rot.........---..--------------- 20 
Vi CSTE T ELSES 1 ETOYS ee ER pO pean 5 a ee ae 20 
POE ine OF Cisease FesIstANCG. 2.22522 25-222 o en en wate conse neon 23 
Disease resistance and vegetative vigor. 2.2. 2.-i2.-2-2--52-2-2+---5--22- 23 
Yue relation of hybridity to disease resistance... ..-..--.-.--....-..5-- 25 

UU SST eg 1 a ee 25 

Are early or late varieties the more resistant?_......-..----------------- 26 
Relation of source of seed and cultural methods to disease resistance - -- - - 26 
Composition and character of tubers as related to rot resistance... -..----- 27 
Character of stem and foliage as related to disease resistance.....-------- 28 
WiseAsc-resishanty yarieties Of Murope <2. 2.2554. ere 28 
nme iGAl Re oe Sele a xe Ae Pe 2 es... Bese Soe ces dae eee eee 29 
MRSC ATI ae cLe) Clete Gy WLAN OL Ss ot er tae yee SEE oo SORE See eee Bae oe Sk 30 
Teen ese OMIM sen oS 2. oe has Oe oe ee Ree eee sso esas s 30 
Drsesse-resisiant varieties of America = 22-2 2.22 _ <2 eset new ost bones 31 
Investigations at the experiment stations-_--.........-.----------/---.--< 31 
WMankat ine. Vermont station. 24.025 55-225 S522 ec ete se cscs ae 32 
iniormation secured by a circular of inquiry-.-:-_-..--.==-----------.-- 33 
Srna MOISCAD-2 sac ek aa Se 2 one waa eA ot oes ae ee eee 35 
SUID TS OTA tee, Sa ee ERC Ao 25 Se ee eee eee 37 


B. P. L.—179. _ VY. P. P. 1.140. 


DISEASE RESISTANCE OF POTATOES. 


INTRODUCTION. 


Potatoes are liable to several diseases, some of greater economic 
importance in one part of the country, some in another.“ The causes 
of most of these have been determined, and remedies of more or less 
practical value, chiefly’ spraying and seed disinfection, have been 
found. The success of the agents of the Department of Agriculture 
in securing disease-resistant varieties of various other plants has given 
an added interest to the question as to what may be expected in the 
way of securing disease-resistant varieties of potatoes. Inasmuch as 
this matter has engaged the attention of the potato specialists of 
Europe longer and more widely than has been the case in America, 
the attempt has been made to glean from their experience whatever 
may prove of assistance in furthering the work in this country. 

In presenting the results it has seemed best for the sake of clearness 
to discuss briefly (1) certain general matters relating to potato culture 
in Europe; (2) the potato diseases which occur there, with comments 
on the resistance shown by particular varieties to each disease, and 
then (3) to summarize the information obtainable in America as to 
disease resistance of potatoes. 


POTATO CULTURE IN EUROPE. 


Potato culture was introduced from America into Europe more than 
three centuries ago. It was slower in its popularization there than in 
this country, but to-day the potato crop is relatively more important, 
both for food and for factory uses, in Europe than in America. This 
is partly attributable to the greater success of maize as a starch- 
producing plant in America and partly to the difference in economic 


«Anyone not familiar with potato diseases should consult Farmers Bulletin No. 
91, Potato Diseases and Their Treatment, by B. T. Galloway, which will be sent free 
of charge upon application to a Senator, Representative, or Delegate in Congress, or 
to the Secretary of Agriculture, Washington, D. C, 


8838—No. 87—05——2 9 


10 DISEASE RESISTANCE OF POTATOES. 


conditions—land and labor values and food habits—between the two con- 
tinents. In the British Islands the annual acreage in the last decade @ 
was more than one and a quarter million; in France three times and in 
Germany six times this amount. The crop in Great Britain is chiefly 
used for human food; in France about 40 per cent is used in starch and 
alcohol manufacture, and in Germany like conditions exist, with the 
use of the potato for alcohol distillation promising to increase largely. 

It is the German national policy so to improve processes of culture 
and manufacture that potato alcohol may rival, if not displace, petroleum 
for lighting and fuel purposes. Various organizations are aiding in 
this movement, including the Deutschen-Kartoffel-Kultur-Station and 
the Institut fiir Gihrungsgewerbe und Stirkefabrikation. These 
institutions, maintained partly by private endowment and membership 
and partly by government aid, have been in operation about fifteen 
years. Under the directorship of Profs. C. yon Eckenbrecher and 
W. Delbriick, respectively, they give attention to all matters pertain- 
ing to potato culture, starch manufacture, and distillation, including 
the breeding, selecting, and trial of new varieties. They work in close 
cooperation with the various German experiment stations and private 
potato specialists. No similar institutions were met with elsewhere in 
Europe, although the potato growers of England have organized dur- 
ing the last year an association of much promise, the National Potato 
Society. 

In any comparison of European with American varieties the differ- 
ence in popular taste must be kept in mind. The English market is 
like the American in giving preference to varieties with white flesh, 
rich in starch, and in making little use of potatoes except for human 
food. On the continent of Europe, however, only the varieties with 
yellow flesh are rated as of first quality for table use. These are rela- 
tively poor in starch and richer in protein than the white-fleshed varie- 
ties, and when cooked they are not sufticiently dry and mealy to suit 
the American taste. On the Continent the white-fleshed, starch-rich 
potatoes are demanded by the starch factories and distilleries, and as a 
result continental potato breeders have aimed to develop white-fleshed 
varieties of high starch content and large productiveness, regardless 
of table qualities, and anyone importing their varieties for use in 
America must consider these facts. It is from England, therefore, 
that we may expect the more promising varieties. 

It is worthy of note in this connection that the difference in char- 
acter of the potato in popular favor in different countries is closely 
related to differences in methods of cooking. The varieties with yel- 
low flesh are inclined to sogginess when baked or boiled, but are admir- 
ably suited for frying. The starch-rich white potato, which is of 


“Sutton, A. W., Potatoes, Jour. Roy. Hort. Soc., XIX (1896), pp. 887-430. 


POTATO CULTURE IN EUROPE. if 


highest quality when baked, may go to pieces if boiled and will not 
hold together sufficiently for frying. In the latter case steam cooking 
gives better results than boiling in water.? 

The methods of culture of potatoes vary somewhat in different Euro- 
pean countries, as they do in America. In general, closer planting is 
practiced than in this country, which is partly explained by the greater 
number of unskilled laborers in Europe. A matter of more pertinence 
to the present discussion is that much more attention is given, espe- 
cialiy in England, to the source and handling of seed potatoes than is 
generally the case in America. The best informed English growers 
considered that this has much to do with the question of disease resist- 
ance, as will be explained later. Indeed, it has come to be accepted 
by them that, for the best results, seed potatoes should be imported 
from some more northern region, at least as often as every third or 
fourth year, and this is done by many large growers every year. 
English growers use Scotch seed very largely. On the grounds of 
Sutton & Sons, Reading, England, a comparative trial was conducted 
during the summer of 1904 with a number of varieties, using seed 
from Scotland, middle England (Lincolnshire), and southern England, 
respectively. When these plats were visited in August, the vigor of 
the plants was in all cases greatest from the Scotch-grown seed and 
least from that grown in the south of England. In certain varieties 
the development of the tops was as two to one. At the Cambridge Uni- 
versity farms, located in the famous potato region of Lincolnshire 
fens, it was said that Scotch-grown seed did better than home-grown 
there and in the Lincolnshire district generally. 

A Jarge potato-growing company of the Canary Islands uses Ger- 
man in preference to English seed, because the growth from the latter 
is too rank and the tubers are too large. Potato growers on the island 
of Jersey in recent years have sent to England and Scotland for their 
seed. Italy and the Mediterranean islands look to France and Ger- 
many largely. 

While some American potato growers recognize the importance of 
the source of seed and attend to this point consistently, there is gener- 
ally indifference, disagreement, or lack of information on this subject, 
as Fraser’ has recently shown. It is a matter deserving of further 
careful inquiry and experiment. 


«The writer is indebted to Professor Petermann for calling his attention to these 
matters. For a further discussion of this subject reference may be made to Peter- 
mann, A., Etudes sur Ja pomme de terre, Bul. de l’ Inst. Chim. et Bact., No. 70, Gem- 
bloux (Belgium), 1901; and Coudon and Bussard, Annales de la science agronomique, 
I, Paris, 1897. To the latter, credit may be given for showing that the behavior of 
the potato in cooking is primarily dependent not so much on the total percentage 
of starch contained as on the relative percentages of starch and protein. The proper 
proportion of the protein is necessary to hold the starchy mass together when cooked. 

bFraser, S., The Potato, New York, 1905, pp. 51-52. 


12 DISEASE RESISTANCE OF POTATOES. 


The relation of the maturity of seed to its value has also received 
considerable attention in Great Britain. Some of the best practical 
potato growers prefer to have potatoes intended for seed dug before 
they are fully mature, i. e., while the tops are still green and the outer 
skin of the tuber rubs up easily. These tubers are then allowed to lie 
on the ground a short time to ‘* harden” in the light, i. e., to dry and 
be slightly sunburned, before they are put into storage. 

The practice of ** boxing” is common and very highly commended. 
This consists of setting the tubers with ‘‘seed end” up in shallow 
boxes or trays in a dry, light room for some weeks before planting, so 
as to have strong, short sprouts started upon them. In this way 
quicker germination, stronger young plants, and a perfect stand are 
secured. ; 

The general preference in Scotland and in England is for planting 
small tubers, one and one-half to three ounces, planted whole. If 
larger tubers are used they are cut, as in this country. 


OBSERVATIONS ON POTATO DISEASES AND DISEASE RESISTANCE 
IN EUROPE. 


Insects cause the European potato grower little annoyance or loss 
as compared with American conditions. Nowhere in Europe, so far 
as learned, is any use whatever made of insecticides upon potatoes, 
leaf-eating insects being practically unknown. On the other hand, 
diseases caused by fungi or bacteria or of nonparasitic origin are even 
more common and destructive than in America. In the following 
paragraphs is a discussion of the nature and relative seriousness of 
such diseases as came under the writer’s personal observation, coupled 
in each case with the evidence secured as to disease resistance. The 
less important and nonparasitic maladies are discussed first, reserving 
until later the consideration of the late-blight and rot. 


CERTAIN MINOR DISEASES. 
INTERNAL BROWN SPOT. 


The peculiar disease of the potato called internal brown spot is as well 
known to plant pathologists and practical potato specialists in Europe 
asin America. The writer learned nothing more about it on the Con- 
tinent than is set forth by Frank,’ who reached only negative conclu- 
sions. It is considered not to be a parasitic disease, and no remedy is 
known and no suggestions are made except the doubtful one of avoid- 
ing the use of diseased tubers for seed. In England and Scotland 
several potato specialists’ of wide experience gave evidence of like 

4¥rank, A. B., Kampfbuch gegen die Schidlinge unserer Feldfriichte, ‘* Bunt- 
werden oder Eisenfleckigkeit,’’ p. 211. 

»Prof. J. H. Middleton, Messrs. Sutton & Sons, and Thomas A. Scarlett. 


CERTAIN MINOR DISEASES. 13 


purport. The trouble is frequently observed, and is most commonly 
termed ‘‘sprain.” It is not propagated in seed or soil and is non- 
parasitic. It is considered to be the direct result of malnutrition 
associated with unfavorable soil conditions, resulting either from too 
dry conditions or from the lack of potash or lime. It is frequent in 
light, dry soils during dry seasons, and is never seen on heavy, strong, 
moist soils. The remedy, in the judgment of the specialists cited, lies 
wholly in attention to cultural conditions and the choice of varieties. 

Some varieties are more liable than others to internal brown spot 
and should not be used on soil that favors the disease; e. g., Mr. Sear- 
lett stated that the British Queen variety is especially predisposed to 
‘**sprain.” The primary remedy, however, lies in selection and treat- 
ment of the soil—i. e., in avoiding dry soil—and in so cultivating as to 
conserve moisture, while using lime and potash liberally. 


FILOSITE, OR GROWING-OUT. 


The names “‘filosité” and ‘‘growing-out” are applied in France 4 
and England, respectively, to various forms of secondary outgrowths 
from tubers. Examples that were shown to the writer were in some 
cases merely “‘prongy” tubers, while in others stolons starting from 
the eyes produced secondary small tubers. Delacroix describes con- 
ditions where the seed tubers send out an abundance of weak shoots, 
both above and below ground, but these soon die without yielding any 
crop. The latter type is therefore allied to the curl disease of the 
potato.’ The less serious forms of growing-out are attributed to cli- 
matic and soil conditions, especially to a long, wet autumn following a 
drysummer. The more serious types of the disease, especially as occur- 
ring in France, are attributed to varietal weakness or ‘‘ running out.” 
In both France and England the use of seed from any plants showing this 
tendency was condemned as tending to give a crop of generally reduced 
vitality. 


LEAF-SPOT. 


Since there has been a difference of opinion among pathologists as 
to the occurrence and destructiveness of the fungus Alternaria solani 
in Europe, the writer was led to keep an especially close watch for it. 
Prof. P. Sorauer kindly sent specimens from Europe some years ago 
corresponding fully with the leaf-spot disease caused by this fungus, 


“See Delacroix, G., Sur la filosité des pommes de terre, Jour. de l’ Agriculture, 
December, 1903. : 

> Frisolée (French), or Kriiuselkrankheit (German), is a common but not very 
serious potato malady of Europe, of which the cause is not clear, but apparently it 
is associated with varietal weakness and malnutrition. It is characterized by dwarf- 
ish plants bearing an excessive number of rather undersized leaves, which are down- 
curled and brittle. The tops have thus a dense or bushy appearance. They may be 
quite as green as normal plants. 


14 DISEASE RESISTANCE OF POTATOES. 


and termed early-blight in America. It is known, therefore, that it 
occurs in Europe, but it can not be as common and destructive as it is 
here, for no trace of it could be found in any of the fields visited. 
Much was seen, however, of a leaf-spot disease which bears a super- 
ficial resemblance to it. This was shown us first by Dr. Otto Appel in 
the experimental fields of the board of health at Dahlem, near Berlin. 
Later it was seen elsewhere in Germany and in England. Apparently 
there has been a confusion of this with our American early-blight,@ 
but both Doctor Appel and Professor Sorauer stated that there was no 
fungus present in this leaf-spot disease, and the writer’s examinations 
confirmed this. The cause of this trouble has not as yet been clearly 
established, but since Appel and Sorauer both have it under observa- 
tion, further advice may be expected upon it in the near future. 
Opportunity was presented to compare the relative development of 
this trouble on different varieties in the trial grounds of the University 
Farm, Cambridge, England, in the latter part of the summer. There 
was a considerable difference in the amount of spotting of the various 
varieties, but none was altogether free from it. 


SCABBINESS OF TUBERS. 
POTATO SCAB. 


Scab-like diseases, i. e., those characterized by surface erosions of 
the tubers, are frequent in Europe and apparently more varied in 
nature than is recognized to be the case in America. Yet nowhere 
does injury to the crop from such diseases approach that which is 
common in this country. This is in some ways surprising and unex- 
plained. A disease closely resembling in appearance the common 
American type of scab was frequently seen, but growers everywhere 
in Germany, France, and Great Britain testified that it is not common 
or destructive enough to be of much practical importance. Scientific 
men are not agreed either as to the cause of the disease or as to the pos- 
sibility of benefit from seed disinfection. But few recommended this 
treatment, and no practical potato grower was met who practiced it. 
The most puzzling thing, in the light of American experience, is that 
potatoes are grown year after year continuously or in the shortest of 
rotations on the same soil without increase of scabbiness. In each of 
these countries potatoes are often grown on the same soil for ten or 
twenty or even forty successive years with practically no trouble 


“See Frank, Kampfbuch, p. 220. Apparently the disease mentioned above is the 
same as Frank’s ‘‘ Pockenfleckigkeit.’’ Frank was altogether mistaken in consider- 
ing it the same as the American ‘‘early-blight,’’ as Sorauer subsequently showed. 
See also Bahna, J. J., Blattbraune der Kart.; Zeitsch. f. Land. u. Fortwirtsch., 2: 113 
(March, 1904). Prof. C. von Tiibeuf comments upon this article and reviews the 
literature in the June, 1904, number of the same journal. 


SCABBINESS OF TUBERS. 15 


from scab.“ Since this is often on rich and heavy land, such as would 
develop scab under like treatment in America, the European experi- 
ence is difficult to understand. Does not our type of disease exist 
there, or is it less virulent because of some difference in the climatic 
and soil conditions? The writer is not prepared to answer with full 
confidence, but his judgment favors the second suggestion. 

Europeans, both pathologists and practical men, recognize more 
than one form of disease under the name of scab (the German 
‘* schorf,” Dutch ‘* schurft,” French ‘‘la gale”). Frank’s subdivisions? 
of ‘‘schorf” into four kinds—fiach, tief, buckel, and buckel-tief 
schorf—were familiar to the continental pathologists interviewed, but 
the general opinion is that these are simply forms of one common dis- 
ease, the variations being attributable to the season of attack or to 
varietal or vegetative conditions of the potato. Frank’s specimens of 
‘‘tief-schorf” were seen in the museum of the board of health at Berlin 
and have the appearance of the common American scab. 

As to the cause, opinions differ.“ The German pathologists who have 
given most attention to this disease consider the commonest German 
form to be due toa fungus similar to Thaxter’s Oospora scabies, but 
perhaps not identical with it. In Holland doubt was expressed as to 
the common Dutch form of potato scab being a fungous disease at all, 
while in Belgium and France it is considered a parasitic disease, but 
due to bacteria (JM/crococcus pellicidus Roze) rather than to fungi.” In 
England Vospora scabies is held responsible for only the minor part of 
the trouble, Sorosporium scabies Fisch. being the commoner parasite. 


VARIETAL RESISTANCE TO SCAB. 


As already stated, none of these scab diseases has proved of sufti- 
cient economic importance to attract much attention from practical 
potato growers in Europe. Probably most of the diseases would yield 
to the seed-disinfection treatment practiced for potato scab in America, 
yet this is nowhere used by European growers, so far as learned. 
The only valuable data as to the relative susceptibility of varieties to 
scab are those furnished by the reports of Professor Eckenbrecher,’ 
whose results point to certain newer varieties as being comparatively 
free from scab, while others are quite susceptible. Age of the variety, 


a@See Sutton, A. W., Potatoes, Jour. Roy. Soe., XIX, pp. 387-430 (1896). 

> Kampfbuch, p. 170. 

«These conclusions are the result of conference’ with a number of men and reference 
to the publications of others. Among those who have given this disease careful con- 
sideration in their respective countries and whose opinions were learned are Professors 
F. Kriiger, Berlin; Ritzema-Bos, Amsterdam; Marchal, Gembloux; Roze and Dela- 
croix, Paris; Massee and Cooke, England. 

4Roze, E., Histoire de la pomme de terre, Paris, 1898, p. 275. 

eC. von Eckenbrecher, Berichte Deutsch.-Kart.-Kult.-Stat., 1903 and earlier. 


16 DISEASE RESISTANCE OF POTATOES. 


however, seems to be of less consequence than it is with the blight, for 
while one of the least resistant is Dabersche, an old standard, one of the 
most resistant is Richter’s Imperator, also an old variety. Other 
varieties reported as preeminently scab resistant are [rene and Pro- 
fessor Wohltmann. Boneza and Pomerania are reported resistant, 
but to a less degree. Seed of all these have been secured for trial in 
this country. Early Rose is found especially liable to scab there as it 
isin America. It will be better, however, to postpone the discussion 
of resistance to scab as shown by American varieties of potatoes until 
the latter part of this publication. 


OTHER SCAB-LIKE DISEASES. 


Rhizoctonia.—This was seen on potato tubers in Europe even more 
commonly than it occurs in America, but most European pathologists 
regard it as nonparasitic. Those who consider it as a facultative para- 
site attribute little injury to it generally, although some think it capable 
of causing rot, as described by Frank.¢ 

Spongospora solani Brun.—This fungus is said? to cause a seab-like 
disease in the north of Europe. This disease occurred” to a consider- 
able extent on potatoes secured by the writer’s order from Groningen, 
Holland. 

Superficial scurvy diseases attributed to other fungi occur in Europe. 
Frank @ describes one such caused by Phellomyces sclerotiophorus, but 
Doctor Appel told the writer that this had not proved serious in Ger- 
many. Johnson’ reports it as causing some trouble in Ireland. In 
England another scurf disease of somewhat similar appearance is attract- 
ing attention. It occurred on the Eldorado seed potatoes imported 
with our order from Scotland, and the cause is apparently the fungus 
Spicaria nivea Hors.’ 

Ocdomyces leproides Trabut.—This is a fungus’ which attacks the 
young sprouts or eyes of the tubers, stimulating them to abnormal, 
cauliflower-like growth, of which the dark color has given rise to the 
popular name black- scab. The writer did not see this disease on the 


“Kampfbueh, p. 194. 

b Tbid., p. 176. 

¢ Found and identified by James Birch Rorer. 

@Kampfbuch, p. 182. 

e Johnson, T., The Diseases of the Potato and Other Plants in Treland, Journal, 
Dept. Agric. Trelinndl Vol. III, No..1 (1902). 

f So identified by Dr. Ernst A. Bessey. Dr. Thomas Milburn, of the Midland Agri- 
cultural and Dairy Institute, England, advises the writer by letter that he is engaged 
in a special study of this disease. He finds it to occur on most English varieties, 
Evergood and Sutton’s Flourball being by far the worst, while East Anglian and 
Sutton’s Discovery are the only ones he has found entirely free from it. 

9 This also occurs on beet roots. See Massee, Geo. A., Textbook of Plant Diseases, 
p. 225. 


— = 


POTATO STEM DISEASES. iif) 


Continent, but while in England Dr. M. C. Cooke sent specimens, as 
did also Dr. John Wilson, from Scotland. It is regarded as capable of 
causing much harm, but fortunately has not as yet become common. 


POTATO STEM DISEASES. 


BLACKLEG. 


_ *Schwarzbeinigkeit,” or blackleg, is a name applied to a disease 
seen commonly in central Europe. It is characterized by the black- 
ening and rotting of the main stem, accompanied by a checking of the 
growth, uprolling, yellowing, and ultimate death of the leaves, and 
more or less rotting of the tubers. It has been exhaustively studied 
in recent years by Appel? in Germany, who concludes that it is due to 
bacteria (Lacillus phytophthorus, and perhaps other allied species), 
which either enter from the soil or are carried in the seed tuber. - 
These start the disease below ground, the rot proceeding, as a rule, 
from the seed tuber to the young plant. Appel’s conclusions are gen- 
erally accepted by pathologists in Germany, Holland, and Belgium, 
and, so far as learned, in England. 

The writer had opportunity to see much of the German disease in 
the vicinity of Berlin, and to verify Doctor Appel’s observations, in a 
measure, in his own laboratory. Later (August) the same malady was 
seen in England, where it is said to be common, though apparently 
less troublesome than Appel reports it from Germany. All of the 
personal observations of the writer therefore lead to an indorsement 
of Appel’s conclusions both as to the bacterial nature of the disease 
and as to its widespread occurrence and economic importance. 

In France a similar if. not identical disease is attributed to. bacteria. 
Delacroix, who has studied this, considers the organism causing the 
disease, which he calls Bacillus solanicola, to be specifically distinct 
from that described by Appel. He kindly showed the writer speci- 
mens of the French disease in his garden in Paris, but it was ina 
stage so much more advanced than that seen in Berlin that a comparison 
of the two does not seem justifiable. 

Of course, the possible occurrence of these stem diseases in America 
was kept in mind. The writer has never seen much, if any, of the 
same trouble in potato fields in the Northeastern States and adjacent 
Canada which have come under his observation. Certainly it is not so 
common in America as it is in Kurope. The symptoms correspond, 
however, somewhat closely to certain diseases attributed to Rhizocto- 
nia in the South and West.? 


«Appel, O., Arb. aus Biol. Abt. Gesundheitsamte, 3: 364 (1903). 
» See especially Selby, A. D., Ohio Exp. Sta. Buls. 139 and 145, and Rolis, F. M., 
Colo. Exp. Sta. Buls. 70 and 91. 


8838—No. 87—05——3 


18 DISEASE RESISTANCE OF POTATOES. 


The only remedies proposed or practiced for blackleg in Europe 
consist in selection of sound seed, careful avoidance of rotten tubers 
or those taken from a diseased crop, attention to rotation of crops, 
since the germs may persist and accumulate in the soil, and the use of 
disease-resisting varieties. None of these methods has been tested 
carefully enough to establish its merits. ° 

The chief evidence as to varietal susceptibilities or resistance to 
blackleg is that of Appel.“ He states that while no varieties have 
been shown to be entirely free from the disease, the evidence is that 
thick-skinned, starch-rich, late varieties are in general more resist- 
ant than thin-skinned, starch-poor, early varieties. The Dabersche he 
finds to be the most resistant of the widely used German sorts, while 
the Rose varieties are especially liable to blackleg. Contiguous plats 
of the Dabersche and White Rose in Doctor Appel’s grounds showed 
this difference very clearly at the time of the writer’s visit in July, 
1904, the Rose being badly diseased, the Dabersche but slightly. At 
the Deutschen-Kartoffel-Kultur-Station distinct evidence of variations 
in disease resistance was found. It was stated at this station that in 
the trial grounds the red varieties were in general found to be more 
resistant than the white. | 

A considerable development of this disease was noticed at the Uni- 
versity Farm, Cambridge, England, where extensive variety trials were 
being conducted. The observations of Mr. H. Henshaw, who had imme- 
diate charge of these, were in agreement with those of Doctor Appel 
as to the association of the disease with the rot of the seed tuber, and 
he regarded it as a bacterial malady. There was a difference in the 
amount of the disease on different varieties, but Mr. Henshaw found 
none wholly free from it. At the time of this visit Factor and Up-to- 
Date were noticeably freer from the trouble than any other varieties 
there under trial. They were about alike in this, as, indeed, in all 
other characters. Doctor Delacroix cited as especially resistant to his 
bacterial disease in France the variety La Czarine. He also stated 
that the variety Geante Bleue is resistant, but to a less degree. Lau- 
rent observed in his work on bacterial diseases that the maximum of 
resistance was shown by the varieties Chardon, Pousse-debout, and 
Chave. . 


OTHER STEM DISEASES. 


Several other potato stem diseases have been reported in Europe, 
but since the writer did not study them in the field and learned noth- 
ing beyond what is recorded in literature they will be passed with brief 
mention. Prillieux and Delacroix have described another bacterial 
stem disease in France attributed to Bacillus caulivorus. Rhizoctonia 
is common on potato stems as well as tubers, but none of the patholo- 


4Arb. aus Biol. Abt. Gesundheitsamte, 3: 408 (1903). 


LATE-BLIGHT AND ROT. | 19 


gists conferred with regards it as a parasite of importance on either. 
Professor Marchal, of Gembloux, said, however, that he sometimes 
found a basidiomycetous fungus, //ypochnis solani Prill.,“ causing a 
blackleg-like stem disease in Belgium. Prof. T. Johnson, of Dublin, 
sent specimens of Sclerotinia sclerotiorum, which at times causes a 
destructive stem disease in Ireland, called ‘‘ yellow blight.” He also 
reports” the occurrence in Ireland of bacterial stem diseases, including 
what he believes to be the malady termed brown-rot in America and 
shown by Dr. Erwin F. Smith to be caused by Bacillus solanacearum. 
In none of these cases was anything learned as to relative varietal 
resistance to the disease in question. 


LATE-BLIGHT AND ROT DUE TO PHYTOPHTHORA INFESTANS. 


Late-blight and rot due to Phytophthora infestans occurs in Europe 
even more widely and destructively than in America, being recog- 
nized as the most important malady in all the countries visited by 
the writer—Italy, Austria, Germany, Holland, Belgium, France, and 
the British Islands. There was less unanimity of opinion than was 
anticipated as to the probable life history of the fungus, and conse- 
quently as to the remedial treatment. While all agreed that it lives 
over in the seed tubers, the opinion was frequently expressed, by 
scientific and practical men alike, that there is probably some hiberna- 
tion in the soil, either in tubers left in the field or in the haulms. 
Possible hibernation in the tubers is to be associated with the fact that 
potato tubers left in the soil are not killed by frost in southern Eng- 
land and the south of Europe. The value of Bordeaux mixture as a 
remedy is recognized generally. Spraying is consistently advocated 
-and practiced in the British Islands—especially Scotland and Ireland— 
the Netherlands, Italy, and portions of France. Germany has been 
surprisingly backward in accepting or even fairly testing this remedy. 
The reason seems to be that some of the experiments conducted by 
scientific men have shown injury to the plants, and so they have pro- 
nounced against it. No other remedial treatments are in common use. 
No one indorsed or practiced any method of seed disinfection. No 
particular culture methods were advocated or condemned other than 
attention to fertilization. There was a general agreement that 
excessive use of nitrogenous fertilizers, either chemicals or composted 
manures, increases the loss from this disease. Much attention is being 
given, however, to the relation of varieties and of source of seed to 
disease resistance, and the results are of sufficient importance to merit 
the somewhat detailed report which follows. 


4Probably identical with Corticium vagum var. solani Burt, which is the fruiting 
stage of the common Rhizoctonia of the potato. 

» Johnson, T., Diseases of the Potato and Other Plants in Ireland, Journal, Dept. 
Agric. Ireland, III, No. 1 (1902) 


20 DISEASE RESISTANCE OF POTATOES. 


RESISTANCE AS SHOWN TOWARD LATE-BLIGHT AND ROT. 
HISTORICAL STATEMENT. 


Doubtless examination of the earlier writings upon the potato disease . 
known as late-blight and rot would show that from the beginning of 
its ravages differences have been observed in the resistance of varieties. 
Any such records of the thirty years from 1845 to 1875 would have no 
practical value now, since the varieties then in use haye passed out of 
culture; nor would they have much scientific significance, owing to the 
lack of exact knowledge then as to the cause of disease. Going back to 
the origin of the varieties still in cultivation, it is found that in the early 
seventies an unusual amount of attention was focused upon the matter 
of the potato disease, as to causes and remedies. Ninety-four essays 
secured by the Royal Agricultural Society of England in 1872 showed 
agreement that an underlying cause was the degeneracy of the varieties 
then in culture.“ The necessity for the production of new varieties 
was emphasized. The introduction of improved American varieties 
into England at about this period was most beneficial and stimulating 
to the potato specialists of that country.” These varieties were made 
the parents in further breeding. The best production of this revival 

yas the Magnum Bonum, originated by James Clark from a cross of 
Early Rose with Victoria and introduced by Sutton in 1876. Experi- 
ence with this variety laid the foundation for belief in‘possible disease 
resistance in potatoes both in England and on the Continent. Magnum 
Bonum soon became the standard main-crop variety of Great Britain 
and so continued until within the last fifteen years, when it yielded to 
Up-to-Date and others. It is still in considerable favor on the Conti- 
nent. 

Charles Darwin in 1877-78 became interested in the possibilities of 
disease-resistant breeding. Francis Darwin® states that Mr. James 
Torbitt, of Belfast, bred and selected varieties to secure disease 


4See Jour. Roy. Agr. Soc. Eng., XX : 291 (1884). 

b Dean, A. Potato Improvement in the Past Twenty-five Years. Jour. Roy. 
Hort. Soc., XII : 41 (1890). 

Mr. ©. G. Pringle, who was then the foremost potato breeder in America, states 
that the demand from Europe for new American varieties was very active at that 
period, and continued until the fear of the Colorado potato beetle led to the prohi- 
bition by European governments of further importations of potato tubers from Amer- 
ica. Thereupon Mr. Pringle supplied B. K. Bliss & Sons with specially hybridized 
potato seed, which was sent abroad in considerable quantity. 

¢ Life and Letters of Charles Darwin, II, pp. 519-522. 

The writer learns from Prof. T. Johnson, of Dublin, that Torbitt has been dead 
some twenty years. He was in the wine trade and raised varieties of potatoes for 
the berries, or seed balls, which he used as a source of material for wine. All of 
his varieties have disappeared except one coarse, red one, which was and is proof 
against disease. 


ee LLL = ee a 


HISTORICAL STATEMENT. 21 


resistance, his method being to cross-fertilize, rear the seedlings, and 
expose them ruthlessly to infection, retaining only those showing 
some degree of resistance. In this work he received much encour- 
agement and some financial aid through Charles Darwin. 

A committee of the English House of Commons, reporting in 1880 
upon the potato disease, found all its witnesses concurring in the 
necessity for the production of new varieties with increased disease 
resistance.“ Parliament was asked to give financial aid for experi- 
ments aiming to produce new and disease-proof varieties, but it did 
not do this. Earl Cathcart, in commenting on this report, states 
that— 

- All potatoes have deteriorated in their disease-resisting powers. A variety from 
seed takes four to six years for its establishment, and under the most favorable cir- 
cumstances a good variety might be expected to degenerate in twenty years. The 
production of new varieties is of national importance. 

Apparently through the influence of Cathcart, Baker’? was led to 
make an exhaustive comparative study of the genus Solanum in order 
to advise as to the relation of the cultivated varieties to the several 
wild species of the American continent preparatory to breeding experi- 
ments in which these might be used. As a result, two species were 
considered worthy of further trial in the attempt to improve disease 
resistance, viz, the Darwin potato, S. maglia, from the Chonos Archi- 
pelago, and the Uruguay potato, S. commersonii. Cathcart furnished 
Sutton with the former and he hybridized it with the common S. tudber- 
osum. Sutton’ reports that, beginning in 1886— 

Although many hundreds of flowers of S. maglia were artificially fertilized with 
pollen from cultivated varieties, only five. were successful, resulting in five berries. 


_ From these but two seedlings were secured and only one of these showed any prom- 


ise whatever, the second having to be grown under glass to prevent its dying. 
* * * This hybrid, although a vast improvement on S. maglia, is far behind the 
ordinary potato in appearance, crop, and quality. The seedling * * * grown for 
eight years, in 1894 was slightly diseased, although previously free from attack. 


Sutton still -has the S. maglia and this hybrid in propagation at 


' Reading, where the writer saw them in August, 1904. Phytophthora 


was then more rampant on the foliage of both of these and on S. com- 
mersonii, Which he also has, than on the average potato plants in his 
fields. No hybrids of S. commersonii had been secured by him pre- 
vious to this. Mr. Lasham, potato specialist for the firm, has been 
giving renewed attention to the possibilities of species hybridization, 
and showed the writer balls which he considered to contain hybrid 
seeds of S. tuberosum * commersonii. 


“Jour. Roy. Agr. Soc. Eng., XX: 291 (1884). 

> Baker, J. G., A Review of the Tuber-Bearing Species of Solanum. Jour. Linn. 
Soc., London, X X, 489-507 (1883-84). 

eSutton, A. W., Potatoes, Jour. Roy. Hort. Soc., XIX, 387 (1896). 


22 DISEASE RESISTANCE OF POTATOES. 


Interest in the possibilities of S. commersonii has recently been 
stimulated by the experiments inspired in France by Prof. E. Heckel,¢ 
who believes in the economie possibilities of this species when improved 
by longer culture and perhaps by hybridizing, and has distributed it 
quite widely in France with this end in view. One French horticul- 
turist, Labergerie,? claims already to have succeeded in producing a 
variety of edible quality, large yield, and superior disease resistance. 
This was seen growing in the grounds of Vilmorin-Andrieux & Co., at 
Paris. This firm was not yet convinced of the practical value of the 
plant, and since Labergerie refuses to send out any of these potatoes 
for trial at present, judgment must be reserved. 

Returning to the consideration of the varieties of the common potato, 
it is found that during the last decade increasing attention has also 
been given to their comparative disease resistance. This has been well 
summarized by Prunet.“ The main facts developed are as follows: 
Sorauer” in 1896 considered the evidence to date as showing that the 
highest degree of disease resistance was possessed by Magnum Bonum, 
the following showing some degree of resistance: Blaue Riesenkartoffel, 
Richter’s Imperator, Athene, Reichskanzler. Rostrup,’ writing: about 
the same time from Denmark, places Magnum Bonum at the head as 
a disease-resisting variety, with Richter’s Imperator and Champion 
as somewhat resistant. 

In this connection it is noteworthy that Magnum Bonum has yielded 
its place in popular favor in Great Britain for main-crop purposes to 
Up-to-Date and other varieties, even while holding its reputation on 
the Continent. During the last decade these standard varieties in turn 
have been *‘running out,” and the demand in England for something to 
take their place has stimulated potato breeders and seed specialists to 
direct their attention very generally to the development of a disease- 
resisting main-crop variety. The greatest efforts in breeding have been 
made during the last four years, while speculation in the most promising 
of the varieties produced has been at fever heat for the last two years, 
during which time many new varieties, more or less disease resistant, 
have been pushed to the front. There are now so many potato special- 
ists in Great Britain breeding and handling varieties of reputed disease 
resistance that it is impracticable to mention all. Archibald Findlay, 


« Heckel, E., Sur le S. commersonii, Rey. Hort. de la Soe. d’ Hort. et de Bot. des 
Bouches-du-Rhone, No. 581, pp. 200-206 (December, 1902); also, Contrib. 4 l’étude 
botanique de quelques solanum tuberiferes, Ann. de la Faculté des Sciences de 
Marseille, vol. 8 (1895). 

» Labergerie, M., Le Solanum commersonii et ses variations, Bul. Soc. Nat. d’ Agric. 
de France, March, 1904. 

¢Prunet, A., Le mildieu de la pomme de terre, Rev. de Viticult., X VII, 663; 
XVIII, 97 et seq. (1902). 

¢ Zeitsch. f. Pflanzenkr., VI, 284 (1896). 

¢ Tidsskrift f. Landbrugets Planteavl, 1895, 1896, 1897. 


—S 


—_— 


DISEASE RESISTANCE AND VEGETATIVE VIGOR. 23 


of Mairsland, Auchtermuchty, Scotland, has originated some varieties 
of high repute; Sutton & Sons, of Reading, England, have also taken a 
prominent part in this work. Scotland-grown seed of all the leading 
varieties can be secured from Thomas Scarlett, of Edinburgh, and Mr. 
Searlett has some promising varieties of his own introduction. Any- 
one desiring more specific information should secure the publications 
of the National Potato Society from its secretary, Walter P. Wright, 
Postling Vicarage, Hythe, Kent, England. 

Meanwhile the German Potato Station has been making extensive 
tests, doing some breeding and encouraging several potato breeders.” 
These efforts have not been directed primarily to disease resistance, 
but the station has taken note of this feature and published the data 
regarding all varieties tested. 

While the results of this work in Europe have been collected or 
correlated by no one, the writer was able to gather considerable infor- 
mation, in addition to that already referred to, from conversation with 
potato specialists, especially in Great Britain. This, together with 
what has been learned in America, is made the basis of the following 
discussion. It is to be regretted that it is impracticable to give detailed 


credit in sone cases to those who kindly furnished the information. 


THE MEANING OF DISEASE RESISTANCE. 


Although potato specialists, especially in England, apply the term 
**disease proof” to their favorite varieties, this is not to be taken liter- 
ally. No variety has as yet shown itself to be absolutely proof against 
disease. The writer personally collected leaves infected with the 
blight fungus from two varieties which were said to be ‘disease 
proof.” Absolute resistance against the blight fungus has not as yet 
been and may neverbe secured. Varieties are known, however, which 
show a relatively high degree of disease resistance. This may be 
shown in the delay in date of appearance of the blight on the leaves 
or its slower progress after appearing, and still more clearly in the 
relatively small amount of loss from rot of the tubers.’ Most of the 
exact observations made in Europe have been based on this latter 
difference. 


DISEASE RESISTANCE AND VEGETATIVE VIGOR. 


Disease resistance and vegetative vigor are closely associated, 
although the factors involved are not necessarily identical. In any 
consideration of the problems of the life and death of the potato plant 
it must be remembered that the potato has two natural methods of 


“@The most active of these are Paulsen, Cimbal, Richter, and Dolkowski. Graf 
Arnim-Schlagenthin, of Nassenheide in Pommern, is an extensive breeder and dealer 
in new varieties, as is also-F. Heine, of Hadmersleben. 


24 DISEASE RESISTANCE OF POTATOES. 


reproduction, the true seed produced in the berries or ‘* balls” follow- 
ing the blossoms, and the tubers produced below ground. Repro- 
duction by seed is a sexual process, that by tubers is vegetative. 
Both are exhaustive of vitality. The two are in a certain sense 
physiologically opposed to each other and can not well be carried on 
at the same time by the plant. Under the natural conditions of the 
wild potato plant in Mexico, and doubtless elsewhere, seed production 
precedes tuber formation. In Europe and northern America, with a 
shorter season and intensive culture, the two processes overlap. As 
a result there is, just after the potato plant comes into blossom, a period 
when the natural tendencies within the plant toward seed production 
above and tuber formation below are such as to subject it to unusual 
physiological stress. This has been termed the ‘‘critical period” @ in 
the development of the potato plant. 

Usually the blossoms fall without the setting of the fruit, and the 
plant then passes into a stage where its energies are devoted to tuber 
formation alone. Once well started upon this vegetative period, its 
growth is more or less indeterminate, i. e., there is no clearly defined 
natural terminus to the life of the cultivated potato plant. Instead, 
there is the gradual decline in vegetative vigor-which may prepare the 
way for early-blight and other diseases characteristic of weakling 
plants. It is noteworthy that the destructive attacks of the late-blight 
fungus occur, as a rule, after the blossoming period has passed. So 
far as the evidence goes it seems to suggest that high vegetative vigor 
enables the plant to ward off in some degree the fungus attack. 

There is, moreover, a natural decline in vigor, or ‘‘running out,” 
with the age of the variety. The length of life of a variety depends 
upon numerous conditions, and is an indefinite matter in any case. It 
is ordinarily placed by potato specialists at from twelve to twenty 
years. Asa variety begins to ‘‘run out” it apparently shows, among 
other things, a lessened degree of general disease resistance. Thus 


Magnum Bonum had the highest reputation in this respect from its: 


origin in 1876 until about 1890 in Great Britain. On the Continent it 
has remained longer in favor. Up-to-Date has held a like place during 
the last decade in Great Britain, and Richter’s Imperator has a similar 
record in Germany. These statements are based upon the popular 
verdict, not upon exact experiments; but the belief that disease resist- 
ance decreases with the age of the variety is firmly established in the 
minds of specialists in potato culture in Great Britain, at least so far 
as concerns resistance to Phytophthora. | 

There is little definite evidence regarding the relation of vegetative 
vigor to resistance to other diseases, but so far as it is formulated it 


4 See Jones, L. R., Certain Potato Diseases and Their Remedies, Vermont Exp. Sta. 
Bul. 72: 4 (1899); also The Diseases of the Potato in Relation to Its Development, 
Trans. Mass. Hort. Soc. (1903), Part I: 144. , 


a 


IMPROVEMENT BY SELECTION. 25 


favors the general applicability of the idea. The production of ber- 
ries, or seed balls, is held by some English breeders to be an indica- 
tion of such vigor, and therefore presumably of disease resistance, 
although no one claims that the absence of these is equally strong 
evidence in the opposite direction. It is worthy of passing note that 
berries are formed much more commonly in Europe, especially in 
Germany and Holland, than in America, apparently because of cli- 
matic differences. 


THE RELATION OF HYBRIDITY TO DISEASE RESISTANCE. 


By a ‘‘new variety” of potato, as the term is commonly used, is 
meant one recently developed from the seed. Sports may appear and 
are indeed frequent in some varieties. These will, however, be men- 
tioned later. The seed in all cases presumably represents a sexual 
origin, i. e., comes from a fertilized flower, but this may have been 
either self-fertilized or cross-fertilized. One would expect greater 
vigor to result from the cross-fertilization, and potato breeders are of 
the opinion that it is secured. On the other hand, while varieties 
recently originated from seed may show a high degree of disease 
resistance, this is not necessarily the case according to the verdict of 
English and German breeeders, many new varieties proving as sus- 
ceptible as old ones. 

Reference has already been made to the work of Cathcart and Sut- 
ton in England and of Heckel and Labergerie in France based on the 
hope of advantage from using one or another of the wild Solanums 
for such hybridizing. There are interesting possibilities along this 
line, since there are many wild forms of S. tuberosum in addition to 
other species of tuberiferous Solanums. Thorough trials of these are 
now being made by Stuart “ at the Vermont Experiment Station which 
will be reviewed later in this paper. It should be emphasized at once, 
however, that while the use of the wild Solanums does offer interest- 
ing possibilities, there is no record of practical success from their use, 
if we except the doubtful one of Labergerie’s variety of S. commer- 
sonii previously referred to. On the other hand, great practical 
improvements have been secured by various breeders from crossing 
varieties of cultivated potatoes. 


IMPROVEMENT BY SELECTION. 


All plants tend to vary. One of the commonest ways to improve 
plants is by selection from among the varying individuals. For the 
first two years after their origin, variants, or ‘“* rogues,” are not uncom- 
mon with the seedling potato, but the variety is ‘‘ fixed” by the weed- 
ing of such rogues before it is distributed for general culture, and 


4Stuart, W., Disease-Resistant Potatoes, Vermont Exp. Sta. Bul. 115 (May, 1905). 


26 DISEASE RESISTANCE OF POTATOES. 


thereafter very little variation occurs. Some modification is to be 
expected, nevertheless, and among other things there may be varia- 
tion in disease resistance. It seems worth while, therefore, to keep a 
lookout for individual plants which show especial resistance to the 
blight when this is epidemic about them. The tubers of such indi- 
viduals deserve to be carefully saved apart and planted, in order that 
it may be seen whether their resistance is a fixed and inheritable char- 
acter or the result of some chance difference in environment. Such 
selection has already been undertaken by three persons to our knowl- 
edge: Appel in Germany,@ Stuart in Vermont,’ and Fraser in New 
York; but no practical results have as yet been secured. 


ARE EARLY OR LATE VARIETIES THE MORE RESISTANT? 


The blight never becomes serious until the midseason of potato 
growth is passed; thus in our Northern States its worst ravages come 
in August and September. Therefore early varieties, as a rule, escape. 
But this is simply because they mature before the blight is epidemic. 
So far as evidence has been secured, both in America and Europe, 
when the early varieties are planted late enough to expose them to 
the disease alongside of the later ones, the early varieties as a class suf- 
fer the worst. For example, the most complete destruction by rot 
which the writer ever saw was with a late crop of Early Ohio potatoes 
attacked by the disease in September. Woods,? of Maine, has also 
found early varieties especially susceptible. 


RELATION OF SOURCE OF SEED AND CULTURAL METHODS TO DISEASE 
RESISTANCE. 


The opinions of highly intelligent potato growers in Great Britain 
are especially worthy of note. One of the first concerns with them is 
the source of their seed. Mention has already been made of the experi- 
ments at Sutton’s grounds, showing the superiority of northern-grown 
seed. It was found to be the general verdict of practical men in 
Europe that northern-grown seed is more highly disease resistant, as 
well as more productive. If one is to aim for the best results in 
health of crop, therefore, attention should be directed to quality and 
source of seed as well as to variety. 

Practical men as well as scientists generally agree that methods of 
culture also determine to a considerable degree liability to disease. 
These act not only indirectly as they affect moisture content or other 
physical conditions of the soil, but more directly as they affect the 


«Appel, Otto. Die diesjiihrige Phytophthora-epidemie, Deutsche Landw. Presse, 
XXIX: 685 (1902). 

> Vermont Exp. Sta. Bul. 115, p. 139. 

¢ Reported in correspondence. 

@ Woods, C. D., Maine Exp. Sta. Rpt., XIX: 181 (1903). 


COMPOSITION AND CHARACTER, AND ROT RESISTANCE. 27 


vigor or inherent disease resistance of the plant itself. Seed dealers 
in this country, as well as in England, have also expressed a prefer- 
ence for seed from a crop that has not been very highly fertilized, 
especially with nitrogenous manures. The most detailed study along 
this line is that of Laurent,’ whose results indicate that nitrogenous 
fertilizers predispose both the foliage and the tubers of the plants to 
the attacks of Phytophthora. This is in harmony with the general 
opinion of practical potato growers, that high manuring increases the 
liability to disease. 


COMPOSITION AND CHARACTER OF TUBERS AS RELATED TO ROT 
RESISTANCE. 


There is considerable evidence that the chemical composition of the 
tubers bears a direct relation to resistance to rot. Paulsen? claimed 
that varieties rich in nitrogen compounds are less resistant to disease 
than those rich in starch. He classed most of the early varieties in 
the first category and found the larger percentage of the second class 
to be of the late varieties. The table varieties of better quality were 
also of the first category. 

Petermann’ has recently made field and laboratory studies at the 
Belgium Experiment Station. These led him to practically the same 
conclusions, viz, (1) that varieties richer in amids are more liable to 
rot, although of superior table quality; and (2) that varieties relatively 
richer in starch, including several recently originated German factory 
varieties, are less liable to rot but are of inferior quality for table use. 

Sorauer”? has published similar’ conclusions, and he made more 
detailed statements as to his belief on this point in conversation last 
summer. In general, he believes that the varieties richer in protein 
are more liable to disease, while those richer in starch are more 
resistant. This probably explains his observation that the yellow- 
fleshed varieties, which are in higher repute in Germany for table use, 
are more liable to disease’ than the white-fleshed varieties which are 
grown for factory purposes. His further observations in harmony 
with this idea are that the thicker and rougher skinned red varieties/— 
e. g., Dabersche—have a higher degree of disease resistance, coupled 
with relatively high starch and low protein content, whereas the thin- 
skinned white varieties, which are more liable to the disease, have 


«Laurent, Recherches exp. sur la mal. des plantes, Ann. Inst. Past., XIII, pp. 
1-48 (1899). 

» Biedermann’s Centralbl. Agr. Chem., 1887, p. 107. 

¢ Bul. Inst. Chim. et Bact. Gembloux, 70 (1901). 

¢d Jahresber. d. Sondersaussch. f. Pflanzensch., XII and XIII (1902 and 1903). 

¢See evidence of this also in Jahresber. d. Sondersaussch. f. Pflanzensch., XIII, 
1903. 

f See also statement that red varieties are in general more resistant; Jahresber. d. 
Sondersaussch. f. Pflanzensch., XII, 1902. 


28 DISEASE RESISTANCE OF POTATOES. 


proportionately less starch and more protein. As a rough empirical 
test he considers that if, when a fresh tuber is cut open, the flesh 
browns quickly on exposure to the airand the vascular bundles darken 
soon, it is evidence of high protein content, and therefore of liability 
to disease, whereas the reverse condition is evidence of probable dis- 
ease resistance. Professor Sorauer also emphasized to the writer his 
belief in the relation of soil, manuring, or other cultural condition to 
disease resistance. The testimony secured elsewhere in Germany, as 
well as from American sources, is in harmony with these ideas so far 
as it goes. 

While character of skin is probably a less reliable index to rot 
resistance than is chemical composition, yet the writer has learned of 
several potato experts in America, as well as others in Europe, who 
regard the red varieties, especially such as have a rough skin, as less 
liable to rot than are the white thin-skinned varieties. The Dakota 
Red has often been cited as an example of this class in America. 
Abundant evidence can be found, however, especially from English 
experience, of high disease resistance coupled with a thin, white skin. | 
Breeders aiming at disease resistance need not turn from the white- 
skinned varieties. 


CHARACTER OF STEM AND FOLIAGE AS RELATED TO DISEASE RESISTANCE, 


In connection with the question of the relation of the character of 
the stem and foliage to disease resistance it is not safe to go far in 
generalizing. Mr. Lasham, potato expert and breeder with Sutton & 
Sons, considers that a stem that is strong, rough, and hard, almost 
woody at the base, is an important character if the plant is to be most 
highly disease resistant. The nature of the foliage is considered by 
others to be of greater importance, the preference being for small 
leaflets, rough and relatively thick rather than large and flabby. A 
rich dark-green is preferred to the lighter colored foliage. It seems 
inherently probable that the character and color of foliage should 
stand in close relation to the other matters which have with more cer- 
tainty been shown to be related to disease resistance, viz, general 
vigor and capacity for starch manufacture. Those giving further 
attention to disease resistance may, therefore, well bear these sugges- 
tions in mind. 


DISEASE-RESISTANT VARIETIES OF EUROPE. 


When all the factors are considered, two important things become 
evident: First, that no variety will maintain its disease-resistant qual- 
ities indefinitely, losing them sooner in one locality than in another; 
second, that no one yariety will be equally disease resistant in all 
countries, i. e., what is best suited to one may not be to another. 


DISEASE-RESISTANT VARIETIES OF EUROPE. 29 


Hence we must expect much conflicting evidence regarding the same 
variety, especially as grown under different conditions and in different 
countries. 

The most satisfactory way will be to summarize separately the evi- 
dence obtained in each country visited in Europe, followed by that 
from America. 

The greatest activity in breeding for resistance occurs in Great 
Britain, and next to this in Germany, while France has recently shown 
special activity in one line. There were secured from each of these 
countries all the varieties of especial promise for trial in America 
comprising a total of nearly one hundred. By no means all of these 
have an established reputation as being highly disease resistant, 
although all will be tested for this as well as other characters. 

Experience has shown that the transference of a variety from Europe 
to America, or the reverse, is likely so to disturb the equilibrium of 
the plant that the developments of the first year are scarcely normal. 
At least two years are necessary for the adequate testing of imported 
varieties. Commercial growers should therefore make only trial 
plantations of any European variety until its adaptability to American 
conditions has been proved. Seed of the varieties secured will be 
under trial at several points this season and next. While it does not 
seem worth while to publish the full list of these in advance, the fol- 
lowing are selected as representing the varieties or types most strongly 
indorsed in their respective countries for disease resistance. 


GREAT BRITAIN. 


More attention has been given to this question in Great Britain 
than anywhere else. Careful comparative observations are recorded 
-as to the relative disease resistance of all the leading varieties. The 
last report of the National Potato Society specifies the following eight 
varieties as the best for disease resistance, named in the order of their 
merit. The writer appended certain facts as to the origin and charac- 
ters of each. All are white skinned, white fleshed, of excellent qual- 
ity, of high general vigor, and heavy yielders. 


TaBLe I.—The most disease-resistant varieties of potatoes in Great Britain, as announced 
by the National Potato Society.@ 


Order When 
of | Name of variety. Originator. | sent Season. 
merit. out. 
HMERU ETO OOUIS ste a cecosc ne stigeecs tes eaies Bin @laviceececcdes ass <aee. 1896 | Medium late. 
2 \fDiseovery ...-- Sutton’ seer con. lees. 1903 | Late. 
‘Royal Kidney. 1) BID Olver eee sss aes aloo 1896 | Second early. 
De PNOLUNOIM NS taD coe 56 hs Jo0c nese soe sna hoe na COM easieddes See ecicecee 1901 Late. 
eared Ole Mew ely ssc = cece ese =| PARIS eee er elr ce-kcewes 1900 | Early. 
pu) eae dco hago ite 004 0 le eee ae \ Butler teen ccc ccmeh hs cee 1901 | Medium. 
Ra PEACLOUAG Outs Ste te cn cos cote eee. waco ke fee wee fan a o)aicis me 1903 | Medium late. 
(1 ADEN 70) Pe ee ee aes ia sa oa Aer DObDIGi ae aseiewicnis 2s owas sgn 1898 | Late. 


a Annual Report, National Potato Society, I: 36 (1904). 


30 DISEASE RESISTANCE OF POTATOES. 


Two points brought out by the foregoing table are worthy of espe- 
cial note: First, the remarkable success of one man in producing four 
of these eight varieties, showing that there is something more than 
chance in their development; second, the comparatively recent origin 
of all. For the latter fact there is probably a double explanation: (1) 
The loss of disease resistance with age of the variety; (2) the remark- 
able activity of British potato breeders during the last few years, with 
the aim above all else of securing increased disease resistance. 

It should be remarked that there are a number of more recent intro- 
ductions which promise to rank with the best of these varieties, but 
they have not as yet been so fully tested. 


GERMANY AND HOLLAND. 


The potato-growing industries of Germany and Holland have much 
in common, and similar or identical varieties are used. Data were not 
available for making so definite a selection as in Great Britain. The 
following are probably representatives of the highest grade of disease 
resistance achieved: 


Tasie I1.—The most disease-resistant varieties of potatoes in Germany and Holland. 


i ae ahed Color of 

Name. Originator. Season. skin. 
PLO OTG tens see noe te one Oe eee teie aaa Dolkowski ixese0 n=2 sce | Daten. .5 52 eee eaemenen White. 
BRCM Gs paces do oe Sane eee mse e- saeee | Paulsenygeesmcssnt eee | Medium late ......... Red. 
Gebeimrat Thiel. = 2: iioteseesee ees sees Richter’. seen ces een GO 505.2286 -beeeeee White. 
Professor Wohltmann .....< 2.2 --.:...2--.- Cimbal .2...-25-......|) Dabe 22 sa.sese eee Red. 
OUuCza ECan N ms ee ee ae reeeMey. 828. | GLO Wwskites een oeeeee | Medium late .....-... Do. 
MiPonhielmer wes acs to leet eee oe oe (Holland) ages eee | Medium early ........ White. 
PARP RTMIPers. 2.256 cence es eeeess ce ose ay =o) eee G0 2 aes eee | Late <:..2..ssecemeneee Do. 


The first five varieties have been imported from Germany. All of 
these are of the white-fleshed, starch-rich type which is being devel- 
oped there primarily for factory rather than for table use. ‘They are, 
however, reputed to be of fair quality except, perhaps, in the case of 
Irene. These, like the British varieties, are of comparatively recent 
introduction. 

The Holland varieties are of a similar type except that Kigenheimer 
has a yellowish-tinted flesh. 


FRANCE AND BELGIUM. 


The conditions are similar in France and Belgium. Neither country 
has varieties of much promise for trial in our Northern States. In 
Belgium the varieties recommended as being most highly resistant to 
blight and rot were the recently originated German factory types like 
those already mentioned, to which might be added Topas and Pro- 
fessor Maercker. The verdict in France was similar, Professor 
Maercker again being commended, along with the English varieties 


DISEASE-RESISTANT VARIETIES OF AMERICA. 31 


Magnum Bonum and Royal Kidney. The indorsement of these com- 
paratively late, white-fleshed, starch-rich potatoes in France and in 
the other continental countries is the more significant when one 
remembers that the chief aim with potato specialists there is to produce 
the yellow-fleshed potato rich in protein. 


DISEASE-RESISTANT VARIETIES OF AMERICA. 


~ Until two or three years ago no systematic attempt was recorded, 
so far as the writer has learned, to determine the relative resistance to 
blight and rot of potato varieties in America. Of course this would 
not imply that individual growers have not observed these differences, 
but they are rarely matters of record, and asa rule are based only 
upon limited observations. Only two or three potato-seed dealers are 
advertising disease-resisting varieties with any prominence this year, 
and these are in all cases comparatively new and little-tried sorts. 


INVESTIGATIONS AT THE EXPERIMENT STATIONS. 


Promising investigations have been inaugurated at several of the 
State agricultural experiment stations which should soon supply data 
for more reliable conclusions than are now possible. Woods,“ of the 
Maine station, and Green,’ of the Minnesota station, have each 
reported results of variety trials as to disease resistance. 

Woods found a marked difference in the ability of varieties to with- 
stand both blight and rot. As a rule the earlier varieties were soonest 
attacked. The variety Rustproof showed the highest degree of foliage 
resistance and also the least rot, viz, only a little more than 1 per cent, 
whereas the average of all varieties under trial was more than 30 per 
cent. 

Green found that the loss from rot varied widely with varieties, 
ranging from 1 per cent in the most resistant to 40 per cent in the 
least resistant types. Potatoes of the type of Sir Walter Raleigh and 
Rural New Yorker resisted rot better than those of any other class. 
Of the 49 varieties tested, only two, Clay Rose and an unnamed seed- 
ling, were practically free from rot. These trials will be continued. 

Observations on disease resistance have also been made by Macoun 
at the Central Experimental Farm, at Ottawa, Canada. He has kindly 
advised the writer, in correspondence, of his conclusions. The follow- 
ing varieties have shown especial disease resistance as judged by 
appearance of blight on the foliage: Holburn Abundance and Pro- 
fessor Maercker are most resistant, with Swiss Snowflake, State of 
Maine, and Rurat Blush only a little less so. 


«Woods, C. D., Maine Exp. Sta. Report, XIX: 181 (1903). 
>Green, 8S. B., Potatoes at the University Farm, Minn. Exp. Sta. Bul. 87 (1904). 


32 DISEASE RESISTANCE OF POTATOES. 


WORK AT THE VERMONT STATION. 


. 


The most extensive work on potatoes has been done by Stuart, who 
two years ago inaugurated at the Vermont station variety trials as to 
disease resistance, supplemented by breeding experiments. Professor 
Stuart has kindly supplied the following summary of his results to 
date:¢ 

Eight varieties were under trial in 1903—Dakota Red, Enormous, 
Green Mountain, Rustproof, Squire, Sir Walter Raleigh, and two of 
Manum’sunnamed seedlings. In 1904 the same varieties were used, with 
the addition of June, Mammoth Gem, Minister, New Queen, State of 
Maine, Sutton’s Discovery, nine more of Manum’s seedlings, S. com- 
mersonii from Doctor Heckel, of France, a Peruvian variety from the 
United States Department of Agriculture at Washington, and four 
Mexican varieties furnished by Mr. C. G. Pringle. The latter included 
two cultivated varieties termed Monterey and Mexican, and the two 
wild species, S. polyadeniwm and S. stoloniferum. Observations have 
been made as to both foliage and tuber resistance. As to foliage, none 
was wholly free from blight, but there was a marked difference, some 
being quickly and entirely destroyed, while others suffered only 
slightly. In 1903 Rustproof headed the list in this respect, and Dakota 
Red was second. In 1904 those showing greatest foliage resistance 
were as follows, in the order of their resistance: Monterey, S. com- 
mersonii, S. polyadenium, Rustproof, Sutton’s Discovery, June, Mex- 
ican, Mammoth Gem, and Manum’s No. 3. Dakota Red did not equal 
its 1903 record. 

Judged by resistance of tubers to rot, Dakota Red made the best 
showing of the varieties which were tested for two seasons, but there 
was some rot in it both years. Of those added to the series of 1904 
several varieties gave a crop of tubers entirely free from rot, namely 
S. polyadenium, SS. COMMETSONAL, Sutton’s Discovery, June, and the 
two Mexican varieties. It is noteworthy that these are likewise the 
varieties which showed the least blighting of the foliage. Possibly 
the absence of rot is in some degree attributable to the lessened amount 
of infection of tubers from vines in consequence of this, although there 
were adjacent plants showing badly diseased foliage. 

Sclection.—In 1903 a few plants in the varieties grown were observed 
to remain green longer than the others. The tubers from these were 
saved and were planted in 1904 along with others of the same variety 
that succumbed much earlier. So far as could be noted no increased 
disease-resistant qualities were transmitted to the offspring of these 
plants. 


“For further details see Stuart, W., Disease-Resistant Potatoes, Vermont Exp. Sta. 
3ul. 115 (May, 1905). 


INFORMATION SECURED BY A CIRCULAR OF INQUIRY. 33 


In view of the limited number of varieties tested and the consequent 
restricted scope of observation, it would hardly be justifiable as yet to 
give an expression of opinion as to the probable outcome of such a line 
of investigation. These observations are to be continued on a much 
larger scale the coming season. 

Hybrid and other seedlings.—Seedlings were grown in 1903 from the 
Mexican species supplied by Mr. Pringle—S. polyadenium, S. stoloni- 
Serum, and S. bulbocastanum. This number has recently been aug- 
mented by S. verrucosum anda wild form of S. tuberosum. In 1904 
some of these were successfully hybridized with the cultivated potato. 

While it is yet too early to say very much regarding the possibili- 
ties of developing a commercially desirable disease-resistant variety of 
potato through the hybridization of wild species with our cultivated 
varieties, there seems good reason for believing that improvement may 
result from such crossing, especially as to disease resistance. 

In addition, a large number of seedlings of the common potato were 
grown in 1904. Among this number several vigorous-growing plants 
were noted, which remained quite green up to the time of digging. 
Some 50 of the more promising of these were saved for trial the com- 
ing season. Selections were based on vigor of vine, size and yield of 
tubers, and freedom from rot. 

One of the most interesting and instructive features of the seedling 
experiment was the object lesson it presented of the extreme vigor of 
some of the plants, showing quite plainly that one of the best sources 
for increasing the vigor lies in the production of new varieties from 
seed. Proper fertilization and good tillage are also important aids in 
increasing the vigor and disease-resisting powers of the vine. 

While the data in hand do not warrant broad generalizations, the 
following inferences are drawn by Professor Stuart: 

(1) Some varieties are less subject to vine injury than others. 

(2) Some show a greater tuber resistance to rot than others. 

(3) With some there seems to be a fairly close relation between resist- 
ance of vine to disease and of the tuber to rot. 

(4) Selection has not given visible increase of resistance. 

(5) Hybridization and the growing of seedling plants, followed by 
careful selection, seem to offer a more logical method of securing 
disease-resistant varieties than does selection. 


INFORMATION SECURED BY A CIRCULAR OF INQUIRY. 


In order to secure all information possible as to the relative merits 
of the varieties now before the public, a circular letter of inquiry was 
sent recently to various experiment station officers and to about two 


34 DISEASE RESISTANCE OF POTATOES. 


hundred potato specialists in the United States and Canada¢ asking 
their opinions and the basis for them. There was a surprising lack 
of agreement upon any one variety as being especially disease resist- 
ant. The replies indicate several things: (1) That very few American 
potato specialists have up to the present time given careful attention 
to this question; (2) that there are few varieties in common use which 
have preeminent worth as disease resisters; and (3) that in so large a 
geographical area local conditions affecting culture and disease are so 
widely variant as to prevent close comparisons or broad generaliza- 
tions. No less than 38 varieties were commended as showing resist- 
ance to blight or rot. Of these, 26 were commended only once, while 
the other 12 were favorably spoken of by two or more persons. Still 
other varieties were mentioned, but not with sufficient positiveness to 
entitle them to a place in the following list. Those named more than 
once, with the locality in which they were indorsed, are as follows: 


Tasie IL1.—The best varieties of potatoes in the United States and Canada, as réported by 
various experiment station officers and potato specialists. 


Number | 
Variety. eae Localities in which commended. 
ments. 
Daluta Red. so 452 nee eS. een = pe eee 10 | Canada (4); Maine (3); Massachusetts; 
: Michigan; New York. 

Trish, Cobbler: -2¢ 22 15s cote an = ote epee ees 5 | New York (2); Maine; Ohio; Rhode 
Island. 

Crean i Moumtalt. <= eos ete see ante tos cade eee eee 5 | Maine (2); Massachusetts (2); Ver- 
mont. 

DOC BIE TIGR. Saino deine maaen oelasele © aeaiaiens caldera ee 3 | Maine (2); Michigan. 

WROTUTORS (oc oa. eens Done ans aetna ee ene eS oee 3 | Maine; New York; Vermont. 

Wihite Beaty o22 35. ase os coyotes inland as csap betas 2 | Maine; Minnesota. 

PYOLCHROP MASTCkK er #24 orca ee ce ee eee ee 2} Canada; Rhode Island. 

Lonig Seed linim. 32409 55) eee tk a eee 2 | New York. 

nick imniClivs ss. cada seer e ee. cee eee eee eee 2 | Pennsylvania; Vermont. 

RBLDEDOL. $n. Siew as oe anak es seene naa 0 oe ae tee 2 | Yermont. 

Sir Waiter Balen: oo 28 Joon. asso. se eerce et esaeeeee 2 | Minnesota; New York. 

Veéermiont Gold Coin J. 3-225 seswisicnin'seoccsama clues = 2 Pennsylvania; New York. 


The following varieties were mentioned once each, the commenda- 
tion coming from the locality mentioned in parentheses: American 
Wonder (Minnesota); Babbitt (Maine); Bonanza (New York); Boss 
(Vermont); Buffalo (Maine); Burbank (New York); Cambridge Russet 
(New York); Carmen No. 3 (Ohio); Clarke’s Pride (Maine); Clay Rose 
(Minnesota); Crines Lightning (New Jersey); Delaware (Minnesota); 
Enormous (New York); Gem of Aroostook (Rhode Island); Gloria 
(Rhode Island); Harris Snowball (New York); Holborn Abundance 


«The writer is indebted to the following experiment station officers for helpful 
advice: Professors Macoun, Canada; Woods, Maine; Rane, New Hampshire; Stuart, 
Vermont; Brooks, Massachusetts; Wheeler and Adams, Rhode Island; Fraser and 
Stewart, New York; Buckhout, Pennsylvania; Halsted, New Jersey; Selby, Ohio; 
Taft, Michigan; and Green, Minnesota. Much information has been secured from 
the replies of leading potato growers and seed dealers. It is regretted that it is 
impracticable to give detailed credit to these correspondents. 


RESISTANCE TO SCAB. 35 


Professor Kuehn (Rhode Island); Million Dollar (Michigan); Orange 
Blossom (Vermont); Rural New Yorker No. 2 (Vermont); Scabproof 
(Wisconsin); Squier (Vermont); Star of the East (Maine); State of 
Maine (Canada); Swiss Snowflake (Canada); Virgirosa (Vermont); 
Westfield (Vermont); White Scotch King (Minnesota). In addition 
three unnamed varieties received favorable mention from Vermont. 

From these reports it is observed that certain varieties like Dakota 
Red, Rustproof, and Keeper have very well proved powers of disease 
resistance, but lack other desirable characters to make them popular 
varieties. Several of the standard main-crop varieties are in some 
degree disease resistant, and doubtless owe their general popularity, in 
a measure, to this fact, although it has not been clearly defined. To 
this group belong Carmen’s best productions—Carmen No. 3, Sir 
Walter Raleigh, Rural New Yorker No. 2, and Rural Blush—Green 
Mountain, State of Maine, Delaware, Enormous, and White Beauty. 
Irish Cobbler is highly spoken of as disease resistant, but it is a 
question whether this may not be in part due to its early maturity, by 
virtue of which it escapes the worst ravages of Phytophthora. 

There is also a very promising series of new seedlings which should 
be carefully watched as to disease-resisting characters, among which 
may be especially mentioned Norcross, Star of the East, and Babbitt 
(Johnson Seed Company); Vermont Gold Coin (Burpee); Ionia Seed- 
ling (Dibble); Harris Snowball (Harris & Co.). 

It is encouraging to learn from the replies that several of the newer 
German and English varieties which are reputed disease resisting in 
their home countries have upon trial in America made a good show- 
ing. This is evidenced by the reports from the Rhode Island and 
Canada stations favorable to Professor Maercker, Gloria, Professor 
Kuehn (German), Holborn Abundance (English), and Swiss Snowflake. 


RESISTANCE TO SCAB. 


It is a matter of common observation that some varieties of potatoes 
are more liable to scab than are others. Reference has been made 
earlier in this publication to conclusions to this effect reached in Ger- 
many. So far as known the only American publication recording the 
results of comparative trials as to scab resistance is that made by the 
Vermont Agricultural Experiment Station in 1901-2.“ Thirteen varie- 
ties were tested in 1901 and fourteen in 1902 in soil badly infested 
with scab germs. While all showed some scab, there was a consider- 
able difference in the amount. Sir Walter Raleigh made a good show- 
ing both years, but an unnamed seedling sent by Mr. A. E. Manum, 
his No. 56, was more highly resistant. These trials established the 
writer’s confidence that still more resistant strains may be secured by 


4 Jones, L. R., and W. J. Morse. Vermont Exp. Sta. Report, XV: 225 (1902). 


36 DISEASE RESISTANCE OF POTATOES. 


breeding and selection. Prof. William Stuart, of the Vermont station, 
has undertaken this work in connection with the development of resist- 
ance to blight and rot.¢ He has made further trials during the last 
two years, but without conclusive results as yet. 

A request for information as to relative scab resistance was inserted 
in the circular of inquiry, already referred to, recently sent to Amer- 
ican potato specialists. Most of the replies to this question were 
negative in character, but a number gave interesting information, 
some of which is especially pertinent. The strongest evidence as to 
scab resistance comes from Mr. Hiram Presley, of Port Huron, Mich., 
a potato specialist, who has tested hundreds of varieties during the past 
thirty years. He commends the Cambridge Russet as practically 
exempt from scab. Mr. Frank Paddock, of Perry, N. Y., gives like 
evidence. 

Carmen No. 3 is highly spoken of as scab resisting by several grow- 
ers in Ohio, New York, and Vermont, but one Michigan correspondent 
condemns it. 

American Giant receives strong indorsement from Freehold, N. J., 
and vicinity; Salzer’s Scabproof from Wisconsin, and Aurora from 
Vermont. Favorable reports come from New York regarding Sir 
Walter Raleigh and Irish Cobbler, and from Canada regarding 
McIntyre. 

The following were each commended by one correspondent from the 
localities mentioned in parentheses: Best (Maine); Doe’s Pride (Maine); 
Early Freeman (Ontario); Keeper (New Hampshire); Seneca Beauty 
(Michigan); Squier (Vermont); White Beauty (Michigan); White 
Elephant (New Jersey); White Scotch King (Minnesota). 

On the other hand, the Early Ohio and some of its seedlings are 
condemned as especially liable to scab. Early Rose, Bliss Triumph, 
and Beauty of Hebron are also reported to scab badly. 

It is encouraging to note that in some cases the same variety is rated 
as ina high degree resistant to both diseases, the scab and the late-blight. 
and rot. A similar coincidence will be found if a comparison is made 
of the lists of German varieties showing resistance to the several 
diseases discussed earlier in these pages, and the same thing was 
observed in some degree with the English varieties. 

This indicates that the attributes which give power of resistance 
against one disease are not incompatible with those operative against 
another. Indeed, it is not unlikely that the general characteristics of 
disease resistance may prove to be similar or the same for these vari- 
ous maladies. If so, it will prove the easier to secure the model 
potato, which, while possessing that which is desirable in quality and 
productiveness, shall, in addition, show the highest degree of resist- 
ance to the various diseases. This is an ideal worth striving for. 


«Vermont Exp. Sta. Bul. 115, p. 139. 


SUMMARY. 37 


SUMMARY. 


The aim of this bulletin is to present in concise form what is known 

about disease resistance of potatoes. Much of this information is from 
European sources. . 
_ Certain minor diseases of obscure nature, but apparently nonpara- 
sitic, are first considered—the internal brown spot, filosité, and leaf- 
spot. Among remedial measures for each is the selection of resistant 
varieties. 

Scab diseases of tubers are in most, and perhaps in all, cases of para- 
sitic origin—fungous or bacterial. Apparently the variety of these is 
greater in Europe than in America, but the severity is less in Europe. 
It is undecided to what extent the American type of scab occurs in 
Europe, so a close comparison of conditions and remedies is not prac- 
ticable. In Germany certain varieties are known to be more scab 
resistant than others, among them being Richter’s Imperator, Profes- 
sor Wohltmann, and Irene. The same is true in America, Cambridge 
Russet leading the list, so far as is known. Other American varie- 
ties showing a considerable degree of resistance are Carmen No. 3, 
American Giant, Sir Walter Raleigh, and'Irish Cobbler. Scabproof 
and Aurora are also highly commended for scab resistance. 

Various stem diseases of the potato are known. The commonest 
type in Europe is termed blackleg (Schwarzbeinigkeit), a bacterial 
disease. It is not known to occur in America, but it resembles certain 
maladies which do occur here and which are as yet imperfectly under- 
stood. Varietal resistance to blackleg is not fully established, but 
apparently Dabersche and certain similar thick-skinned, starch-rich 
late varieties are more resistant than thin-skinned, starch-poor early 
varieties of the Rose type. Factor and Up-to-Date showed a consider- 
able degree of resistance to blackleg in England. La Czarine and 
other varieties are reported to show resistance to a bacterial stem 
disease in France. 

The late-blight and rot due to the fungus Phytophthora infestans 
occurs more commonly in Europe thanin America. Attention has been 
given for many years to relative varietal susceptibility to this disease, 
especially in Great Britain and in Germany. Varieties of superior 
disease resistance are known in both countries, and a number of the 
most promising from these and other European sources have been 
imported for trial. 

The following statements are tentatively formulated as to the nature 
of resistance toward blight and rot and the character of the varieties 
exhibiting it: 

(1) Disease resistance in potatoes is relative, not absolute, no variety 
known being wholly proof against late-blight and rot. 


38 DISEASE RESISTANCE OF POTATOES. 


(2) It seems related to general vegetative vigor, and is therefore in 
& measure dependent upon cultural and developmental conditions and 
tends to decrease with the age of the variety. 

(3) It can be restored by originating new varieties from seed, 
especially of hybrid origin. Not all seedlings show superior disease 
resistance. 

(4) The use of other species of tuber-bearing Solanums for hybrid- 
izing offers some promise, but no practical results have yet been 
secured. 

(5) Possibly the disease resistance in established varieties can be 
improved by selection, but this has not been proved. 

(6) Early varieties may escape the disease by maturing before it 
becomes epidemic, but when similarly exposed they are as a class less 
resistant than late varieties. 

(7) The source of seed tubers is a matter of importance, northern- 
grown seed giving plants of superior disease resistance in Europe. 
Seed from a crop that was not too highly fertilized is probably prefer- 
able. Possibly tubers are better for seed purposes if dug before they 
reach full maturity. . 

(8) High fertilization, especially with nitrogenous manures, lowers 
the power of the plant to resist both blight and rot. 

(9) Varieties relatively rich in starch are more resistant to rot; those 
richer in protein are more susceptible to it. 

(10) So far as skin characters are an index, the red varieties with 
thick and rough skin seem more resistant as a class than the thin- 
skinned white varieties. 

(11) So far as stem and foliage characters are concerned, the evidence 
favors the stem that is hard, rough, and rather woody at the base, and 
the leaf that is small, somewhat rough, and dark colored. 

The varieties rated highest as to disease resistance in England are 
Evergood, Discovery, Royal Kidney, Northern Star, Sir John Lle- 
welyn, King Edward VII, Eldorado, and Factor. 

In Germany and Holland the following represent the best types: 
Mohort, Irene, Geheimrat Thiel, Professor Wohltmann, Boneza, 
Eigenheimer, and Paul Kriiger. 

In Belgium and France no improvement as to disease resistance has 
been made over the best English and German types. 

In America, trials as to disease resistance have been conducted at 
some of the experiment stations, notably in Vermont, where experi- 
ments in breeding and selection for increased resistance are under way. 
These results have been correlated with information recently secured 
by a circular of inquiry addressed to a large number of potato special- 
ists in the Northeastern States and in Canada. From these it appears 
that a wide variation is shown in disease resistance among the varieties 
now in cultivation in America, but that no one variety is preeminent. 


SUMMARY. 39 


Among those which have been widely tested, the following deserve 
mention as of the resistant class: Dakota Red, Rustproof, Irish Cob- 
bler, Sir Walter Raleigh, Doe’s Pride, and White Beauty. Certain 
European varieties of the disease-resistant type seem to retain that 
character when grown in this country, e. g., Professor Maercker and 
Sutton’s Discovery. There is much of promise in certain new varie- 
ties under trial at the Vermont station. Several new sorts of reputed 
disease resistance have recently been placed on the market by American 
seedsmen, e. g., Harris’s Snowball, Dibble’s Ionia Seedling, Burpee’s 
Vermont Gold Coin, and Johnson’s Norcross, Star of the East, and Bab- 
bitt. Those having opportunity should carefully observe the relative 
disease resistance of these and also of other new varieties. 

The evidence at hand seems to justify the hope that the coordinated 
efforts of potato specialists working from both the practical and the 
scientific standpoints may soon result in the development of varieties 
of potatoes combining general excellence with a high degree of disease 
resistance. All who can do so are urged to aid toward the accomplish- 
ment of this end. 


O 


o* Speer 


PLATE I. 


Bureau of Plant Industry, U. S. Dept. of Agriculture. 


8 


‘NOLLOD DNILSISSY-TIASSM HLIM SLNAWIYSdxX9 JO 3N3S0G SHL WWIVWALYNYD ‘'ZVd VYSA VLITV ‘WINDNVOAS LY AATIVA 


U. S. DEPARTMENT OF AGRICULTURE. 
BUREAU OF PLANT INDUSTRY—BULLETIN NO. 88. 


B. T. GALLOWAY, Chief of Bureau. 


WEEVIL-RESISTING ADAPTATIONS 
OF THE COTTON PLANT. 


BY 


Or COOK. 
BIONOMIST IN CHARGE OF INVESTIGATIONS IN THE AGRICULTURAL 
Economy oF TROPICAL AND SUBTROPICAL PLANTS. 


IssuED JANUARY 13, 1906. 


SS alts AS 
WZ MUTATOR 
ATT 3S 
Sine <a> 


WASHINGTON: 
GOVERNMENT PRINTING OFFICE. 
L906. 


BUREAU OF PLANT INDUSTRY. 


B. T. GALLOWAY, 
Pathologist and Physiologist, and Chief of Bureau. 

VEGETABLE PATHOLOGICAL AND PHYSIOLOGICAL INVESTIGATIONS. 
ALBERT F. Woops, Pathologist and Physiologist in Charge, Acting Chief of Bureau in Absence of Chief. 
BOTANICAL INVESTIGATIONS. 

FREDERICK Y. COVILLE, Botanist in Charge. 

FARM MANAGEMENT. 

W. J. SPILLMAN, Agriculturist in Charge. 

POMOLOGICAL INVESTIGATIONS. 

G. B. BRACKETT, Pomologist in Charge. 

SEED AND PLANT INTRODUCTION AND DISTRIBUTION. 

A. J. PIETERS, Botanist in Charge. 

ARLINGTON EXPERIMENTAL FARM. 

L. C. CORBETT, Horticulturist in Charge. 


INVESTIGATIONS IN THE AGRICULTURAL ECONOMY OF TROPICAL AND SUBTROPICAL 
PLANTS. 


O. F. Cook, Bionomist in Charge. 
DRUG AND POISONOUS PLANT INVESTIGATIONS, AND TEA CULTURE INVESTIGATIONS, 
RODNEY H. TRUE, Physiologist in Charge. 
DRY LANP AGRICULTURE AND WESTERN AGRICULTURAL EXTENSION, 

CARL S. SCOFIELD, Agriculturist in Charge. 

EXPERIMENTAL GARDENS AND GROUNDS. 

E. M. BYRNEs, Superintendent. ; 

SEED LABORATORY. 


EDGAR Brown, Botanist in Charge. 


J. E. ROCKWELL, Editor. 


_ JAMES E. JONES, Chief Clerk. 


INVESTIGATIONS IN THE AGRICULTURAL ECONOMY OF TROPICAL AND SUBTROPICAL 
PLANTS. 


SCIENTIFIC STAFF. 
O. F. Cook, Bionomist in Charge. 
G. N. CoLuins, Assistant Botanist. 
F. L. Lewton, Scientific Assistant. 


H. PITTIER, Special Agent. 


.¥ 


LETTER OF TRANSMITTAL. 


U. S. DerarrMent oF AGRICULTURE, 
Bureau or Piant Inpustry, 
OFFICE OF THE CHIEF, 
Washington, D. C., September 26. 1908. 

Sir: I have the honor to transmit herewith a report on “ Weevil- 
Resisting Adaptations of the Cotton Plant,” and to recommend it 
for publication as Bulletin No. 88 of this Bureau. This report has 
been prepared by Mr. O. F. Cook, bionomist in charge of investiga- 
tions in the agricultural economy of tropical and subtropical plants. 
Tt contains an account of his observations and experiments which 
show that some of the varieties of the cotton plant have definite 
weevil-resisting characters. The establishment of these facts opens 
new and unexpected lines of approach to cultural solutions of the 
weevil problem. 

The investigation of cotton referred to in this report was begun 
in March, 1904, through the Laboratory of Plant Breeding, there 
having been set aside for it from the emergency cotton boll weevil 
appropriation a part of the funds which had been devoted to the 
breeding of weevil-resistant cotton. The existence of a field culture 
of cotton in the presence of the boll weevil had been ascertained by 
Mr. Cook during a visit to Guatemala in 1902, and it was hoped that 
‘the immunity of the cotton might prove to be due to some weevil- 
resistant quality. 

The first result of detailed observations was the discovery of the 
weevil-eating kelep or so-called Guatemalan ant, which has been 
made the subject of previous reports through the Bureau of Ento- 
mology. It now appears that the usefulness of this insect is not 
limited to the boll weevils which it catches and kills. By making a 
regular field culture of cotton possible in the presence of the boll 
weevil it has contributed in an important manner to the development 
of the weevil-resisting characters here described. The cotton plant, 
it seems, has been greatly modified in protecting itself against the 
ravages of its insect enemy. Not only has it attracted the kelep to 
its service and developed other means of defense which are more 


3 


4 LETTER OF TRANSMITTAL. 


direct, but even the lint, on the peculiar character of which the com- 
mercial value of the crop depends, appears to find its chief use to the 
plant in excluding the weevil larvee from the seed. Our Sea Island 
and Upland varieties have been raised for long periods in regions 
where the boll weevil did not exist and, as was to have been expected. 
are largely lacking in protective features. The Kekchi cotton, on the 
other hand, which has continued its development in a weevil-infested 
region under the protection of the keleps, has by far the largest 
number of weevil-resisting characters. 

The fact that weevil-resisting adaptations really exist, as shown in 
numerous instances in the present report, emphasizes the necessity of 
a thorough study of our cultivated cottons for the purpose of taking 
advantage of any and all protective characters. 

It is possible, as Mr. Cook suggests, that the Guatemalan variety 
of cotton which he has discovered, and which has such a surprising 
number of weevil-resisting adaptations, may not prove suited to culti- 
vation in the United States, but even in that case the value of the 
present paper on weevil-resisting characters would not be diminished, 
for it will serve as a help to all who may engage in seeking and 
developing such characters in the types of cotton now cultivated in 
our country. * 

Respectfully, B. T. Gattoway, 
Chief of Bureau. 


Hon. JAmMes Witson, 
Secretary of Agriculture. 


taprodnehlon= 2!) 2-2 ..... Se ae oe RS IAS od oe ON Ie SS, eee ee eee 
Paueantnence Of the boll weevil s.2.- -.2.-2.5..--.----2+2---------.-- 
Seemet MemaneciiVe-CNuraAcvors;. \- <2. 22 32. 2-- 2 . eae ee eae 
Dwarf habit and determinate growth of Kekchi cotton__________-___-- 
Mart anionsyin wheswekchiGOuOn 2-0. ees ee tk 

Effects of Guatemalan conditions on United States varieties_ .____- 
Acclimatization of Kekchi cotton in the United States _____. _____- 

Early bearing facilitated by long basal branches ----_------------------ 
Hari rejection of superfluous squares -:-....----.--.----------------- 
Seasonal bearing of perennial varieties ______-__.----------- fis Te eh ee: 
Annual cutting back of perennial varieties _-____------------- it eee Ye 

Pe avai Us ane Cet SbOIMN Se -- ah. - SE en EL See 2 St ee ee ee 
TEGO BEATE (DO DIS ics 92 Sree Gah deh an SS 2 Ra a pe Anna a LONE es 
Extrafioral nectaries _-_-__- Che ages a RE a as a a 
Le RCPOTEN BS: UE (ELUTE IGE ING ae a a a eo 
Pmaemamnectaries of the involueré. 22. 2-22 22 25-222 tee ene - 
renee UTIs Ol HO INVOMUGCTOl -—. = 52 2a es ok ieee we 
Nectaries of Guatemalan Sea Island cotton _____---_____---------- 
Mmmmimnrecdisecrenion Of NeClar 2249-=*=, .- 22s 8 228 ee se ee 
Peachicpisubienaine immer nectaries: 3.52. o.-> os ee ee 
Biteinicy Ol tie Kelep provectOmn.. 25. Si 9 Soe 2 eo elle k ee 
Other nectar-bearing plants visited by the keleps___-__-_-._-____-- 
Mheunvolucre as a protective structure ...-.+.....-.-<--_...--.-..-=.--.- 
Kamlocralpracks STown together 2.3.0 2.5-- =. =--.4- 52222 sence t 222 
PERO RCEACOMNALCINIS OF DTACUS., 2 22. 0 2a= eae eosin 5 ace teas rege 
MacesMnvOluGresiOL WekChi COtbON 2 222-525. 225-2 F208 oie ee ee 
Gannon oriarins. Of bracts avoided... 25) 2.2 S25 1 2. ie ee 
Eenininarcins OL imvolucral bracts. . =: .....=-=-2 222-22. -<- 5.2.2 E 
Panominoa protection by-involucre: .....22..2222--4.- += 262 Sseee--24- 
Pein Oe OiOPenviabvVOlUCLeS .<<5.. ooo 25- foe oe ee epee nek ee 
SeEmeEeMbLe TRAratitized WUdS —.-_._.-. .._-__--_--.-.--4---~----es-2--- + 
paccdm= of weevil-infested squares _____...-..-.-.-----.-------------- 
Cognmnes of flared and fallen squares ......-.=-.....-.--..----------- 
Eroierraion. oO: internal tissues of buds..2.22-.2.=22.--5. 2-24 ..2.--2-- 
Causes and conditions of bud proliferation ______......---------------- 
Proliferation in other varieties ____--_- spat) 2 Nae ee ee Gages Oe 
NNEC RCERNE DON es Ts ee as tet. aclo mae See de 


mummy or very young bolls-. +. .22-2. 22.¢.le22 4: 22-2. 2 - rhe 


i wuberdmooL-young bolls’: . 02.20.25 lees 32. 2 -ee. e222 
LEE PE LDS) 0 DS NR oer? AS ee 


~ OT 


io-3) 


occ caose 


aor awo-m 


or 


6 


CONTENTS. 


Protection of the bolls—Continued. Page. 
Proliferation from the wall of the boll._..-.------ code cheec. Soe 58 
Time required for proliferation. .....-..-=:--..02--2 --2s.sesene eee 60 
Efficiency of adaptive characters of bolls... ..:.:.._...-_2_. Soe 61 
Bacterial diseases following weevil injuries_..........-.---.--.---.--- 62 
Breeding in buds'a ‘derived habit ...:..2..)_/: 22-232 2 eee 62 
Relation between proliferation in buds and in bolls ______--__--.-_.--- 64 

Protection of seeds by lint._.. ____ _- aan + ears oon 65 
Protective seed arrangement in Kidney cotton: ace cece denne 66 

Cultural value of Kidney cotton. 2-2... .22___ 2-4. 2 67 

The nature and causes of adaptations. __-- -22......-2_ 2-22.22 2e 67 

Conscious and unconscious’ selection.--_-.. . 2-22-3222 ee 70 

Summary of adaptations -_.._-_....-.-..-.25.-2-22-225-.08=s= a 72 
Classification of adaptations -----....-..25..-21-2-2- Se 72 
Adaptive characters of different types of cotton_............-.-.------ 73 

Concluding remarks . --...-22....-.22---22 212.4... 22 74 

Description of plates _-.--*-+--.-= - 2% ‘ol steer Seer = 78 

Index - 242.5 5232022022008 2251 ea Se 79 

ILLUSTRATIONS 

PuateE I. Valley at Secanquim, Alta Vera Paz, Guatemala, the scene of 
experiments with weevil-resisting cotton_-_-.__..--__-- Frontispiece. 

Il. Fig. 1.—Mature plant of Kekchi cotton. Fig. *.—Kekehi cotton 
plant with polls -__.:.. ..2 22: [22 2 ee ee 78 
III. Involucres of Kekchi cotton, showing nectaries and bractlets___- 78 

IV. Fig. 1.—Involucres of Rabinal cotton, showing connate and ap- 
pressed margins. Fig. 2.—Open involucres of Egyptian cotton - 78 

V. Fig. 1.—Young buds of Kekchi cotton with weevil punctures. 
Fig. 2.—Buds of Kekchi cotton with proliferation. -_.--.-.-.-- 78 
VI. Large buds of Kekchi cotton with proliferation -.______-...----- 78 
VII. Weevil-infested bolls of Kekchi cotton -_.........-------------- 78 
VIII. Carpels of Kekchi cotton, showing proliferation -__._........--- 78 

IX. Fig. 1.—Kekchi cotton, successive stages of the boll. Fig, 2.— 
Kekchi cotton bolls (right) compared with King bolls (left) - -- 78 

X. Fig. 1.—Rabinal cotton with bolls. Fig. 2.—Bolls and seeds of 
Kadiiey Conon, - 252225 eese Pe 78 


B. P. I.—180. 


WEEVIL-RESISTING ADAPTATIONS OF THE 
COTTON PLANT. 


INTRODUCTION. 


The fact that Central American varieties of cotton have developed 
weevil-resisting adaptations has already received preliminary notice.’ 
A third visit to Guatemala, in the spring of 1905, has given opportu- 
nity for further studies of the protective characters of the native 
varieties and for comparing them with the types of cotton now cul- 
tivated in the United States. For this purpose plantings of Upland 
and Sea Island varieties have been made in Guatemala, and as the 
season advanced other tests of the Guatemalan and United States 
varieties were arranged under very different climatic conditions in 
Texas and at Washington. 

These opportunities of comparative observation have revealed a 
series of protective adaptations of such number and nicety as to fur- 
nish a unique and well-nigh incredible instance of selective develop- 
ment. The statement of the former paper may be repeated with 
emphasis, that the presence of the weevil-eating kelep has enabled 
the Indians of eastern Guatemala to maintain since very ancient 
times field culture of cotton in the presence of the weevils, with the 
result that there has been developed a dwarf, annual, short-season 
variety with numerous features which, in the absence of sufficient 
numbers of keleps, afford material assistance in protecting the crop 
against the ravages of the weevil. 

Whether this Guatemalan cotton can be made of direct use in the 
United States or not, it demonstrates the existence in the cotton 
plant of weevil-resisting characters. The new variety has lint of 
good length and quality, so that its utilization in the United States 
depends upon its adaptability to our climate and methods of culture. 

As already explained in publications devoted to the kelep, the 
weevil-eating propensities of that insect were discovered in 1904 
during a visit to Guatemala which had been undertaken in the hope 
of finding a weevil-resisting variety of cotton. It had been observed 


2 Cotton Culture in Guatemala. Yearbook of the United States Department of 
Agriculture for 1904, 475-488: Science, N. S., 20: 666-670, November 18, 1904. 


‘ 


8 WEEVIL-RESISTING ADAPTATIONS OF COTTON. 


two years before that a field of dwarf cotton cultivated by the Indians 
did not suffer from the boll weevils, though these pests were abundant 
on a “tree cotton” a short distance away. 

The kelep afforded an entirely unexpected and yet very striking 
explanation of the fact that cotton was being grown as a regular 
field crop in a region which had probably been infested with weevils 
for many centuries, if it were not, indeed, the original home of the 
species. That there was an insect in existence specially qualified 
by structure and habits to attack, disable, and devour the boll weevil, 
was welcome news in the United States, and in accordance with cabled 
instructions from the Secretary of Agriculture numerous colonies 
of the keleps were brought home and colonized in the cotton fields 
of Texas. 

The finding of the kelep explained the failure of the weevils to 
prevent cotton cultivations in eastern Guatemala, and seemed at first 
to diminish the prospects of weevil resistance in the cotton itself. 
Nevertheless, the intention of studying Guatemalan varieties of cot- 
ton and the cultural methods in use in that country was not aban- 
doned, and the results are not without bearing on the original ques- 
tion of the causes of the apparent immunity of the Guatemalan 
cottons, and also upon the more practical question of securing cotton 
varieties and cultural methods by which the injuries of the boll 
weevil in the United States may be reduced to a minimum. 

The Guatemalan cotton protected by the keleps is a genuine Up- 
land variety, very early and productive, with a fiber of good length 
and texture, as already stated. In addition to features which di- 
rectly favor the keleps, it has many other qualities which may 
render it useful, even without its insect guardians. In former reports 
it has been compared with the very early Upland varieties, such as 
King and Parker; but comparative tests made in eastern Guatemala 
show that the native variety, which it is proposed to call Aekehd, 
vepresents a very distinct type of this important cultivated plant. 
It belongs to Gossypium hirsutum, the Upland species or series of 
varieties, in the sense that it is not a Sea Island, Egyptian, or Kidney 
cotton,” but it is distinctly more different from any of the Upland 
varieties now cultivated in the United States than these are from each 
other. It has not been ascertained that the Kekchi cotton in its 


aThe Sea Island cotton is so called because cotton of this type is cultivated 
en the Sea Islands of South Carolina, long famous for the excellence of their 
product. The Sea Island cotton came originally from Barbados, whence also 
its botanical name, Gossypium barbadense. 

Upland cotton gained its name as a means of distinguishing it from the Sea 
Island, being cultivated in the interior, or “ upland,” districts of the Southern” 
States. The Upland type of cotton was recognized as a distinct species by 
Linnseus under the name Gossypium hirsutum, but many subsequent writers 


INTRODUCTION. 9 


present form is suited to cultivation in the United States, but it has, 
without any doubt, new and significant characters which must be 
regarded as factors in cultural solutions of the weevil problem. (PI. 
II, fig. 1.) 

Although cotton was not found to be planted as a regular field cul- 
ture in any localities in Guatemala where the keleps do not exist, 
small quantities are produced in the interior plateau region about 
Rabinal by what may be called dooryard cultivation, and these, too, 
have suggested cultural factors and expedients which may not be 
without practical bearing. 

The present paper can claim to make only a beginning in the 
bionomic study of the question, but it shows at least that the weevil 
problem has many avenues of approach on the botanical side. 

The cotton of Guatemala and neighboring countries has maintained 
an existence, at least, in the presence of the weevils, and has suffered 
an acute natural selection with reference to its ability to protect itself 
against the weevil or to secure the assistance of allies, such as the 
keleps. That no commercial cotton crop is raised or exported from 
such districts does not prove that they are unworthy of scientific 
investigation, or that they are not likely to yield materials and sug- 
gestions of practical value in meeting the invasion of weevils which 
is Now so serious a menace to the cotton industry of the United States. 

Some of these weeyvil-resisting adaptations have been of use in 
securing for the cotton the assistance of the keleps. There are others 
which, if properly utilized, might render these interesting insects 
unnecessary. Tropical America has been serving for thousands of 
years, evidently, as a laboratory for this class of experiments. Texas 
was invaded only yesterday—a decade ago. Now that we are forced 
to engage in the strife, the first preliminary should be, it would seem, 
to take stock of the weapons which nature has forged. 

The present report was planned and partly written before the dis- 
covery of the true nature of the best of the weevil-resisting adapta- 
tions—the proliferation of the tissues of the buds and bolls. Some 
of the characters here described may have no value except as sug- 
gestions, but taken together they may be of interest as an outline of 
the results of the very long period of selection to which the presence 
of the boll weevil has subjected the Central American varieties of 
the cotton plant. 


liave erroneously confused it with the Old World species Gossypium herbaceum, 
which is not cultivated in the United States, though often so reported. 

The Egyptian and Kidney cottons belong to the Sea Island series, and are of 
American origin. The Kidney cottons seem not to have been cultivated on a 
commercial scale, but they are very widely distributed in tropical America. The 
name refers to the fact that the seeds ef each compartment of the boll are 
grown together into a small compact mass, in shape suggesting a kidney. 


10 WEEVIL-RESISTING ADAPTATIONS OF COTTON. 


SELECTIVE INFLUENCE OF THE BOLL WEEVIL. 


The boll weevil exerts a most prejudicial effect upon the cotton 
crop, but, unlike most parasites, it does not cause disease or debility 
in its host plant. The young buds and bolls are merely pruned away, 
as it were, the purposes of the weevil being the better served when the 
plants remain vigorous and continue to produce more buds and bolls, 
in which more eggs can be laid and more larve brought to maturity. 
Nevertheless, if no bolls are allowed to develop no seed can be set. 
The fate of the cotton crop in wet seasons in Texas shows that with- 
out some form of protection the plant would have been extinct long 
since in all localities reached by the boll weevil. 

The long contact between the boll weevil and the cotton plant in 
Central America has given ample opportunity for the latter to profit 
by the selection which the insect itself has provided. Every differ- 
ence by which a cotton plant was able to resist or to avoid the weevil 
and thus ripen more seeds than its fellows would give it a distinet 
advantage, quite as if the selection were consciously carried on by the 
planter or the plant breeder. The case is different from that of the 
recent improyements of many of our cultivated plants by selection 
for the increase of some particular quality already existing. Such 
improvements can often be made appreciable, or even highly valu- 
able, in comparatively few years, but under the desultory Indian 
methods of cultivation long periods of time would be required for 
the origination and accumulation of such characters as these pro- 
tective adaptations. 

Climate and other local conditions must also be taken into consid- 
eration. An adaptation which would be effective in one set of cli- 
matic conditions may be of little use, or even a positive disadvantage, 
in others, as, for example, the prompt shedding of the parasitized 
buds. In a dry region the falling of a bud to the superheated, 
sun-baked earth insures the death of the weevil larva, either by the 
heat directly or by the complete drying out of the tissues in which 
the larva is embedded. In the moist districts of eastern Texas, how- 
ever, this expedient is quite ineffective, the larvee often developing 
even better when the buds fall off and he on moist soil than when they 
‘remain attached to the plant. 

It need not surprise us to learn also that the weevil-resisting adap- 
tations shown by the Kekchi and other cotton varieties of Central 
America are shared, to some extent, by those already known in the 
United States, since the whole Upland type of cotton appears to have 
been, originally, a native of the Central American region. Varieties 
which reached the United States from Mexico and the West Indies 
may, however, have had little or no contact with the weevil for many 
centuries, while in Central America the struggle for existence has 
remained severe and continuous down to the present day. 


GENERAL PROTECTIVE CHARACTERS. 11 


It is now known that in the plateau region of Mexico the long dry 
season effectually excludes the weevil, so that varieties of cotton from 
the Mexican highlands, instead of being weevil-proof, as sometimes 
represented, may have no immunity whatever when brought into 
the much more moist climate of the cotton belt of the United States. 

The Kekchi cotton of Guatemala, on the other hand, has to a much 
greater degree than any of the varieties now grown in the United 
States the very qualities which experiment has shown to be effective 
for the mitigation by cultural means of the injuries inflicted by the 
boll weevil. That it has, in addition, other features not possessed 
by our United States varieties, or not hitherto interpreted as weevil- 
resisting adaptations, need not be looked upon as anything outside 
the normal order of nature, but is entirely in accord with what 
appears to be the biological and agricultural history of the cotton 
plant in Central America. 


GENERAL PROTECTIVE CHARACTERS. 


DWARF HABIT AND DETERMINATE GROWTH OF KEKCHI COTTON. 


Although Guatemala is a tropical country and the climatic condi- 
tions are suitable for the growth of cotton throughout the year, the 
Kekchi cotton is cultivated only as an annual, and is smaller and 
more determinate in its habits of growth than the Upland varieties 
now known in the United States. It soon attains its full height, 
and after a crop of bolls has set on the lower branches there is a 
definite tendency to cease growing or producing new buds. The 
later upward growth of the plants seems to be supplementary, as it 
were, to the formation of the bolls; often there appear to be no 
more flowers formed, and many of those which come seem to be 
undersized, as though the plant were really mature and were 
«pproaching the natural termination of its existence. Our Upland 
varieties, on the contrary, continue to produce throughout the season 
hundreds of small squares on each plant which serve only as breed- 
ing places for the weevils. 

The explanation of the high development of these short-season 
qualities of the Kekchi cotton is doubtless to be found in the custom 
of the Indians, who pull up the cotton as soon as the bulk of the crop 
has ripened to make room for the peppers, which are always planted 
with the cotton. For the Indians the peppers are an even more 
important crop than the cotton, so that when the time comes for 
clearing away the cotton they do not wait for the plants which may 
have delayed maturity. Late bolls, even, would never come to 
maturity or furnish seed for planting. The result has been a very 
long-sustained selection for early bearing and uniform ripening of 
the crop. Some of our earliest Upland sorts may begin blossoming 


1g WEEVIL-RESISTING ADAPTATIONS OF COTTON. 


as soon as the Kekchi, but they show far less tendency to determinate 
growth. 

The development of earliness has been assisted, no doubt, by the 
climatic conditions which prevail in eastern Guatemala. The rainy 
season oftens begins before the cotton harvest is completed, so that 
the later bolls are very likely to become diseased, or, if they reach 
maturity and open, the lint is often beaten to the ground and made too 
dirty for use in spinning and weaving. In either case the seed is not 
harvested. 

The Indians believe that even if they did not pull the cotton up it 
would not become a perennial, but would die out completely, even to 
the roots, during the rainy season. Seeds scattered accidentally in 
the plantation at harvest time are rotted by the rain and do not germi- 
nate, so that little or no volunteer cotton is carried over from one 
season to another. : 

If the Kekchi cotton were the only variety planted in Guatemala 
and the weevil had there, as in the United States, no other food plant 
than the cotton, the insects might all die off between April or May, 
when the cotton is pulled up, and October, when the next crop is 
olanted. There is, however, enough perennial “ tree” cotton in the 
country to keep the pest from becoming exterminated. Moreover, 
the question of additional food plants in Guatemala is still open. 

The importance of securing short-season varieties of cotton for the 
United States can hardly be overestimated, since, as already intimated 
elsewhere,“ there is no longer any reason to hope that the more severe 
winters of the northern districts of the cotton belt will give any pro- 
tection against the weevils. 

As long as the weevil was confined to the southern part of Texas, 
where the cotton could survive the winter, the destruction of the 
plants as soon as possible after the maturing of the crop was the only 
measure calculated to seriously reduce the number of weevils. It 
was also essential to plant cotton as early as possible in the spring to 
avoid the weevils bred on the volunteer, or hold-over, cotton which 
negligent planters had left in the ground. The extension of the pest 
farther north and the possibility of securing cotton varieties with 
determinate habits of growth introduce several new considerations. 
The hold-over cotton is eliminated from the problem, but in the more 
northern latitudes, where the cold comes earlier and the temperature 
remains lower throughout the winter, it may often happen that there 
will be no period in which the weevils can be reduced by starvation, 
unless time can be secured for this purpose in the spring by the plant- 
ing of short-season varieties of cotton. 


«Cook, O. F., 1905. Progress in tue Stuuy of the Kelep, Science, N. 8., 21: 


052. 


PROTECTIVE CHARACTERS OF KEKCHI COTTON. 13 
? 

Instead of colder winters being unfavorable to the weevils, there is 
every probability that cold sufficient to keep them in a torpid, inactive 
condition will preserve their noxious lives much better than warm and 
pleasant weather, which enables them to continue active and thus 
deplete their vital energies. The winter of 1904-5 was one of un- 
precedented severity in Texas, both in absolute temperature and in 
continued cold and wet, and yet the weevils were able, in many locali- 
ties, to infest heavily the early plantings of cotton to a far greater 
extent than in previous years. 

The farther north the locality the more will the efficiency of cul- 
tural methods of avoiding the boll weevil depend upon the plant- 
ing of quick-maturing varieties of cotton. It is true that in a 
favorable season the cotton planted first would set its crop soonest, 
and thus escape a part of the damage suffered by adjoining fields 
of later growth, the earlier fields breeding weevils to attack in 
larger force the later plantings. But instead of insuring a decrease 
of the number of weevils in a given locality and checking the 
propagation of the pest, very early planting by a part of the farmers 
of a community might tend, after an early fall and a cold winter, 
to the opposite result, since it would save the lives of large numbers 
of weevils which would otherwise perish before the cotton, if sown 
a few weeks later, would be large enough to furnish the weevils 
with food. Dr. Herbert J. Webber states that planting ‘could 
probably be deferred even to the middle of June without impair- 
ing the chances of a crop as large as that which can be obtained 
in the presence of the weevil. 

There would seem to be little object in planting cotton where 
the weevils are as abundant as in some places in southern Texas in 
the spring of the present year, 1905. Nevertheless, the opportune 
occurrence of a few weeks of dry weather was able, even then, to 
greatly improve the prospects of @ crop. No matter how bad the 
weevils, the planter still has hope that dry weather may come and save 
his crop from being a total loss. .As long as indeterminate varie- 
ties are planted this possibility will always make it difficult to carry 
out a general policy of early destruction of the plants. 

Some of our Upland varieties of cotton are early enough in the 
sense that they begin flowering and fruiting very promptly, but 
unless the season is very dry they will produce a continuous succession 
of buds until they are pulled up or frost cuts them off. The earli- 
ness of practical value is not to be shown merely by the date of 
flowering, but by the date of ripening the crop of bolls and of 
ceasing to form new buds in which weevils can breed. If the im- 
provements noted in other parts of this report can be realized in 
practice, it would no longer be necessary to destroy the cotton plants 


14 WEEVIL-RESISTING ADAPTATIONS OF COTTON. 


in order to’ put an end to the breeding of the weevils. It would 
then become practicable and desirable to regulate planting so as to 
bring the growing period of the cotton at the most favorable season 
for a rapid development of the crop, and thus to give ‘the weevils 
the shortest possible opportunities for breeding. If the fall and 
winter had favored the survival of many weevils, planting could well 
be deferred until the weevils had disappeared, a fact which could be 
ascertained by starting early a few observation plants from which 
the weevils could be carefully picked by hand as long as they con- 
tinued to appear. 

The extent of the mortality of the boll weevil in the spring has been 
well shown in the investigations reported by Mr. W. D. Hunter on 
the effects of applying Paris green to the very young cotton as a 
means of destroying the weevils which had lived through the winter. 
Numerous dead weevils were found in the poisoned fields, but equal 
or even greater numbers were found in those to which no Paris green 
had been apphed, and the conclusion was drawn that a large propor- 
tion of the weevils, which pass the winter in a state of hibernation 
or torpidity induced by the cold, perish through starvation or other 
causes in the spring, after the weather has become warm enough to 
render them active again and permit them to renew their search for 
cotton plants on which to feed and lay their eggs.” : 

It is easy to understand, too, that after the weevils have been re- 
duced by the cold to a condition of inactivity involving an almost com- 
plete suspension of the vital functions, the lack of food and the lapse 
of time can make very little difference with them. Starvation comes 
much quicker during warm weather while they are going actively 
about, so that it is the autumn and spring which must be relied upon 
to reduce the numbers of the weevils rather than the cold periods of 
the winter months. Messrs. Hunter and Hinds have also noted as 
significant the fact that of weevils captured at the middle of Decem- 
ber, 15.8 per cent passed the winter successfully, while of another lot 
captured a month earlier, only 1 per cent survived. Their conclu- 
sions were as follows: 

It is evident that the weevils which pass the winter and attack the crop of 
the following season are among those developed latest in the fall and which, in 
consequence of that fact, have not exhausted their vitality by oviposition or any 
considerable length of active life. 

With these facts in mind it becomes plain that no objections need 
be raised on general biological principles to the introduction of new 


aA determinate vantety of cotton would also avoid the “cuiltwel disadvan- 
tages incidental to very early planting, for if the weather happens to turn 
cold and wet the cotton is often either killed outright and has to be replanted 
or, what is still worse, it becomes permanently stunted and unproductive. 

b Hunter, W. D., 1904. The Use of fies Green in Controlling the Cotton Boll 
Weevil, Farmers’ Bulletin No. 211, U. S. Department of Agriculture. 


VARIATIONS IN KEKCHI COTTON. — £5 


. 


quick-maturing varieties of cotton from tropical countries on the 
ground that cold weather will exclude them from the United States. 
The early spring is the only time in which they will be likely to 
encounter adverse conditions in this respect, and if varieties can be 
secured which are able to mature a satisfactory crop in a short season, 
these quick-maturing qualities will far more than compensate for any 
lack of ability to withstand cold weather in the early spring. 

The Kekchi cotton may prove, however, to be quite as tolerant of 
cold as the other Upland varieties now cultivated in the United 
States.* In its native country it is planted in October and grows 
throughout the winter months in mountain valleys where tempera- 
tures of between 40° and 60° F. are not infrequent. (Pl. L.) 


VARIATIONS IN THE KEKCHI COTTON. 


Very great diversity of size, habit of growth, and other features 
exists in the Indian cotton of the vicinity of Secanquim and Cajabon. 
The plants cultivated by Mr. John H. Kinsler on the United States 
system were also very different from any grown by the Indians, 
being much more robust and compact than in the more crowded 
native fields. The spreading lateral branches and low, compact 
growth of the Kekchi cotton, as shown in Plate IT, figure 1, might have 
cultural disadvantages if these tendencies were to be maintained in 
regular field cultures. Such, however, is not likely to be the case. 
When growing closer together the plants are more upright and less 
leafy below. 

To what extent the differences observed thus far represent varietal 
characters can scarcely be determined without a field test of the 
apparently different strains, side by side. The broken, precipitous 
nature of the country renders it impossible to rely upon comparisons 
of the conditions of the different fields. 

The conservative agricultural habits of the Indians would tend to 
the continued planting by one man or family of the same seed for 
long periods of years, which might well conduce to the formation of 
separate strains. The low germinating power of the seed may pos- 
sibly be due to such inbreeding, though it is more likely that it deteri- 
orates because of the humidity of the climate.’ Nevertheless, our 
experiments were sufficient to prove that even among plants grown 
from seed raised by the same Indian there were very appreciable 


«This was shown to be a fact before the report was printed. See p. 18. 

» The Indians appreciate the fact that the cotton seed does not germinate well. 
They are accustomed to plant six seeds together, from which two or three plants 
usually reach maturity, often with one or two insignificant dwarfs underneath. 
The yield per plant in these crowded fields is naturally very small, but the larger 
individuals often bear from 20 to 30 bolls. At Rabinal from 6 to 10 plants in a 
cluster is the rule, the product of the individual being still further reduced. 


16 WEEVIL-RESISTING ADAPTATIONS OF COTTON. 


(lifferences, sufficient to have a very practical bearing upon the ques- 
tion of securing strains having the special characters required in the 
United States. Indeed, there was nearly as much diversity among 
the Guatemalan plants as among all the Upland varieties, though 
these were in some cases unusually variable, as a result apparently 
of the transfer to new and unwonted conditions of climate and soil. 

The usual number of locks or cells in a boll of the Kekchi cotton is 
four, but bolls containing three or five are not uncommon; often they 
are on plants which have otherwise the usual number. 

There is also considerable diversity on the same plant in the shape 
of the bolls, some, for example, remaining quite conical and pointed, 
while others round-out to near the apex. One plant was observed 
in which the bolls were very nearly spherical. The involucre was 
also unusually large. The plant had an unusually deep red or black- 
ish color, and was distinctly more vigorous than its neighbors, as 
often happens with mutations. 

It is not at all probable that a close selection has ever been prac- 
ticed by the Indians, so that a wide diversity of mutational charac- 
ters may be expected when once the variety has been brought under 
careful observation. 

The stems and petioles of the Kekchi cotton plant are dark red, 
or at least spotted with red, and the leaves turn dull red with matu- 
rity. The bracts and bolls are green when young, but with age and 
exposure to the sun become more or less tinged or spotted with red. 
The outer involucral nectaries also turn deep red, especially the two 
upper ones, even while the buds are still very young. The great 
majority of the leaves are simply three-pointed, but many of them 
have an additional smaller lateral point on each side near the base. 


a@QOne plant at Secanquim showed a very decided instance of variegation 
with white and red, though the latter color might have been due to an increased 
tendency of the white portions to take the red discoloration common on normal 
leaves. The lower branches of the plant show only normal green coloration, 
and a part of the upper branches is also normal in color and size, and with 
fruits rather above the average size. The variegated branches do not regu- 
larly alternate, nor do they come all from one side, but they might still have 
connection with the phyllotaxy. There’ seem to be two stages of the variega- 
tion, a white and a light greenish-yellow; the latter may belong only to young 
leaves. soth are distributed with the utmost irregularity, and both may 
affect the upper surface of the leaf while the under surface remains green, or 
vice versa, though the latter condition is much less common than the former. 
The etiolated portions of the leaves, involucres, and fruits do not attain the 
full size of the corresponding normal organs, so that the parts affected are 
more or less unsymmetrical, though where the variegation is slight this result 
may be apparent, or if it be complete the symmetry is not affected. Except 
for two premature bolls the seed was not ripe, and these were from the nor- 
mal lower part of the plant. 


’ ACCLIMATIZATION OF KEKCHI COTTON. 17 
EFFECTS OF GUATEMALAN CONDITIONS ON UNITED STATES VARIETIES. 


The behavior of the United States varieties under changed climatic 
conditions in Guatemala is interesting in several ways. The “ King,” 
which in the United States appears to resemble the Guatemalan 
variety most nearly, here loses most of its distinctive characters and 
breaks up into’a variety of types, many of which would not be recog- 
nized in the United States as at all related to King. One of these is 
« “limbless” or “cluster” variety, which for a time appeared to 
Mr. Kinsler as a very promising new sort. It was smaller and dis- 
tinctly earlier than King plants of the normal type, and seemed 
likely to be more productive, but only a few bolls developed, and 
these proved to be of abnormal form, with deep grooves or notches 
across the tip. 

One of the features in which the change of climate seems to pro- 
duce remarkable effects is that of earliness. The King, which in the 
States is looked upon as the earliest variety, is found by Mr. Kinsler 
to be somewhat exceeded in this respect by “ Allen,” which has not 
been looked upon as a competitor. The Sea Island and Egyptian 
varieties, too, prove to be much more precocious than was expected. 
Some of them begin flowering almost as soon as the Upland sorts. 
The Rivers variety of Sea Island cotton, in particular, was very 
early, robust, and productive, distinctly ahead of the near-by Janno- 
vitch, though not so tall. 


ACCLIMATIZATION OF KEKCHIT ‘COTTON IN THE UNITED STATES. 


It was not unexpected that the Kekchi cotton would show a change 
in its method of growth on being transferred to Texas. New condi- 
tions of soil and climate often cause notable disturbances of the 
organism. Some of the tropical cottons planted in Texas for experi- 
mental purposes have grown into large bushes without showing the 
shghtest tendency to produce fruit or even flowers. In 1904 cotton 
from Peru planted at Victoria, Tex., grew most vigorously to a 
height of 18 feet, but remained quite sterile. Jt is possible, however, 
that even in their own country these were what are called * tree 
cottons,” which usually grow to considerable size before beginning 
to flower. Letters from Mr. Kinsler, in charge of our experimental 
plot at Pierce, Tex., relate a similar behavior on the part of the 
Kekchi cotton, which at that place has grown large and rank; but 
toward the end of July it was beginning to fruit, so that the ripening 
of seeds in Texas is to be anticipated. 

Two or three years will probably suffice to diminish this abnormal 
vegetative vigor, due to the stimulus of the new conditions, and per- 
mit a return to the normal earliness of the variety. Similar results 


9962—No. 88—05 M——2 


18 WEEVIL-RESISTING ADAPTATIONS OF COTTON. 


have attended the introduction into Texas of Mexican varieties of 
corn. The plants grew 14 feet high the first year and bore very little 
seed; in the following seasons they became smaller, earlier, and more 
productive. 

The probability that the Kekchi cotton can be grown even at the 
northern limits of cotton cultivation is strongly indicated by the 
results of an experiment at Lanham, Md. (1905). In favorable sea- 
sons cotton can be grown to maturity as far north as Washington, but 
the present year has been very unfavorable, the summer months being 
for the most part cool and rainy, and with several intervals of 
unusually low temperature. The cotton, which was planted intention- 
ally in rather poor soil, to avoid too great luxuriance of growth, ger- 
minated very badly and remained small and stunted until August. 
The Kekchi rows have, however, produced more plants, and more of 
these have grown to maturity than with any of the domestic or for- 
eign varieties included in the test. The Kekchi type has also remained 
more constant in Maryland than did the King variety when grown in 
Guatemala, though there are obvious differences between individual 
plants. Two plants in particular were found to have numerous buds, 
some ready to blossom before any of the others had begun to show 
signs of productive maturity. 

It might be feared that a variety newly introduced from a tropical 
country would be likely to suffer more from low temperatures than 
our United States varieties, but this seems not to be the case with the 
Kekchi cotton, even when the cold is carried down to the freezing 
point. There were light frosts in Lanham about the end of Septem- 
ber, just sufficient, as it happened, to do appreciable damage to cotton 
in low ground. The Kekchi plants did not suffer more than the 
American Upland varieties. The difference, if any, was in favor of 
the Kekchi cotton, perhaps on account of the closer foliage. 

Many annual plants, even those of tropical origin, are most vigor- 
ous and productive at their northern limits of growth, not, as has been 
supposed, because this is the coldest part of their range, but because 
the heat and sunlight, necessary to plant growth, are greater during 
our summer months than can be secured in a similar time in the 
Tropics, owing to the much longer days of our northern latitudes. 

The Pachon cotton from western Guatemala, though it has grown 
taller at Victoria, Tex. (52-79 inches), than at Lanham, Md. (80-40 
inches), has produced numerous buds in Maryland, but none in 
Texas. The Kekchi cotton also appears to have been more productive 
at Lanham than at Victoria, to judge from a recent partial report 
from Mr. Argyle McLachlan. 


a Cook, O. F., 1902. Agriculture in the Tropical Islands of the United States, 
Yearbook of the United States Department of Agriculture for 1901, p, 367. 


EARLY BEARING AIDED BY LONG BASAT. BRANCHES. 19 


It is very possible, therefore, that if the Guatemalan variety is able 
to thrive in the United States it will ripen its crop here in even less time 
than it requires in Guatemala, and this is rendered the more probable 
from the fact that in Guatemala the cotton has to be planted in the 
rainy season and is obliged to exist for the first few months under 
conditions of excessive moisture. The dry season of this district is 
short and uncertain. For two years, 1903 and 1904, the Indians were 
unable to burn their clearings, so that the corn crop failed and the 
community was reduced to the verge of starvation. The cotton crop, 
in normal seasons, is said to be planted in the latter half of October 
and ripens in March. 

The introduction of a dwarf, short-season cotton would require, of 
course, something of a change in cultural methods in the South, since 
the smaller size of the plants will need to be compensated by closer 
planting. It will be readily understood that to secure the setting of 
a crop in the minimum of time as many plants as possible should 
be set at work. The question is not that of the maximum product for 
each plant or for a given area. With the weevil in the field the time 
factor becomes of chief importance. . 

Little is gained in reality by the rank growth of the larger varie- 
ties; in fact there is a distinct loss in earliness, even though some 
bolls are set in the early part of the season. If these are overshad- 
owed and starved by the continued upward growth, the crop is delayed 
and the lower part of the plant becomes, on the whole, distinctly 
unproductive. 


EARLY BEARING FACILITATED BY LONG BASAL BRANCHES. 


The earliness of the Kekchi cotton is made possible by the fact that 
the bolls are nearly all borne at the base of the plant, the upper 
branches and their folage serving merely to assist in bringing to 
maturity the fruits which are set while the plant is still very young. 

Like several other tropical economic species, such as coffee, cacao, 
and the Central American rubber tree, the cotton plant has two kinds 
of branches—the true or primary branch, which arises in the normal 
position of branches in the axil of the leaf, and the secondary or fruit 
branches, one of which arises at the side of each primary branch. In 
most varieties only a few of the true branches are developed: often 
none at all. They are almost always plainly indicated, however, by 
a small bud or a stunted leaf or two, in case the bud has not remained 
entirely dormant. 

Cotton plants are either right-handed or left-handed in the sense 
that on the same plant all the secondary branches come out on the 
same side of the primary branches. It is possible, therefore, to de- 
termine by its position whether any particular branch is a primary or 


20 WEEVIL-RESISTING ADAPTATIONS OF COTTON. 


a secondary. But the function of the two sorts of branches does not 
always remain as distinct as in the coffee and cacao. A primary 
branch, like the main stem, never bears any flowers; it produces only 
leaves and other branches, mostly secondary. 

Secondary branches, on the other hand, produce normally a flower 
bud at the axil of each leaf, and this rule holds very generally, except 
that at the lower part of the plant it sometimes happens that a branch 
which has the secondary position functions as a primary; that 1s, 
instead of bearing buds and flowers it produces only leaves and sec- 
ondary branches. In the Kekchi cotton, as grown crowded together 
in the Indian fields, the primary branches seldom appear, but when 
more space is allowed and the soil is fertile it is usual for two branches 
to start from the axil of each of the lower leaves, one promptly pro- 
ducing flowers, the other assisting in the rapid increase of the leaf 
surface of the plant and of its power to elaborate food. 

Under the popular idea that plants draw their food from the 
ground the possession of branches which bear little or no fruit might 
be looked upon as an undesirable character, but when we take into 
consideration the fact that the leaves instead of the roots are the true 
assimilating organs of the plant it becomes apparent that a variety 
of cotton which develops its lower primary branches may have an 
advantage in earliness over one which is obliged to depend for its 
foliage upon secondary or fruit-bearing branches. In the matter of 
determinate habits of growth these primary branches are also a fea- 
ture, because they enable a plant to produce a full quota of leaves 
without unduly increasing the number of fruiting branches and thus 
continuing to add to the number of superfluous buds. 

The most obvious characteristic of the Kekchi cotton as it grows in 
our experimental plots is the long basal branches, which often equal 
or exceed in length the main stem itself. The most prolific branches 
of the United States varieties are those which come out from the main 
stem at the height of about a foot, but the bulk of the crop on the 
Kekchi cotton is borne much closer to the ground. (PI. II, fig. 2.) 
The long basal branches facilitate the early ripening of a uniform 
crop of cotton, but they will not be an advantage under all cireum- 
stances; as, for example, in dry regions where the weevil can be held 
in check by open culture. The necessary exposure of the fallen 
squares to the full sunlight on hot, dry soil would be interfered with 
by a plant of low spreading habit and dense foliage. 


EARLY REJECTION OF SUPERFLUOUS SQUARES. 


That the Kekchi cotton has a limited or determinate growth and 
does not take advantage of the perpetual summer to become a tree or 
even a large bush is evident from the fact that in the latter part of 


EARLY REJECTION OF SUPERFLUOUS SQUARES. 21 


the season most of the flower buds and leaf buds blast and fall off 
while still very young, before the weevil would give attention to them. 
By the time the first of the cotton is beginning to ripen, most of the 
plants have ceased flowering and no new leaves are being put forth. 
Generally there are bolls only near the base of the plant. 

It is a normal character of the cotton plant that the fruiting 
branches shall produce a bud at each node or joint; that is, at the base 
of each leaf. If all these buds were to be retained and treated impar- 
tially to the food materials which the plant is able to supply, the 
result would undoubtedly be disastrous, since the plant would be able 
to bring very few of its fruits to maturity, perhaps none at all, unless 
a part of the burden were removed by the weevils or by other outside 
causes. It is under the necessity of throwing off a part of its load of 
fruit at one stage or another of its development, the younger the 
better. 

The rejection is accomplished by the formation at the base of the 
peduncle, or fruit stalk, of special layers of cells of soft texture, 
which soon disintegrate and allow the bud or young fruit to fall off. 
This is one of the many instances of the prodigality of nature, which 
makes so many allowances in advance for the accidents which beset 
the existence of all living things. The waste of buds is, perhaps, not 
so large in proportion among the perennial “ tree” cottons, which 
form a considerable shrub before beginning to blossom. In cultiva- 
tion, however, the tendency has always been to encourage early bear- 
ing, and thus reduce the early vegetative period of the plant and 
bring it to a precocious maturity. The result is that fruiting branches 
are produced, even on young plants, and buds are formed out of all 
true proportion to the actual productive power. 

The babit of rejecting a large part of the squares and bolls is espe- 
cially obvious in the “ cluster cottons,” varieties in which the branches 
are abnormally shortened, so that the leaf surface of the plant is still 
further reduced. This cuts down still more the productive power of 
the individual plant, though there may be a gain in the number which 
‘an be grown on a given area. 

But cluster cottons have not learned to moderate their promises to 
correspond with their powers of performance, and continue to set 
vast numbers of buds, flowers, and bolls, which they are unable to 
ripen. The same is true to a less obvious extent of all our Upland 
varieties, but until the advent of the boll weevil the superfluous buds 
were not a serious factor, and the waste under favorable conditions 
was often well compensated by the power to recover and set a new 


“In Texas it is believed that rain at the time of flowering reduces the crop to 
half the normal quantity, or even less. The explanation given is that water 
settles in the flowers and prevents fertilization. This might serve as an addi- 
tional indication that cotton originated in a dry climate. 


22 WEEVIL-RESISTING ADAPTATIONS OF COTTON. 


crop when in unfavorable seasons the earlier buds were lost, or when, as 
occasionally happened in southern Texas, there was a liberal top crop, 
or second period of bearing, late in the autumn months. 

The presence of the weevil alters all these factors. The superfluous 
buds become positively detrimental, for they furnish the breeding 
grounds for successive generations of weevils and enable the pest to 
attain in the latter half of the season such numbers that a top crop 
not only becomes utterly impossible, but a menace is prepared for the 
cotton of the following year. For, although only a small proportion 
of the weevils live through the winter, the number of survivors un- 
doubtedly has a very practical relation to the supply maintained at 
the end of the previous season, and this again is merely a question of 
this persistent production of buds, now much worse than useless. 

A short-season variety of cotton having a sufficiently determinate 
habit of growth would by itself constitute a solution of the weevil 
problem. The Department’s entomological investigations in Texas 
indicate that it is only the weevils hatched in the last month of the 
growing season—in October or November—which have a prospect 
of surviving the winter. A cotton which ceased to produce buds 
after July or August would remove the chance of wintering over 
from all the weevils except the few that might develop in the bolls, 
an almost infinitesimal number compared with those that now attain 
maturity in the squares. Much would be gained, of course, if all 
planters would promptly pick their cotton and then pull up and 
destroy the plants, being especially careful to collect the infested 
bolls. But to carry out efficiently such a programme is difficult and 
expensive. 

To what extent, if any, the Kekchi cotton will meet this need of a 
short-season determinate variety, it 1s too early to form an opinion, 
but the fact that it has these qualities to a higher degree than any of 
the varieties hitherto known in the United States must be accepted 
as evidence, at least, that the possibilities of this method of protection 
have not been realized. In the latter part of the season the Kekchi 
cotton ceases the upward growth of the main stem and its branches 
and regularly drops the greater part of its buds before they are large 
enough to be entered or fed upon by the weevils, and the analogies 
to be drawn from the habits of other plants will justify persistent 
efforts toward the development in this and in other stocks of the 
habit of rejecting the buds still earlier or of not forming them at all 
after the first crop of fruits has set. Many plants have, in fact, 
exactly this habit so desirable in cotton; they continue to flower until 
permitted to set seed. 


SEASONAL BEARING OF PERENNIAL VARIETIES. 23 
SEASONAL BEARING OF PERENNIAL VARIETIES. 


The continued existence of perennial cottons in weevil-infested 
countries, like Guatemala, proves the presence in these also of means 
of protection. One of the most important is, doubtless, the produc- 
tion of an annual crop at a definite season, leaving the weevils with- 
out opportunity to breed in the intervening months, thus greatly 
reducing their numbers. 

The popular impression that tropical plants take advantage of the 
continuous summer climate and blossom continuously is correct only 
for a small minority. Where there are definite wet and dry seasons 
many tropical plants have-alternating periods of growth and rest 
almost as pronounced as in temperate climates, and even in regions 
of continuous humidity there are some species which shed their leaves 
annually and rest for a time. 

A further general reason for a simultaneous annual blossoming of 
all the flowers of a species is undoubtedly to be found in the greatly 
increased opportunities of cross-fertilization, just as»many insects 
swarm and many birds and mammals collect in flocks before the 
breeding season. Simultaneous flowering is carried to a remarkable 
extreme among the bamboos, where whole species grow for long series 
of years without flowering, and then flower and die at once over long 
distances and in spite of local diversity of conditions which might be 
expected to advance or retard maturity. 

Accordingly, while it would not be reasonable to insist that peren- 
nial varieties of cotton have adopted the habit of annual flowering 
only because of the boll weevil, the analogy of other plants may be 
invoked to show that such a character can be brought about by select- 
ive influence. The weevil could certainly assist in the development 
of such a tendency, especially if there were a season of the year in 
which the insects were less numerous, from climatic or other external 
causes as yet unknown. 

The tropical varieties of cotton are, as is well known, mostly peren- 
nial, and some of them develop into trees of considerable size, the 
trunk attaining a diameter of 6 or 8 inches, and the main branches a 
length of 15 or 20 feet. The existence in Mexico of tree cotton 
immune to the weevils has been reported, but as yet this has not been 
substantiated. Possibly the weevil has not yet penetrated some of the 
remote and arid parts of the republic. In eastern Guatemala, at 
least, the tree cottons appear to enjoy no immunity from the weevil, 
and at the time of the visit of the writer it was often impossible 
to secure uninjured bolls, even as samples of the varieties. The 
native cottons of the island of Cuba, according to Mr. E. A. Schwarz, 
also have the habit of annual blossoming, in the intervals of which 
the number of the weevils becomes greatly reduced. The cutting back 


24 WEEVIL-RESISTING ADAPTATIONS OF COTTON. 


of the cotton by the Indians at Rabinal, as described in the next 
paragraph, is an artificial means of attaining the same end, but the 
native Sea Island cotton, found at San Lucas, and the Kidney cotton, 
at Tucuru, are the best Guatemalan examples of this protective habit. 


ANNUAL CUTTING BACK OF PERENNIAL VARIETIES. 


While the annual variety of cotton protected by the keleps is the 
basis of the only field culture found in eastern Guatemala, the Indian 
population of the central plateau about Salama and Rabinal raise 
small quantities of cotton in their dooryards by means of another 
cultural expedient, apparently of great antiquity, as indicated by the 
extent to which the plant is adapted to the cultural conditions. The 
variety is perennial and has very small and inactive nectaries, pos- 
sibly as an adaptive result of the dryness of the climate. 

Most of the perennial varieties begin bearing only after the plants 
have attained considerable size, but the Rabinal cotton is a notable 
exception to this rule and avoids injury from weevils by the very 
prompt flowering and fruiting of the new shoots. 

The weevils are present in numbers, and are frequently seen crawl- 
ing about on the plants in a leisurely manner quite different from 
that which they affect in regions stocked with keleps. At the time 
of our visit not a single boll or bud of any except the smallest size 
could be found which had not been attacked by them. Nevertheless, 
a crop of cotton is secured at another season. In the month of April 
the Indians cut back all the bushes to the ground, and as the cotton 
is always planted immediately about the doors of their houses, where 
the chickens and turkeys congregate, the mortality of weevils at this 
time is probably very great. The protection of the domestic birds 
doubtless continues until the new shoots have grown out of reach. 

As soon as the plants are a few inches high they begin flowering, 
and before the weevils are sufficiently increased in numbers to become 
injurious a crop has been set. Flowers and fruit are commonly borne 
on the lower branches, only 6 or 8 inches from the ground. The 
Indians say that if the cotton is not cut back, but allowed to grow 
tall, they get no crop. The fact is that by that time the weevils are 
too numerous to permit normal bolls to be formed. Our search for 
such was quite in vain on both our visits to Rabinal. One boll which 
gave no certain external proof of injury was wrapped up in a paper 
and retained as a sample, but was overlooked in packing and not 
transferred to the preserving fluid. When the paper was unwrapped 
a few weeks later three dead boll weevils were found. 

The Rabinal cotton crop is evidently not large, but the harvest is 
said to be regular, and the area of fertile land in this district is so 
small that none of it is wasted. Much foreign thread is now 


HAIRY STALKS AND LEAF STEMS. 25 


imported, however, for weaving in the native looms. The industry 
has greatly declined in the last century, perhaps because chickens 
have been generally substituted for turkeys, which were formerly the 
only domestic fowl possessed by the Indians. 

All attempts at establishing field cultures of cotton in this region 
have failed. The local public, which does not take the weevil factor 
into consideration, is firmly persuaded that cotton will not bear ex- 
cept in the heavy, rich soil of the dooryards of the Indian villages. 


HAIRY STALKS AND LEAF STEMS. 


The weevil on foot is a rather slow-moving, clumsy insect, and it 
has been ascertained in the course of the investigations conducted by 
Messrs. Hunter and Hinds that its movements on the plants are to a 
great extent impeded by hairy stalks and leaf stems. The smooth 
Egyptian and Sea Island varieties were found to be more susceptible 
to weevil injuries than the hairy Upland sorts. The Kekchi cotton 
is still more hairy, however, than the United States varieties, and 
gains an added advantage from this fact. The longer it takes the 
weevils to climb from one bud to another the greater are the chances 
of their being caught by the keleps. The latter insects, owing to 
their much longer legs and the claws with which their feet are armed, 
are not only able to travel readily over the hairs, but find them of 
definite assistance. On smooth surfaces they are much less adroit 
in catching and stinging the boll weevils. In our experiments, too, 
they seemed to prefer the hairy Upland cottons to the.smooth Sea 
Island varieties. 

The difference between the two insects in this respect may also be 
illustrated by the fact that the keleps are unable to ascend a perpen- 
dicular surface of clean glass, a feat which the weevils accomplish 
without difficulty. 

That the Guatemalan cotton was more attractive to the keleps than 
the United States Upland and Sea Island varieties planted in ad- 
jacent rows seems to be indicated by a census of our plot experiment, 
taken April 19 by Mr. Argyle McLachlan. Kelep nests were 
found at the bases of 41 per cent of the plants of the other varieties, 


«Though distinctly hairier than our ordinary Upland varieties, the Kekchi 
cotton is exceeded in this respect by two other Guatemalan types, as well shown 
in a field test at Lanham, Md. The Pachon cotton obtained by Mr. William R. 
Maxon in the Retalhuleu district, of western Guatemala is distinctly more 
hairy than the Kekchi variety, though it seems to be lacking in other weevil- 
resisting features. The involucral bracts are not closed any more than in the 
Sea Island or Hgyptian types. The most hairy cotton of all is the Rabinal 
variety, at least in the form it has taken at Lanham. The plants are very much 
more robust in every respect than at home in Guatemala, and the hairy covering 
shares in this increased vigor. 


26 WEEVIL-RESISTING ADAPTATIONS OF COTTON. 


while 76 per cent of the plants of the Kekchi cotton were favored with 
kelep nests. This apparent preference may be somewhat exaggerated, 
perhaps, in view of the fact that the plants were often farther apart 
in the rows of the Kekchi cotton, the seed having germinated very 
irregularly. Moreover, the superior attraction of the Kekchi cotton 
for the keleps may not have consisted entirely in the greater hairiness 
or the more abundant nectar. The compact foliage and spreading 
lower branches of the Kekchi cotton give greater protection from the 
midday sun, which the keleps utilize by greater activity in the middle 
of the day. 

With the Sea Island varieties it seemed obvious, however, that the 
smooth stems, more open habit, and smaller supply of nectar result in 
distinctly less attention from the keleps. From 9 or 10 o’clock on hot 
days they foraged very little, and seemed to have quite disappeared 
from these varieties, though still to be found in considerable numbers 
on the stems of the Upland varieties and most of all on the Kekehi 
cotton, which appears especially adapted for the comfort and con- 
venience of the keleps. 

It was noticed, however, that the keleps went much more often into 
the involucres of the Sea Island and Egyptian varieties than into those 
of the Kekchi cotton, for the simple reason, probably, that they can 
get in more easily. 

In the latter part of the season, after the weevils had gained a foot- 
ing in this field, Professor Pittier noticed a very decided preference 
on their part for the Egyptian varieties, though it seems certain that 
this type of cotton had never been planted in the country before. The 
partiality of the weevils might be explained, perhaps, on such grounds 
as the relative absence of the keleps, and also the ease of access to the 
buds of the Egyptian cotton allowed by the more open involucres. 
However, a slight change of food or of conditions of growth is often 
a distinct advantage to plants and animals, so that a direct preference 
for a new variety as food might reasonably be expected, and similar 
instances are known. 

The greater hairiness of the stems and the presence of the keleps 
may also explain why the weevils in Guatemala were seldom seen 
walking about on the cotton plants as they do in Texas. On the 
other hand, they take to wing very readily and seem to prefer to 
alight in the open flowers, the only places on the cotton plants where 
they are safe from the keleps. 

The petals are so smooth that the keleps seldom descend into the 
flowers, and when they do sometimes appear to be unable to climb 
out. The petals of the Sea Island sorts are smooth even on the mar- 
gins, sometimes entirely so, while those of the Upland varieties are 
fringed with fine hairs well up on the sides, if not all the way round 
the apex. 


PENDENT BOLLS. 27 


The liability to capture by such an insect as the kelep may also 
afford an explanation of the peculiar sedentary habits of the male 
weevils, which often remain stationary in one involucre for long 
periods, or as long as their food supply lasts. It is necessary for the 
females to go about in search of fresh squares for egg laying, but 
similarly active habits on the part of the males would subject them 
to unnecessary danger. 

PENDENT BOLLS. 


The early bearing of the Kekchi cotton is made possible, as already 
noted, by the unusual development of the lower lateral branches, 
which often have a drooping habit, leaving the buds and bolls in 
pendent position, intead of upright. There are several advantages 
in this arrangement, one being that the instinct of the weevils leads 
them to the upper portion of the plant. In a very badly infested 
field without kelep protection, the only bolls which escaped the 
weevils were a few lying close to the ground on these lower pendent 
branches of the Kekchi cotton. Only at the time of flowering does 
the peduncle curve upward and give the flower its normal upright 
position. Thus these drooping lateral branches of the cotton, which 
seem to hide the buds and bolls away from the weevil, may be looked 
upon as a short step in the direction of such phenomena as the 
cleistogamous flowers of violets which remain buried in the ground, 
or those of the peanut which, after flowering, burrow into the soil to 
ripen their seeds. ° 

The flowers of the cotton plant open in a more or less directly up- 
right position, and this is retained by the boll in most varieties. In 
the so-called “ stormproof ” sorts, however, the bolls hang down, and 
this is looked upon by many planters as a distinct advantage, since 
when the boll is ripe and open the rain does not beat into it and wet 
the cotton or wash it out, but is shed by the protecting outer shell and 
involucre. 

On pendent bolls the external nectaries are brought upward, so 
that there is no danger of an abundant secretion of nectar being lost 
by dropping off. The surface of the nectary is papillate and has a 
somewhat waxy appearance. The secretion often collects as a dis- 
tinct drop. The nectaries are also more readily visited by the keleps. 
and the young bolls are likely to be better protected by them. If 
these remained upright, the weevils would be more likely to alight 
and enter the involucre at once. 

The drooping habit may have a mechanical explanation as the re- 
sult of the weakness of the comparatively slender lateral branches. 
It is also to be connected, perhaps, with the habit of early flowering 
and fruiting, since this would bring heavier bolls upon smaller and 
softer branches which would be twisted over by their weight. In 


28 WEEVIL-RESISTING ADAPTATIONS OF COTTON. 


the later and more upright varieties the flowers are not formed until 
the wood of the branches has hardened and become strong and rigid. 
Pendent bolls may thus be said to be incompatible with the cluster 
habit, which is brought about by the abnormal shortening and thick- 
ening of the lateral branches, which are able to hold their flowers and 
fruits rigidly upright, except as they may be turned sidewise by bemg 
crowded together. The cluster cottons, too, have the undesirable 
tendency to an abnormal multiplication of squares and young bolls, 
many more than the restricted leaf surface of the plant will enable 
it to ripen. This superabundance of flowers and fruits gives, how- 
ever, the greater encouragement to the weevil, and uses up vegetative 
energy which could be better employed in the prompt ripening of the 
bolls already set. It is no uncommon thing, however, for even half- 
sized bolls of cluster cottons to die without any sign of external 
injury or disease, while other varieties close by remain perfectly 
healthy. The cause is probably to be found in inadequate nutrition, 
but this might also be expected to give them increased susceptibility 
to injury from parasitic enemies of every kind. 

It is not unlikely, too, that the drooping habit may be connected 
with the greater size of the inside nectaries of the Guatemalan vari- 
ety. These are, as far as we have seen, larger than in any other 
American variety yet known; but the Asiatic cottons, which have the 
inside nectaries still larger and more active, are also more definitely 
pendent. The involucre is grown together at the base, as though to 
more thoroughly protect the nectaries from above—from the sun, 
which would dry up the secretion, and from the rain, which would 
wash it off. 

The nectar is formed in great abundance, and Mr. F. J. Tyler, of 
this Department, has called attention to the fact that the surface of 
the nectaries of the Asiatic cottons, instead of being merely papil- 
late, as in the American Upland varieties, has a covering of close- 
standing fine hairs, to which its velvety appearance is due. 

Finally, it may be remarked that for cotton with upright bolls the 
inside nectaries are often an element of danger, since when the secre- 
tion is abundant and is not removed it flows along the bases of the 
involucre and may serve as a medium for the germination of parasitic 
fungi or bacteria. Bolls are not infrequently found diseased around 
the base, apparently from this cause. 


EXTRAFLORAL NECTARIES. 


The cotton plant is not without floral nectaries similar to those of 
related genera, consisting of fringes of nectar-secreting hairs lining 
the pits inclosed between the bases of the petals. The nectar serves, 
doubtless, the same purpose as in other plants, the attraction of the 


EXTRAFLORAL NECTARIES. 29 


honey-loving insects through which cross-fertilization is secured. 
It does not appear, however, that the floral nectaries of the cotton 
have any connection with the problem of weevil resistance, although 
the weevils seem in Guatemala to spend a considerable part of their 
time in the flowers, which are indeed the only safe places for them 
on plants protected by the keleps. It had been noticed from the first 
that the keleps seldom visit the cotton flowers, and Mr. Kinsler has 
learned a very adequate explanation of this fact, namely, that they 
are able to climb out of the flowers only with considerable difficulty, 
and sometimes remain imprisoned in spite of all their efforts to 
escape. 

The functions of the extrafloral nectaries of plants are, as far as 
‘an be ascertained, similar to those of the floral nectaries to the extent 
that they attract insects, but beyond this there is a fundamental dif- 
ference; the floral nectaries and highly colored floral organs serve to 
secure visits of flying insects and thus maintain intercommunication 
and cross-fertilization between the different members of the same 
speciés, in spite of the fact that the individual plants are rooted fast 
in the ground. The extrafloral nectaries, on the other hand, attract 
to the plants insects which will remain upon them as permanent resi- 
dents, and this is the end secured by the extrafloral nectaries of the 
cotton. 

It may be objected by some that no use or benefit to the plant has 
been ascertained in the case of many species which have extrafloral 
nectaries and other insect-attracting devices. Much remains to be 
learned concerning these marvelous biological specializations, and 
there are two obvious alternatives which need to be canvassed before 
belief in the adaptive nature of extrafloral nectaries and analogous 
structures can be destroyed. The character and extent of many such 
specializations show that they have existed for a long time. They 
may have served protective purposes no longer apparent. The other 
consideration is that some of the symbiotic specializations existing 
between such plants as Cecropia and Acacia and their insect inhab- 
itants have arisen through selective encouragement, much as the 
special characters of our domestic plants and animals have been 
developed. It may be sufficient, in other words, that the nectaries or 
other structures be of use to the insects which have done the selecting. 
It may seem absurd to think of bushes or trees as having been domesti- 
cated by ants many thousands of years ago, but the wonder is no 
greater than that ants and termites regularly maintained subter- 
‘anean fungus gardens ages before mushroom culture was undertaken 
by man. 


30 WEEVIL-RESISTING ADAPTATIONS OF COTTON. 
NECTARIES OF THE LEAVES. 


The midrib of each leaf bears on the under side an oblong pit, 
from which a drop of nectar may often be seen to exude. This is 
collected and eaten by the keleps, which are thus induced to visit 
all parts of the plants, especially while they are still small. 

The habit of collecting the nectar was not previously known to 
exist among the insects of the family (Poneride) to which the kelep 
has been referred. Nevertheless, the fact is not open to question. 
The process is easy of observation in even greater detail than with 
the true ants or the bees, because the keleps do not, like these insects, 
have the art of regurgitating their food. They merely lap the nectar 
up to form a drop, which, protected by the widely opened mandibles, 
is carried into the nest to feed the queen and the young. 

Nectaries, or at least nectary-like depressions, are to be found 
probably on the leaves of all varieties of cotton, though very small 
and apparently inactive on some of the larger tree sorts.¢ The 
shape of the nectaries also varies greatly in the different species and 
varieties, some being longitudinal, others transverse, and still others 
crescentic or even sagittate. Some varieties have nectaries on the 
three principal veins, and some even on five veins. 

The leaf nectaries of the Kekchi cotton are to be found on the 
midrib of the leaf about 1.5 em. from the base. They consist of a 
rather shallow longitudinally oval depression surrounded by a broad 
raised rim. The midrib often appears distinctly narrower above the 
depression than below it, as though there were extra tissues to supply 
it. The secretion is quite active, nearly all the nectaries showing a 
small amount of liquid, which sometimes spreads out on the adjacent 
surfaces. 

These nectaries furnish, as might be expected, a medium favorable 
for the growth of molds or fungi, and there is often a considerable 
network of dark-colored fungus mycelium creeping in and about the 
moistened depressions, and with occasional erect, needlelike points, 
which may be fruiting bodies. 


«This was not true, however, of a Mexican “tree cotton” of the Upland 
type grown in the Department’s experimental plots in Texas last year. Large 
nectaries were generally present on three veins of each leaf, and the midvein 
often had two. They were of the crescentic or sagittate type, but often 
extremely long and distorted. Another Mexican tree cotton, with a different 
type of lighter green foliage, suggesting that of Bixa, “had nectaries only 
on the midvein and these reduced to a narrow groove. The vein was not 
thickened nor the margins raised. The two varieties were about as different 
as could well be with respect to nectaries. Neither produced either flowers 
or fruit, so that their true relationships were not to be ascertained. 


NECTARIES OF THE INVOLUCRE. 31 
EXTERNAL NECTARIES OF THE INVOLUCRE. 


The Guatemalan cotton protected by the keleps has three broadly 
oval or reniform pits at the base of the involucre, one at the middle 
of the base of each of the involucral leaves.*. These are larger, deeper, 
and more active than the nectaries of any of the Texas varieties as 
yet observed, though there is very great diversity of size and nectar- 
secreting activity. In some of the varieties these nectaries are 
reduced to mere rudiments or are entirely wanting. The depres- 
sion may be present, but with no secreting tissue. The variety nearest 
approaching the Guatemalan cotton in having large and active necta- 
ries is the Redshank, but the King and other related sorts also have 
fairly large nectaries. 

The drooping or pendent position of the bolls in the Kekchi cotton 
may be correlated with the special development of these nectaries, as 
already noted. In the middle of the day the keleps are not very 
active, but the nectaries are sometimes full to overflowing. If the 
bolls kept the erect position usual in the varieties cultivated in the 
United States the nectar would frequently drop off and be lost, but 
when the fruits hang down the cuplike nectaries are brought upper- 
_most and hold the liquid much longer. 

The evolutionary origin of these nectaries is fairly obvious. The 
bracts are to be looked upon merely as modified leaves, with nectaries 
which have increased in size and activity as the leaves have become 
smaller and more specialized. , 


INNER NECTARIES OF THE INVOLUCRE. 


As though to induce the keleps to come inside the involucre and 
thus more effectually protect the young buds and bolls against the 
weevil, the Guatemalan cotton is also provided with unusually large 
interior nectaries, alternating in position with those of the outer 
series and thus placed opposite the edges of the involucral leaves or 
bracts. These inside nectaries, like the outside ones, are larger and 
more active than those on most of the cottons cultivated in the 
Southern States, but the closing of the involucre and the devolopment 
of the inside’ nectaries have been carried much farther in the Old 
World cottons belonging to the species Gossypium herbaceum. 
Here the external nectaries are quite wanting, but the internal ones 
are enormously larger and heartshaped, and secrete nectar in such 
quantities that it often flows out in the groove between the adnate 


#Instances are occasionally found where only two nectaries are developed, 
but such deficiencies are much less frequent than in other varieties of the 
Upland and Sea Island series. The Rabinal cotton commonly has only two 
external nectaries. The Old World cottons thus far observed have no nectaries 
in this position. 


32 WEEVIL-RESISTING ADAPTATIONS OF COTTON. 


bracts to moisten the edges of the involucre. As yet, however, the 
purpose of these adaptations in the Asiatic cottons is entirely 
unknown, both the boll weevil and the kelep being absent in the 
Eastern Hemisphere. 

The botanical homology of the inner nectaries is somewhat different 
from that of the outer. They correspond in all probability with the 
nectaries which are found on the calyx of some of the species of 
Hibiscus, but there the calyx is large and covers the buds and each 
sepal bears a nectary near its middle. 


NECTARIES OF GUATEMALAN SEA ISLAND COTTON. 


A variety of Kidney cotton planted in small quantities by the 
Indians at Trece Aguas, Guatemala, has the outer nectaries very 
variable in size and commonly quite wanting. The inside necta- 
ries seem always to be developed and are unusually large, being ex- 
ceeded, as far as known, only by those of the Asiatic varieties. The 
nectar secretion is also very abundant. No weevils were found upon 
this cotton, nor any keleps. 

On the other hand, the free-seeded Sea Island cotton found by 
Mr. Kinsler in the San Lucas” neighborhood, not far from the 
kelep cotton culture of Secanquim, reverses again the tendency of the 
Kidney cotton to the great development of the inner nectaries and 
the suppression of the outer. The latter are, in the San Lucas cotton, 
nearly always present, of rather large size, and of a red color. The 
inner nectaries are often rudimentary or quite absent. 


CONTINUED SECRETION OF NECTAR. 


Our Upland varieties commonly secrete nectar only at the time of 
flowering, but in the Kekchi cotton the liquid continues to exude 
until the boll is nearly or quite full grown, thus securing the protec- 


«This variety not infrequently produces flowers with only two bracts, closely 
appressed, like a clam shell. In one such instance there were two nectaries at the 
base of each bract, or, to be more exact, two separate nectaries on one side and 
one partly divided nectary on the other, as though the nectary belonging to the 
deficient third bract had separated into two parts and joined the other necta- 
ries. 


’This San Lucas Sea Island cotton is probably the variety in which the 
weevils were found abundant in 1902, when the first intimation was gained 
that the Kekchi cotton had means of protection against the weevil. The San 
Lucas cotton is attacked not only by weevils, but by another long-bodied in- 
sect larva, evidently lepidopterous, that gnaws through the boll at the ends, 
both from above and below, and eats out the seeds. Nothing of the sort has 
been seen in the fields protected by the keleps. There was also noticed in this 
cotton an occasional abnormality closely comparable to the navel orange. 
Rudimentary parts like a small secondary boll were found in the middle of 
bolls otherwise normal. The orange tree and the cotton plant belong, it may 
be remembered, to related families. 


BRACTLETS SUBTENDING INNER NECTARIES. 33 


tion of the keleps for a longer period. The temporary character of 
the secretion in our United States sorts was reported by Professor 
Trelease several years ago. 

In Guatemala, however, the young bolls seem to be quite as effi- 
cient as the flowers. It is even possible that this generosity on the 
part of the plant is excessive, since if the number of keleps is small 
they may find all the nectar they need on the lower bolls, and hence 
have less inducement to inspect other parts of the plant. Under 
favorable conditions in Texas the cotton plant produces a much 
larger number of flowers than in Guatemala, so that what is lacking 
in quantity may be made up by numbers, in case it should become 
possible to utilize the keleps in Texas. 


RACTLETS SUBTENDING INNER NECTARIES. 


The Kekchi cotton is distinguished from all our Upland and Sea 
Island types by the more regular presence and much larger size of 
a series of bractlets, a pair of which usually subtends each of the 
inner nectaries. In other varieties these are either wanting entirely 
or are rare and rudimentary.” The bractlets are inserted somewhat 
obliquely, with their margins in contact below the nectary. 

Sometimes they serve to conduct nectar to the edge of the involucral 
bracts, the nectar following along between the slender bractlets like 
ink between the nibs of a pen, as though to coax the keleps inside the 
inmvolucre. This must happen rather infrequently, however, to judge 
from the great irregularity in the size of the bractlets. Sometimes 
they are half an inch or more long, and extend well into the angles of 
the involucre, or even project outside. (Pl. III.) Nevertheless. it 


«Professor Trelease, who studied the American Upland varieties, appears not 
to have found the bractlets in pairs. He says: “ These glands (the inner nec- 
taries) belong in reality to an inner whorl of three bracts, alternating with 
the outer ones, but generally wanting. In stunted plants, especially as cold 
weather comes on, one or more of these inner bracts may be found.” (See 
Comstock, 1875, Report upon Cotton Insects, 324.) 

The shape and position of the bractlets seem to warrant the suggestion that 
they represent the stipules of the outer bracts instead of an independent inner 
whorl of bract leaves which has first become specialized and then become rudi- 
mentary. The suggestion has the further warrant in that it may help to explain 
the numerous inyolucral appendages of some of the related plants, which range 
about the number 9—that is, 3 leaves and 6 stipules. The normal number should 
be 6, if the two whorls of leaves were represented. One of the Guatemalan 
species of Hibiscus examined with this interpretation in mind seemed to con- 
firm it by showing very often 3 of the appendages broader than the others, 
though-the total number varied from 8 to 11, with an irregularity quite compar- 
able to that of the bractlets of the cotton. Even the bracts of the cotton some- 
times vary, involucres of 2 bracts being found occasionally, and in rare 
instances 4. 


9962—No. 88—05 mM——3 


84 WEEVIL-RESISTING ADAPTATIONS OF COTTON. 


may well be questioned whether these inner bractlets have remained 
unusually large in the Kekchi cotton because they have a definite 
function or because of the greater size and activity of the adjacent 
nectaries. 

A variety of cotton called Pachon, planted rather extensively in the 
Retalhuleu district of western Guatemala, and likewise protected by 
the keleps, is similar to the Kekchi cotton in many respects, including 
the possession of these large stipular bracts subtending the inner 
nectaries, but with the addition that the bracts are fringed with long 
hairs, as though to hold the nectar the better. This may also be the 
function of the hairs which cover the nectaries of the Old World 
cottons 

EFFICIENCY OF THE KELEP PROTECTION. 


The special development of the extrafloral nectaries in the Kekchi 
cotton has been noted in former reports, it being the nectaries which 
attract the keleps to the cotton plant. That the kelep preys upon boll 
weevils and protects the cotton ‘crop was learned last year, but it was 
still possible to question the practical value of this form of defense. 
Such doubts would not have survived an inspection of our recent 
experiments in Guatemala. A small field of cotton just outside the 
kelep area was attacked by the weevils in such numbers that not a 
single normal boll developed on any of the United States Upland and 
Sea Island varieties. In the field protected by the keleps the weevils 
obtained no footing until the plants were well grown and an excellent 
crop of full-sized bolls had been developed. 

To test the efficiency of the keleps as destroyers of boll weevils and 
as protectors of cotton would be possible in Texas only by stocking a 
large area with keleps—a difficult and expensive undertaking. No 
small tract would give a fair indication, since the weevils from the 
whole neighborhood would continue to come in, and, although they 
might soon be captured, would be able to do vastly more damage than 
would be possible if the whole region were stocked with keleps. 

In Guatemala, however, it was quite possible to contrast a protected 
with an unprotected piece of cotton by the simple expedient of plant- 
ing outside the area occupied by the keleps. A more striking result 
could hardly be imagined. For several weeks, during which the two 
plots were under continuous observation, the one remained almost 
entirely free from weevils and weevil injuries and set an excellent 
crop, while in the other scarcely a flower opened or a boll developed. 
The very few exceptions were on the concealed drooping branches of 
the native Kekchi cotton. 

The weevils became, indeed, too numerous for their own prosperity 
and fed upon and destroyed the very young buds before they were 
old enough to breed larve. Twenty-five fallen squares collected and 


EFFICIENCY OF THE KELEP PROTECTION. 35 


examined from under the plants of the plot without keleps yielded 
only 6 larve, or 24 per cent. They even attacked the young leaf 
buds, as observed last year at Rabinal. 

A large proportion of the injuries were caused by feeding punc- 
tures, but this only emphasizes the fact that the number of weevils 
which migrated into this plot was sufficient for a complete destruec- 
tion of the crop, and since the other experiment protected by the 
keleps was much nearer to the fields of the Indians there is every 
probability that the weevils would have been, if possible, even more 
numerous if the keleps had not been at hand to catch them. 

The unprotected plot was located at about one-quarter of a mile 
outside of the belt of Indian cotton culture, on land not inhabited by 
keleps. The weevils lost no time in finding the new field. Infesta- 
tion was complete, and quite as destructive as in Texas, the weevils 
being so numerous ,as to overcome whatever resistance the cotton 
might have been able to oppose to smaller numbers of the pests. 
The Sea Island, Egyptian, and United States Upland varieties were 
not permitted to produce flowers or even full-sized buds, and even 
the native Guatemalan varieties shed their squares before the per- 
sistent onslaughts of the weevils. 

Cotton is regularly cultivated by the Indians in this immediate 
neighborhood, and Indian plantings more or less infested with 
weevils were to be found within short distances of the protected field. 
Nevertheless, the keleps proved to be sufficiently abundant on this 
piece of ground to completely exclude the weevils. There were 
enough, indeed, to protect with apparent impartiality all the kinds 
of cotton included in the experiment, but if the numbers had been 
less and the plants had been closer together, as in the Indian fields, 
we may be sure that those producing the most nectar would have 
received the most protection from the keleps. 

The weevils were seldom to be found in the plot stocked with 
keleps as long as the Indian cotton remained in vigorous growing 
condition, but about the time the Indian cotton ripened, the weevils 
seemed to make a more determined raid on our field, and along one 
side nearly every plant suffered somewhat, though the weevils 
could rarely be found except in the open flowers, which seem 
to be recognized as their only safe roosting places. In a week or 
ten days there was a distinct falling off, so that very little damage 
was being done, and there was another short interval of practically 
complete protection. But after this a renewed onslaught began and 
‘the numbers of weevils gradually increased. the Upland and Sea 
Island plants continuing to produce thousands of new squares in 
which the weevils were able to breed, quite as in the United States. 

That the keleps are definitely attracted to the cotton plants. as 
stated in previous reports, is fully demonstrated by the fact that 


36 WEEVIL-RESISTING ADAPTATIONS OF COTTON. 


many of the colonies moved their nests to new burrows excavated 
immediately at the bases of the cotton plants. In some parts of the 
field the proportion of cotton plants having kelep nests established 
about their roots reached nearly 75 per cent, whereas the chance 
that the positions of the cotton plants which stood in regular rows 
would coincide with those of kelep nests would not be one in hun- 
dreds. 

The success of this experiment would seem to justify fully the 
suggestions made in connection with the first announcement of the 
discovery of weevil-resisting adaptations of the cotton plant, namely, 
that the protection which these Central American varieties had 
been able to secure from the kelep had afforded them an opportunity, 
perhaps unique, of developing other resisting adaptations. The 
Kekchi and other related cottons, though having no monopoly of 
weevil-resisting characters, furnish, however, the only instance as 
yet known to scientific observation in which a field culture of cotton 
has been maintained for long periods of time under climatic condi- 
tions favorable to the boll weevil. 

In Central America, at least, the secretion of nectar by the cotton 
is not a useless or meaningless function, as observers of the plant in 
other parts of the world have sometimes supposed. The cotton is 
not the only plant upon which the kelep can live, nor the boll weevil 
the only insect upon which it preys. To secure the attention and 
obvious preference of the kelep the cotton has been obliged to put 
forth the superior attractions provided by its numerous extrafloral 
nectaries. 

This additional proof of the value and efficiency of the kelep does 
not affect, of course, the possibility of acclimatizing it in the United 
States. A more extended search in Guatemala resulted in finding the 
insects under a wide range of conditions, and at altitudes of from 200 
to 2,000 feet. It lives and thrives, moreover, in soils very much drier 
than those to which it was supposed last year to be confined. Last 
year’s experiments in Texas indicated likewise that the kelep with- 
stands drought much better than it does standing water in its burrows, 
and care is being taken this season to locate colonies with a view to 
adequate drainage. 

OTHER NECTAR-BEARING PLANTS VISITED BY THE KELEPS, 
’ 

The honey-colleeting habits of the keleps are not confined to the 
cotton. Another favorite is a species of Bidens (2B. pilosa) called by 
the Indians “tshubai,”’ which has considerable value as a forage 
plant, being of quick growth and succulent texture. 

The preference of the kelep for the tshubai as a second choice after 
cotton was noted last year, but no explanation was found, though 


THE INVOLUCRE AS A PROTECTIVE STRUCTURE. 37 


the plant was searched for nectaries. It was noticed by Mr. Kinsler 
that the keleps seemed to be giving especial attention to the midrib 
near its junction with the veins of the lower divisions of the leaf. 
Our lenses then revealed the fact that there are two minute raised 
wings or margins running along the upper side of the midrib and 
petiole, forming two narrow grooves in which the nectar is.evidently 
secreted. The grooves are also protected by a row of fine hairs which 
project across them from the raised margin. The behavior of the 
kelep thus receives a practical explanation, and the tshubai finds a 
regular place next to the cotton among the plants protected by the 
kelep. The nectar-secreting habit of the tshubai may also explain 
its being eaten so readily by stock, and may help to give it standing 
as a forage plant, in spite of its weedy and unpopular relatives. 

A second member of the composite family often visited by the 
keleps is the “ sajal,” a species of Melanthera (probably J/. deltoidea), 
which also has local value as a forage plant, being eaten greedily 
by horses and mules, even in preference to grass. No nectaries have 
been found on this. <A third composite, not yet identified, produces 
nectar in small depressions at the base of the leaf on the under side. 


THE INVOLUCRE AS A PROTECTIVE STRUCTURE. 


Cotton is the only plani known to be attacked by the boll weevil, 
and it is also unique among its relatives tn the possession of a large 
leafy involucre. This may be a mere coincidence, or it may be that 
the weevil has had a considerable influence in the development of the 
involucre, depending upon the antiquity of the contact between the 
insect and its host plant. The involucre has, it is true, functions 
other than the exclusion of the weevils, since it takes the place of the 
calyx in protecting the young bud, but the reduction of the calyx 
probably followed the enlargement of the bracts, instead of preceding 
it. But however originated, the large bracts have, at the present 
time, a definite value in the problem of weevil resistance. There are 
several specialized characters which appear as though definitely cal- 
culated to increase the efficiency of the involucre in excluding the 
weevils from the young buds. 


INVOLUCRAL BRACTS GROWN TOGETHER. 


Both the Kekchi and Rabinal cottons frequently have the involucre 
closed at the base, the three bracts being grown together, thus making 
it impossible for the weevils to enter from below. In the Sea Island 
and Egyptian varieties, as well as in some of the Upland sorts, the 
bracts are not merely divided to the base, but they often have the 
lower corners rolled back, thus leaving an open passage for the 
weevils. The Rabinal cotton much excels all the other varieties thus 


: 


38 WEEVIL-RESISTING ADAPTATIONS OF COTTON. 


far studied in the extent to which the bracts are grown together at 
the base. Sometimes they are united for a quarter or even a third 
of their length. (PI. IV, fig. 1, and Pl. X, fig. 1.) 


APPRESSED MARGINS OF BRACTS. 


In both of these Guatemalan varieties the margins of the bracts of 
voung involucres are firmly and closely appressed, in striking con- 
trast with the Sea Island and Egyptian varieties, where the bud is 
commonly exposed even when very young. This form of protection 
is effective while it lasts, but in the Rabinal cotton the involuere is 
too small, and the growth of the young bud soon separates the bracts 
and permits the entrance of the weevil. The United States Upland 
varieties are intermediate between the Sea Island and the Kekchi cot- 
tons in the degree to which the involucres are closed and the margins 
fitted together. A large proportion of the Upland involucres give 
ready access to the weevils, while most of those of the Kekchi cotton 
remain effectively closed for a longer period, as will be understood 
after a survey of the other involucral characters which conduce to 
the same result. 

In one respect the firmly closed involucres of the Rabinal cotton 
seemed almost like an advantage to the weevil rather than the con- 
trary, for the insect is not admitted to the bud until it is about large 
enough to furnish a place of development for a larva. The plant 
having taken control, as it were, of this relation, the weevils have not 
needed to possess an instinct against the destruction of young buds. 
Those of the open involucred Sea Island varieties often were attacked 
while still altogether too small to bring a larva to maturity. The 
advantage of the closed involucres les, no doubt, in the fact that they 
shorten the period of access and allow some of the buds to escape 
which would be punctured either for feeding or for egg laying if the 
weevil has a longer opportunity. (PI. IV.) . 

The Rabinal cotton culture is that in which the plants are cut 
back yearly to the ground. During the next month, or until the buds 
begin to develop on the new shoots, the weevils have no breeding 
places and nothing to feed upon except the leaves and leaf buds. In 
patches where the weevils are abundant the leaf buds are eaten out 
so persistently as to seriously interfere with the growth of the plants, 
and the very young flower buds were also reached in some instances 
by boring through the involucres. When attacked at this stage the 
buds wither and drop off. They serve the weevils only for feeding 
surposes, and their use in this way only postpones the time when 
breeding can be resumed. 

The cotton at Rabinal was often overrun by two species of small 
black ants, identified by Dr. W. H. Ashmead as belonging to the 


LARGE INVOLUCRES OF KEKCHI COTTON. 39 


genera Solenopsis and Tapinoma.* There was no indication, how- 
ever, that these afforded any protection against the weevils, although 
they might, perhaps, act as watchmen and scare weevils away when 
they happened to be present on buds or bolls where weevils had 
alighted, like other small ants which have been reported as attacking 
the boll weevil. The keleps belong in an entirely distinct category 
in being able to sting and carry off the weevils and make regular use 
of them as food. Instead of being of service to the cotton these 
small ants at Rabinal were a distinct injury; the Solenopsis was 
taking care of plant lice,” which often infested the cotton to a 
decidedly harmful extent. It continues and supplements the work 
of the boll weevils in stunting and distorting the plants. When the 
aphids are very numerous, the leaves are badly curled and growth is 
greatly impeded. 


LARGE INVOLUCRES OF KEKCHI COTTON. 


The Kekchi cotton has the bracts of the involucre much larger in 
proportion to the contained bud than the Rabinal cotton or than any 
of our Upland varieties. The possession of larger bracts constitutes 
a distinct weevil-resisting adaptation, since it permits the involucre 
to be more effectively closed and the protection to be continued for a 
longer time. Sooner or later, of course, the bracts must be separated 
by the growing bud. The larger the bracts the longer the bud can 
continue to grow before spreading the bracts apart. (PI. IX, fig. 1.) 

Prof. H. Pittier, who had charge of the Secanquim experiment in 
the latter part of the season, was especially impressed with the pro- 
tective utility of the larger bracts of the Kekchi cotton, as shown by 
the following summary of his observations: 

The large sizé of the bracts in proportion to the floral bud is a very important 
protective feature. In the Kekchi cotton the amplitude of these bracts is such 
as to completely inclose the bud at all times before the anthesis, and even in 
eases when they happen to be slightly separated the occlusion is maintained by 
the long hairs which fringe them on all sides. The length of these hairs con- 
stitutes a serious obstacle to the progress of the weevils, whose tarsi can not 
obtain a firm hold on the solid surface. I have seen them drop to the ground 
after many awkward attempts to gain access to the squares, while on the other 
hand the keleps did not seem to be impeded at all by the bristles. 


«The material was not sufficient for a conclusive determination of the species. 
Doctor Ashmead says: ‘“*‘ You have two distinct species of ants here. One, No. 
1, belongs to the family Myrmicids and is apparently the worker of Solenopsis 
picea Emery; the other, No. 2, belongs to the family Dolichoderidz and is 
apparently the worker of Tepinoma ramulorum Emery. I am sorry you did 
not have the different sexes, so that I could make positive of the species. In 
Solenopsis, as you probably know, there are four or five different forms, and it 
is not easy to identify from a single form.” 

b These have been identified by Mr. Theodore Pergande as Aphis gossypii, a 
species well known in the United States. 


40 WEEVIL-RESISTING ADAPTATIONS OF COTTON. 


To show the increased size of the bracts in the Kekchi cotton, I have carefully 
measured over 250 squares of five of the most promising varieties of the Upland 
species. The dimensions taken were the length of the floral bud, and the length 
and breadth of the bracts. The table, in which these data are condensed in a 
comprehensive form, shows a decided advantage in favor of the Kekchi cotton. 


TABLE I1.—Dimensions of floral buds and bracts of several varieties of cotton 
compared. 


Kekchi. Parker. King. Allen Jewett 
ol al el ~ ~ eal al al ol el ~ | je eal | Sol eal 
Length of SA ead a) en es a Peg te “dlas| sel elas) ee 
oral buc §69)95/98/ SG /g8/98) 9G) gh (85) on] ob /s5|salggigs 
(millimeters)., 20 /2ai58 SU Es Se et Selgglavlsgisg avlsslsg 
§2)M2 eh gB Me eH | ge) WH ae) gs] Ma| aa] ge | ton 25 
Bo ee }ee) er) ee) ee) eo) ee ee) 3°) ee) ee 
4 Ja a 14 fA |e 1] J4 18 |@ 18 eee 
| } 
nin. ment nent, neat mn, Went mn. nine mm mm 
Be os Da AN (Ba 35 Daeg Ee Pn, aR ee | eereye eens Roe ekey Rees; ests 
7- 8_. 1 28 18 2) & 19) 2522 ee ie ee 1 26 | 20:2 
S210) Sse 6| 39) 27 13} 31 20 5 | 33 19} 18} 34] 21 3| 38 | 26 
iB ES ee a 5; 42; 380| 16| 36) 24 7| 34| 2] 10) 34)| 22 2] 36) 21 
ib Sy eee es 3} 42) 30| 18|] 39) 2 9) 40") 241 Pats 7 | 23) 10) 39 | 26 
1S | a re. + 42 30 8 38 23. 6 44 25 13 | 39 24 7 41 | 28 
jy Cit. ae ee | 3| 47| 33 6| 39) 26 6| 40] 24 5 | 39] 25 5 | 39) 30 
git | pee Pred | 2] 52) 30 3} 43] 24 4| 42) 26 1| 49] 29 1} 52) 38 
dE Es i Ud [oma Y= 5-76 3] 48| 26 1 43] 25 1} 40) 2 2| 47 | 34 
Pee 3| 47 36 1 36 | 25 2} 41| 2 5 | 40) 2522222 
25-26_....-..- 2| 42) 30 Bi Ol.) V2 eae fee ee Th) Sul) Hae 1 | 48 | 33 
af 2 IED IY EL ae SS A DE 2| 44) 2% 2| 40| 2% 5 | 42.) - 2a. eee 
AES Res et td Pee ae Ee aes 2| 45 | .24 1 AGS) 2S. eer |. nc 2|-cee ee oe ae 
5 ae ee ee a = 1 Ai |. SO | aaa eee ee eee eee en) er Et 
5 3, ee 5 eee Deke An) Ae Mel Pacman MPM) fe Sa fee) ene ope yes 1 | 42°| 32 
Total FW) Pepe see) | Se They Peed ese 4355.2 (oie mee | fess 3d a id 


The advantage is particularly notable with respect to the greater 
width of the bracts, which enables them to remain much more effect- 
ively closed at the angles. In the Parker, King, and Allen varieties 
the bracts very seldom attain a width of 30 mm., while in the Kekchi 
cotton the average width for all except the smallest buds is above 


30 mm. 
OPENING, OR FLARING, OF BRACTS AVOIDED. 


The unusually large and well-closed bracts of the Kekchi cotton 
have another practical use in keeping the bud from drying out, as 
explained in the discussion of proliferation. 

The external indication of this difference is that in the Kekchi 
cotton punctured squares commonly do not open, or flare, by the 
spreading apart of the involucral bracts, while among the Upland 
and Sea Island varieties flaring is the regular rule. Quite a per- 
centage of the squares of Abbasi, Parker, King, and other varieties 
stand well open normally before any injury has occurred, but the 
Kekchi cotton seldom or never exposes its squares before flowering. 
The larger and broader involucre is also able to permit the protrusion 
of the flower without losing the power of closing and remaining 
shut for a considerable period after flowering, while the Parker and 
King varieties often remain quite open, so that the young boll is 
fully exposed to the weevils. 


sg 


EXTENT OF PROTECTION BY INVOLUORE. 4] 


An example of the promptness with which weevil injuries cause 
the involucres of our Upland cotton to open is well shown in a note 
by Mr. McLachlan: 


On August 8, at 2 p. m., a small cage was placed over a small plant of Parker 
cotton, and 5 female and 2 male weevils were introduced. The plant possessed 
86 squares, 4 flowers, and 9 bolls. The morning after the weevils were put 
into the cage several of the squares had flared and one had fallen. It would 
seem that the mechanical forces of the square are quickly affected by the work 
of the weevils. Here, of course, the punctures were numerous, because of the 
many weevils on the plant. Some of the squares were riddled with feeding 
and egg punctures. 


The buds of Kekchi cotton often recover from three or four punc- 
tures, though they might not do so if these were all made at the same 
time. But it often happens that squares with numerous feeding 
punctures remain closed and wither up without flaring. 


HAIRY MARGINS OF INVOLUCRAL BRACTS. 


‘In addition to their larger size, the bracts of the Kekchi cotton have 
the marginal teeth or laciniz more numerous and more hairy than 
those of our Upland varieties and able to afford more of an impedi- 
ment to the entrance of the weevils. The difference was very pro- 
nounced in our experimental plot, where King, Parker, and other 
familiar American sorts were planted beside the Kekchi. It is as 
superior in this respect to the other Upland varieties as they are to 
the Sea Island. 

The Kekchi and Rabinal varieties, though both belonging to the 
Upland series and having many similarities, have also very distinct 
differences, as, for example, in the present character. The small, 
firmly appressed bracts of the Rabinal cotton have the marginal 
lacinie few and small; sometimes the edges are nearly entire, or 
merely toothed. The hairy covering is also reduced to a fine, short 
coat, which can afford little or no impediment to the weevils. 


EXTENT OF PROTECTION BY INVOLUCRE. 


That the closed involucres do indeed contribute to the protection of 
the young buds from the weevils became very obvious in one of our 
experimental plots at Secanquim, located about a quarter of a mile 
outside the belt of Indian cultivation of cotton. There being no 
keleps to afford protection, the cotton soon became thickly infested 
with weevils, and very few bolls were allowed to develop on any of 
the plants. There was a notable difference, however, in the age at 
which the buds were punctured. As already stated, the edges of the 
bracts of some of the Sea Island and Egyptian varieties separate at 
a much earlier period than those of the Upland varieties, and the 


492 WEEVIL-RESISTING ADAPTATIONS OF COTTON. 


weevils commonly attack them in their very early stages, and even 
while they are altogether too small to permit the development of a 
weevil larva. It has been pointed out already by Messrs. Hunter 
and Hinds that the smooth stems and petioles of the Sea Island and 
Egyptian cottons render them much more readily susceptible to 
injury by the boll weevil than are the Upland types, and if we add 
to this the disadvantage arising from the later development and the 
more open involucres the possibility of protecting the long-staple 
cottons against the weevils seems small indeed. 

Instead of being immune to the boll weevil, as at one time hoped. 
the Egyptian and Sea Island varieties seem to be most lacking in 
weevil-resisting adaptations, as might, indeed, have been expected in 
view of the fact that they have been developed in regions to which the 
weevil has not yet penetrated. The Kidney cottons, which may be 
looked upon as representing the Sea Island type on the mainland of 
the American continents, have, as will be seen later, a peculiar 
feature of protective value. 


ADVANTAGE OF OPEN INVOLUCRES. 


It will be apparent from the facts already recited that the partly 
closed involucres of the Sea Island and Upland varieties now culti- 
rated in the United States serve little or no purpose in resisting the 
boll weevil. On the contrary, they often appear to be an advantage 
to the insect, serving, as they do, to hide the parasite from its enemies 
and protect it against the application of insecticides or capture by 
insectivorous birds.* 

The great variation in the size and shape of the involucre in the 
different varieties of cotton suggests the practicability of securing 
sorts with open involucres or with these structures reduced to small 
dimensions. If the weevils were to be caught by insectivorous birds, 
like the Cuban oriole, whose weevil-eating habits have been discovered 
by Mr. E. A. Schwarz, open involucres would be a distinct advantage. 
It might then be possible also to apply Paris green or other insecti- 
cides to young buds which are, except in the early spring, the 
exclusive feeding places of the weevils. 

The practicability of an open involucre will need, however, to be 
considered from another standpoint. It must be ascertained whether 
the young buds will bear full exposure. Unlike most of the related 
plants, the cotton bud is not protected by a calyx. The involucre may 
be necessary as a substitute, especially in dry climates. In humid 


aDr. H. J. Webber states that the desirability of open involucres has been 
appreciated and that selections of Upland varieties with a view to the develop- 
ment of this character have been made. 


BEHAVIOR OF PARASITIZED BUDS. 43 


regions, however, this requirement might be relaxed, and it is in such 
places that the injuries of the weevils are the greatest.” 


BEHAVIOR OF PARASITIZED BUDS. 
: SHEDDING OF WEEVIL-INFESTED SQUARES. 


In a dry climate, like that of the Mexican plateau region, the drop- 
ping of the squares in which the weevils have deposited eggs would 
constitute a very effective adaptation. The weevil larve do not sur- 
vive a thorough drying out of the squares. It is only in the arid 
districts of Mexico that the cotton plant has shown its ability to 
escape from cultivation and maintain itself without human assistance, 
if indeed it be not in some places a truly indigenous wild plant, as 
several botanists have reported. But in a moist region like the cotton 
belt of eastern Texas this habit of the plant has no practical use, 
since as many of the weevils die when the injured squares remain 
attached to the plant as when they fall to the ground. 


“It is generally true that squares seriously injured by the weevil sooner or 
later fall to the ground. Some plants, however, shed the injured squares more 
readily than do others. It seems to be a matter of individual variation rather 
than a varietal character. _Thus occasional plants retain a large proportion of 
their infested squares, which hang by the very tip of the base of the stem. 
Normally the squares are shed because of the formation of an absciss layer 
of corky tissue across their junction with the stem. In the case of the squares 
which remain hanging, the formation of this layer seems to be incomplete, or 
else it becomes formed in an unusual plane, so that while the square is effectu- 
ally cut off, it merely falls over and hangs by a bit of bark at its tip. In this 
position it dries thoroughly and becomes of a dark brown color. Plants 
showing 6 or 8 of these dried brown squares are quite common in infested 
fields. Although exposed to complete drying and the direct rays of the sun, 
the larve within are not all destroyed. * * #* 

“It seems a conservative estimate, therefore, to say that fully one-third of 
,these exposed dried squares may be expected to produce adults. Considering 
the exposed condition of such squares this seems to be a very high percent- 
age. * * * The observations made, however, certainly show that a complete 


aAfter the above had been written it was observed that the Pachon cotton 
from western Guatemala, grown in an experimental plot at Lanham, Md., has 
the peculiar feature of a large calyx, which completely covers the young bud~ 
and extends above it into long, slender, hairy tips. It may be that this is to be 
looked upon as still another weevil-resisting adaptation. The weevils would be 
able, undoubtedly, to bore through the calyx, but the hairy tips might hinder 
their access to the bud. The bracts are much smaller and much more open 
than in the Kekchi and Rabinal varieties, but the lacinize, or teeth, along their 
margins are rather stiff and are clothed with numerous hairs, stronger and 
more bristlelike than in the Kekchi and Rabinal varieties, and able to keep the 
laciniz from closing together. It may be that the greater rigidity of the lacinize 
and the bristles gives better protection than the open position of the bracts 
would indicate. The case is in reality quite different from that of the Sea 
Island varieties, where the bracts are both naked and open. 


44 WEEVIL-RESISTING ADAPTATIONS OF COTTON. 


drying of the square does not necessarily destroy the larva, and that a square 
may undergo far more exposure to direct sunshine than had been supposed 
possible without causing the death of the larva or pupa within.” @ 

It is to be remembered, however, that such disconnected squares are 
thoroughly dampened every night by the dew, and that a small 
amount of moisture may pass out from the plant through the shred 
of dead tissue. In either case the hanging boll might get more moist- 
ure and less heat than if lying on the dry ground, exposed to full 
sunlight. Suspended bolls are exposed to air temperatures only. | 

If no other means of avoiding the weevil becomes practicable a 
great extension of the cotton production into the semiarid districts of 
western Texas, Oklahoma, and even Kansas is to be expected. The 
long days of the more northern districts will conduce to the shorten- 
ing of the growing season, and if dry weather cuts down the yield the 
loss is likely to be neutralized by more or less complete protection 
against the weevils. ; 

These contradictory effects of the same adaptation depending upon 
climatic condition may render necessary a complete differentiation of 
the cotton varieties of wet and dry regions. 

It is not improbable that the Upland varieties previously known in 
the United States came originally from the more or less arid regions 
of Mexico, where absence or very small development of the basal 
branches keeps the ground from being constantly shaded and gives 
better chances for the weevils to be killed by the drying out of the 
fallen squares. 

Our Upland cottons are undoubtedly of American origin, but the 
region from which they came has not been ascertained. Some of the 
Texas varieties are said to have been brought from Mexico. Coro- 
nado’s Journal of the earliest Spanish exploration in Arizona and 
New Mexico contains many references to the cultivation of cotton by 
the Indians. There can be little doubt that the agricultural Indians 
of the Gulf region also cultivated cotton, though no documentary 
evidence of the fact seems to have come to hght as yet. 

It is highly probable that the original home of the cotton plant, and 
of the boll weevil as well, was in a somewhat arid region, since it is 
only under such conditions that the weevil would be effectually pre- 
vented from increasing to the fatal degree of destroying its host 
plant, and thus cutting off its only means of subsistence. On the 
other hand, it was only in a humid country like eastern Guatemala 
that many of these weevil-resisting adaptations would be likely to 
develop if, as now appears, it has required the selective influence of 
the boll weevil itself to bring them to their present advanced develop- 
ment. 


a Tiunter, W. D., and Hinds, W. FE., 1904. The Mexican Cotton Boll Weevil, 


Bul. 45, Division of Entomology, U. S. Department of Agriculture, pp. 78 and 74. 


COUNTINGS OF FLARED AND FALLEN SQUARES. 45 


The adaptive character of this habit of shedding the parasitized 
squares seems to be confirmed by the fact that it depends upon the 
existence of a special layer of soft cells which readily break down 
when the bud is injured. Many plants have such cells as a means of 
shedding their fruits, but they seem not to be prevalent among the 
relatives of the cotton. The cotton itself does not drop the ripe bolls, 
and even the empty shell often remains long after the seeds are gone. 

The drier the climate the more effective is the prompt shedding of 
injured squares. Whether there are other adaptations thus especially 
suited to dry climates 1s not yet known, our studies having been con- 
fined mostly to humid regions. 

Dr. Edward Palmer, who has spent many years in botanical ex- 
plorations of the dry plateau region of Mexico and who discovered 
that the boll weevil was a cotton pest, states that in several localities 
where the cotton was formerly grown without difficulty the introduc- 
tion of irrigation improvements has proved disastrous. With the 
assistance of the moist soil the weevils are now able to reach maturity 
in large numbers and complete the devastation-of the crop, quite as 
in Texas. The irrigated soil affords a situation favorable for the 
development of the larvee in the fallen squares. 

This is said to have been the case about Parras, and at Rio Verde, 
below San Luis Potosi. The culture of cotton has declined also in 
the “Huasteca Potosina,” the tropical district between San Luis and 
Tampico, and on the Pacific side of Mexico, along the Santiago River 
above San Blas, as well as about Tepic. Doctor Palmer saw cotton 
growing in a wild condition in the fences at the old mission, San José 
de Guaymas, 6 miles from the commercial port; again at Mulege, 
Lower California, across the Gulf from Guaymas, the latter a much- 
branched, prolific tree, producing a nankeen-colored lint. About 
Guaymas cotton was formerly utilized by the Indians as tinder, after 
being dipped in a solution of saltpeter. The same facts were observed 
by Dr. L. O. Howard in 1899 at San José de Guaymas. 


COUNTINGS OF FLARED AND FALLEN SQUARES. 


An attempt was made in connection with our Guatemalan experi- 
ment to secure data on which a definite statement might be based 
regarding the extent to which the different varieties were protected 
by their involucral characters, but the problems are too complex to 
be reached except by more elaborate statistical studies than were prac- 
ticable at that time. 

Countings were made, for example, of the flared and _ fallen 
squares—that is, of those which it might be supposed that the weevils 
have injured—and of the number of weevil larvie, proliferations, etc., 
found inside them. The results in percentages do not agree, however, 


46 WEEVIL-RESISTING ADAPTATIONS OF COTTON. 


with the facts obvious in the fields; indeed, they greatly misrepre- 
sent them. Thus the percentage of weevil injuries in flared and 
fallen squares does not appear very much higher in the Kekchi cotton 
than in the Sea Island and Upland varieties; yet as a matter of 
fact the squares of the Kekchi cotton seldom flared for any other 
reason than weevil injuries, and much less often for this cause than 
did those of other varieties. Many small squares of the Kekehi cot- 
ton fall off, however, before they are large enough or open enough 
to be attacked by the weevils.*| This takes place in the other varie- 
ties to a much smaller extent, but with them the apparent percentage 
of weevil injuries among flared squares is much diminished, because 
many squares stand open and appear as though beginning to flare, 
even before the weevils have attacked them. 


PROLIFERATION OF INTERNAL TISSUES. OF BUDS. 


The protection of the buds does not end with devices for the exclu- 
sion of the adult weevils, nor with the rejection of those in which they 
have laid their eggs. It is also possible for the plant to heal the 
wound, and bring the injured bud to maturity by preventing the 
growth of the weevil larva. Where the climate is dry the weevil 
larvee in the rejected buds are killed, as already explained. The 
humid climate alternative of the falling of the parasitized squares is 
proliferation, the growth inside the bud of loose, watery tissue in 
which the larva does: not develop. Whether the larva is killed by 
smothering, starving, or poisoning, or by some combination of these, 
is not yet known. Starvation is a sufficient explanation, since the 
material with which the larva becomes surrounded can be no adequate 
substitute for the highly nutritious pollen grains on which the infant 
larva would otherwise feed. 

Proliferation is much more frequent in the Kekchi cotton than in 
any of our United States varieties, as far as known. ‘The first and 
second punctures are commonly resisted successfully, but the third, 
fourth, or fifth attempt may succeed in the development of a larva. 
The proportion of weevil punctures rendered ineffective by prolifera- 
tion was found to run well above 50 per cent, sometimes between 80 
and 90. (PI. V.) 

The promptness and efficiency of proliferation bear an inverse pro- 
portion to the size of the buds. As the latter grow larger the mass of 
anthers inside becomes less compact, and the other tissues become too 


aProfessor Pittier found in the latter part of the season that the buds of 
the Kekchi cotton were sometimes cut away at the base and left hanging in a 
wilted condition. These were at first taken for flared squares as the result 
of weevil injuries, but it was later ascertained that this was not the case, 
though the true cause was not learned. The damage was done in the night, 


PROLIFERATION OF INTERNAL TISSUES OF BUDS. 47 


nearly mature to put forth new growth. If the presence of the larva 
ut this stage is sufficient to cause the bud to fall off, the development 
of the parasite to maturity is well assured, the large bud affording 
good protection and adequate food. 

In the Kekchi cotton, however, such late attacks very. seldom cause 
the bud to fall off. Larvee developed in the larger buds are turned 
out of doors, as it were, by the opening of the flower. The tendency 
of injured buds to persist is notably greater than in the United 
States, either because of some physiological difference between the 
varieties, or because of the larger and more firmly closed involucres 
of the Kekchi cotton, which keep the buds surrounded with a moist 
atmosphere and protect it against drying out while the new tissues 
are forming to heal the wound and encyst the egg. 

In the closely planted Indian fields the squares seldom flare as in 
the Texas varieties. They generally remain in place and continue to 
grow until the bracts have reached nearly their full normal size. In 
fields partially protected by the keleps the weevil larve do not 
seem to develop in buds as small as in Texas. Proliferation may 
partly explain this delay and also the more firmly closed involucres, 
but in our unprotected plot the weevils were able by repeated punc- 
tures to infest smaller squares and reach maturity in them, after they 
had fallen to the ground. 

The behavior of weevil larve inside the squares in Guatemala 
seems also to differ appreciably from that observed in Texas where 
younger squares are usually much more accessible to the weevils, and 
are commonly punctured. In Texas the larve regularly grow to 
maturity, depending for food upon the pollen, which is completely 
eaten out. In Guatemala this very seldom occurs. Small squares 
with well-developed weevil larve are rarely found under normal con- 
ditions, nor do the larvee depend upon the pollen as their principal 
article of diet, as jn Texas. 

Several reasons for this difference may be considered. The first is 
that the larger and more firmly closed involucre of the Kekchi cotton 
gives the buds several days of protection, so that the average size 
would naturally be larger. The examination of large numbers of 
squares picked at random from the Indian cotton fields by Messrs. 
Kinsler and McLachlan show also that a very large proportion of the 
punctures are followed by proliferation, and that this means of pro- 
tection is much more efficient in the younger squares. Another rea- 
son must be sought, however, for the failure of the larva to eat the 
pollen of the large buds where proliferation is less prompt and less 
frequent. The impression might be gained that the pollen of the 
Kekchi cotton is in some way not acceptable to the weevils, since even 
when there is an abundance of pollen at hand they prefer to eat out 


45 WEEVIL-RESISTING ADAPTATIONS OF COTTON. 


the style and central column of the flower, and thence down into the 
ovary or young boll. After this has been consumed the larvee return 
to the upper part of the bud to finish the remainder of the pollen. 
Nevertheless, this suggestion of a protecting quality in the pollen 
itself can not be accepted with much confidence because the weevils 
showed in numerous instances that they could live and thrive upon the 
pollen of the young squares, quite as in the United States. This oc- 
curred in the experimental plot where there were no keleps, and the 
weevils were very numerous and persistent in their attacks. After 
two or three punctures the squares flared and fell to the ground in 
the usual manner, and in these the weevil karve were able to reach 
maturity. 
A more probable reason for the usual failure of the larvee to eat the 
pollen as freely as in the United States is furnished by the opinion of 
Mr. W. D. Hunter, that the original habit of the weevil was to attack 
the bolls, like related species of Anthonomus, which live upon various 
kinds of fruits.*. If this be true with reference to the boll weevil we 
may think of the Guatemalan members of the species as having 
retained somewhat more of the ancestral habits which with them are 
definitely useful, because the cotton variety with which they have to 
deal has perfected, to 2 larger extent than the Texas varieties, the art 
of proliferation. 
As a further indication of the greater strength among the Guate- 
malan weevils of the instinct of attacking the ovary of the bud 
may be mentioned the fact that a very large proportion of the 
punctures occur low down—that is, on or below the level of the apex 
of the young boll. The larva commonly eats directly to the center of 
“the bud and hollows out the apex of the young boll. This habit 
gives rather less opportunity for successful proliferation than in 
Texas, because the cavity hollowed out by the larva lies below the 

fevel of the staminal tube, the tissues of which are the most active 
in proliferation. The Kekchi cotton shows occasionally another 
dorm of proliferation not recorded from Texas, namely, that of the 
ise of the corolla. Sometimes this enlargement takes place in an 
outward direction, forming a wart or protuberance on one side 
of the bud, as shown in Plate VI. In other instances the direc- 
tion is reversed and the ingrowing edges of the wound made by 
the weevil fill the internal cavity and prevent the development 
of the larva. The proliferation of the corolla, besides being less 

aA new species of Anthonomus with habits closely identical with those of the 
boll weevil, but parasitic on the pepper plant (Capsicum), has been discovered 
recently in Texas by Mr. BE. A. Schwarz. This gains an added interest from the 
fact already noted that it is the regular custom of the Indians of Alta Vera 
I’az to plant peppers among the cotton. 


CAUSES AND CONDITIONS OF BUD PROLIFERATION. 49 


frequent than that of the staminal tube, is probably also less effect- 
ive, since the weevil larve could escape before it into the center of 
the flower while the proliferation from the staminal tube grows 
outward, as though to meet the intruder and keep him separated 
from the more special organs. 

The habit of the larve to seek the center of the bud and gnaw 
off the style is responsible for the loss of large numbers of younger 
bolls which have suffered no direct injury from the weevil. Even 
though the larva be subsequently killed by proliferation or though 
the flower drops off and carries the larva with it, the lack of polli- 
nation must prevent the development of the young boll unless par- 
thenogenesis takes place, which seems improbable. 

Larve were found in several instances in nearly full-sized buds 
about to open, and in another case a more than half-grown larva 
was found inside the central column of an open flower. More or less 
distorted flowers with unmistakable signs of previous proliferation 
in the bud stages are commonly found in the Kekchi cotton fields. 

Summarizing the results of the study of proliferation in the 
Kekchi cotton, it may be said that although the frequency of pro- 
liferation in the young squares is very great, its efficiency in prevent- 
ing the breeding of the weevils is somewhat less than might be ex- 
pected in Texas, owing to the difference of food habits among the 
weevils. If the Texas weevils are as consistent in their habits as 
now supposed, the introduction of the Kekchi cotton or of a similar 
proliferating variety might be of great benefit as a preventive 
measure. The extent, however, to which it could be made to compass 
the complete destruction of the weevil would depend somewhat upon 
the degree, if any, to which they might return to the habit shown in 
Guatemala of feeding upon the ovaries or boll rudiments rather 
than upon the pollen of the young buds, an important and hitherto 
unsuspected difference in habits between the weevils of Texas and 
those of Guatemala. 


CAUSES AND CONDITIONS OF BUD PROLIFERATION. 


That the proliferation is occasioned by the injuries of the weevil is 
too obvious to admit of doubt, but it may be of much practical 
importance to learn the exact way in which the new growth of tissue 
is brought about. The disturbing: factor might be either mechanical 
or chemical. The new growth may be a direct response to injury of 
the weevils in feeding or laying eggs, or it might be stimulated indi- 
rectly by the secretions of the young larva, or by chemical changes 
or decay of the damaged tissue. A second mechanical possibility is 
that of pressure developed in the young and rapidly growing bud. 


9962—No. 88S—05 m——4 


50 WEEVIL-RESISTING ADAPTATIONS OF COTTON. 

The burrowing of the weevil relieves this pressure at one point, and 
may thus furnish the exciting cause of the rapid growth in this diree- 
tion of the tissue of the staminal tube. , 

It seems not improbable that a relation will be found between the 
method of culture and the extent and frequency of proliferation. 
Open-field conditions, with much bare ground about the plants, would 
increase the daily exposure of heat and dry air, and this would con- 
duce to the wilting of the punctured squares, which might then be 
expected to flare and fall off instead of remaining to proliferate. The 
result of weevil work tn our open-culture plots was obviously differ- 
ent from that in the more crowded cotton fields of the Indians. On 
the widely separated plants the squares often fell off and permitted 
the larvae to develop, as in Texas, except that there was still a distinct 
tendency on the part of the larve to attack the pistil and ovary first, 
before eating out the pollen. 


PROLIFERATION IN OTHER VARIETIES. 


Proliferation is by no means confined to the Kekchi cotton, but 
probably occurs, occasionally at least, in all the Upland and Sea 
Island varieties. A noteworthy Guatemalan Sea Island cotton was 
found by Mr. Kinsler in the aldea of San Lucas, a few miles from 
Secanquim.* Both the buds and the bolls afforded fine examples of 
effective proliferation. Even the Egyptian varieties showed a dis- 
tinct ability in this direction. In one instance no less than 17 of 23 
punctured squares of Jannovitch had proliferated, and 15 cases 
seemed to have been effective. 

Proliferation ceases to occur when the bud has become too large. 
The anthers are no longer so closely packed together and the tissues 
of the staminal tube are too nearly mature. By that time, however, 
the style may be sufficiently developed to furnish adequate food. 
It is well known, however, that the period of development of the 
weevil larve may be greatly prolonged, and this would seem likely 
in the present instance, since the tissues of the styles must be less 
nutritious than the pollen. The delay also would be advantageous, 
since it would permit the young boll to become larger. 


a This variety is peculiar in having about half of each seed covered only with a 
very fine, short, bright bluish-green lint. The upper half bears the long white 
fiber, and is smooth and black when this has been removed. Some of the 
plants had excellent crops of bolls, unusually uniform in size and apparent age, 
as though the habit of seasonal flowering were well accentuated. The variety 
is evidently perennial and grows to a height of from 6 to 8 feet, but on the 
other plants the leaves, flowers, and bolls were much reduced in size. The 
plants were all occupied by small black ants. On some of them no weevils nor 
any indications of weevil injury were found, but others only a few rods away 
were badly infested, 


PROTECTION OF THE BOLLS. 5 


But as the power of effective proliferation declines in the larger 
buds another factor of protection comes into play. The later the 
attack of the weevil the greater is the chance that the bud will mature 
and the flower will open and turn the weevil larva out of its quarters 
to die. And since buds commonly mature which have been attacked 
while still young enough to proliferate, it is easy to understand why 
attacks made in the later stages seem to be effective only in excep- 
tional instances. 

An element of uncertainty often attaches to the enumeration of 
weevil injuries because of the difficulty of finding the egg or very 
young larve of the weevil in the squares which have been only 
recently attacked. This is especially true in small squares where the 
anthers are still white and of about the same color, size, and general 
appearance as the eggs. The possible error does not, however, mate- 
rially affect the result, since it is to be expected that the same propor- 
dion of bolls will proliferate and the same percentage of weevil 
larve develop as in the squares which are far enough advanced to 
show definite results. 


PROTECTION OF THE BOLLS. 


If it be true, as already intimated, that the original habit of the 
weevil was to attack the boll instead of the bud, the opportunity for 
the selective development of protective characters of the boll has 
been greater. This suggestion seenis to accord with the results, 
since the boll of the Kekchi cotton has a series of protective characters 
even more striking and effective than those of the involucre and the 
bud. 


PERSISTENCE OF FLOWERS. 


As long as the flower remains in place the young boll is thoroughly 
protected, the weevils having no means of access except by boring 
through the withering tissues, which seems not to be attempted. In 
the Kekchi cotton the flower falls only when detached by the swelling 
of the young boll. This may also be true of other varieties. (See 
Pl X. ) 

The frequent sequel of proliferation in the bud, as noted above, is 
the loss of the young boll through lack of pollination. This is espe- 
cially true in Guatemala, owing to the tendency of the weevil larvee 
to eat away the style. On one occasion Mr. Kinsler collected from a 
field of Indian cotton 28 young bolls showing signs of debility. These 
measured from 13 to 20 mm. in length, most of them about 15 mm. 
None of the smaller bolls showed signs of weevil injury, but in many 
of them the ovules were already shriveling up. A few punctures were 
found in some of the larger bolls, and in some of these proliferation 
had occurred. The development of the weevil larve to maturity 


OZ WEEVIL-RESISTING ADAPTATIONS OF COTTON. 


seemed unlikely in any case, because the unfertilized ovules were 
already withering. 

Presumably there are various stages and degrees of fertilization. 
Some of the stigmas of proliferated buds seem to have adequate 
pollen, so that the bolls can develop normally, while others obtain 
none at all or only a little. The persistence of injured flowers is 
much greater. They may not fall off at all, and often remain at- 
tached by the withered style to the boll when nearly full size. 

It thus happens that injured flowers protect their young bolls 
longer than the others, but in most instances such bolls remain small 
or unsymmetrical, presumably as a result of inadequate fertilization. 
It is quite possible, however, for normal bolls to develop occasionally 
from weevil-infested buds which never open, for the style often 
pushes through and becomes fully exposed, so that fertilization by 
pollen from another flower might readily take place. 


IMMUNITY OF VERY YOUNG BOLLS. 


For reasons not yet ascertained, the weevils in Guatemala seldom 
or never attacked the very young bolls. This may be due to a con- 
servative instinct on the part of the weevil, like that which forbids 
the laying of any additional eggs in a bud already parasitized.* It 
is not impossible, however, that the oil glands with which the sur- 
face of the young boll is very thickly beset may have a protective 
function. As the boll grows larger the glands do not appear to 
increase in numbers, but become separated much more widely. On 
bolls of the Kekchi cotton the oil glands are usually absent from a 
distinct longitudinal band running down the middle of each carpel. 
(Pl. VII.) A large proportion of the weevil egg punctures are 
made along this naked band, although: very few of them take effect. 
The wall is thicker here, and the weevil in boring meets the tough 
lining of the boll chamber at an angle, and is seldom able to penetrate. 
If this interpretation of the facts be correct, the naked band consti- 
tutes a veritable weevil trap, a device for inducing the weevil to 
make its punctures and lay its eggs in the part of the boll where they 
can do no harm.” 

To ascribe a protective value to the oil glands is not unreason- 
able in view of the fact reported by Messrs. Quaintance and Brues, 


«Hunter, W. D., and Hinds, W. E., 1905. The Mexican Cotton Boll Weevil, 
Bul. 51, Bureau of Entomology, U. S. Department of Agriculture, p. 78. 

’This peculiarity of a glandless longitudinal band in the middle of each 
carpel was also noticed in a variety of cotton cultivated by the Moqui Indians 
of Arizona, grown in 1904, in the Department’s plant-breeding experimental 
field at Terrell, Tex. The Moqui cotton is interesting also by reason of its | 
short, squarish, distinetly apiculate bolls, more like some of the Old World 
cottons than are those of other members of the Upland series. 


IMMUNITY OF VERY YOUNG BOLLS. 5a 


that the Egyptian cotton, the bolls of which are excessively oily, is 
on this account immune from the bollworm.* The oi] contained in 
the glands has a deep-brown color, a sticky, molasses-like consistence, 
a disagreeable, pungent odor, and a sharp, resinous taste, suggesting 
turpentine or Canada balsam. 

The development of the oil glands seems to be especially great in 
the Egyptian variety known as Mit Afifi, and the glands are more 
superficial. By slight pressure, or by drawing the nail across the sur- 
face, the oily liquid is freely obtained. Most of the Upland varieties 
have the oil glands much more scattering and deep set than the Egyp- 
tian sorts, and it is not possible to. squeeze the resin out of them in 
any such manner. 

On Redshank and other Upland types the resin glands are marked 
by shght superficial depressions, but a cross section shows them to be 
well below the surface, with several layers of chlorophyll-bearing 
cells between. On the Egyptian sorts the glands are also set in de- 
pressions, but the gland itself is very close to the surface, and makes 
the bottom of the depression again convex, the superficial layer of 
cells being very thin. It seems to break spontaneously in some in- 
stances; at least there are frequently small spots of hardened resin, 
and very slight pressure brings out the dark, gummy fluid. The 
fingers receive a permanent brownish stain, which with the acrid, 
biting sensation experienced when the liquid is applied to the tongue, 
increases the probability that substances of a definitely protective 
character are present. It is well known that many of the aromatic 
oils are for some reason highly distasteful or even fatal to many 
insects. 

The Sea Island and Kidney cottons have the oil glands conspicu- 
ously developed, like the Egyptian varieties, but the Old World 
cotton (Gossypium herbaceum) is in this, as well as in other respects, 
more nearly related to the American Upland cotton (Gossypium hir- 
sutum). The Aidin (Asia Minor) variety of Gossypium herbaceum 
has the oil glands rather small and deep set, with the superficial pits 
rather shallow, more so than the Ceylon or Korean types. 

Even the petals of the Guatemalan Kidney cotton found at Trece 
Aguas” contained oil glands. The color of the petals was a uniform 
pale yellow, without purple spots on the inside, but in the upper 


a@Quaintance, A. L., and Brues, C. T., 1905. The Cotton Bollworm, Bul. 50, 
Bureau of Entomology, U. S. Department of Agriculture, p. 71. 

+ The Kidney cotton at Trece Aguas is called paiyi, and seems to have little or 
no relation in the minds of the Indians with the dwarf Upland cotton, which is 
called nok. In the Secanquim district, only a few miles away, this name paiyi 
(pronounced like the English words pie ye) is not recognized. Kidney cotton, 
though apparently not now planted by the Indians, is not entirely unknown to 
them. They call it simply che nok, or tree cotton. 


54 WEEVIL-RESISTING ADAPTATIONS OF COTTON. 

half specked with minute brown glandular dots.*. The oil glands of 
the bolls of this Kidney cotton are apparently quite as strongly devel- 
oped as in the Egyptian varieties, or even more so. They are distrib- 
uted very irregularly over the surface, and are not lacking above the 
dissepiments, along the middle of the carpels. The position and 
structure of the glands seem also to be the same as in the Egyptian 
cottons. They are close to the surface and show as distinct black 
spots, there being no green tissues over them as in the Upland and 
herbaceum types. 

I am indebted to Mr. Guy N. Collins for the suggestion that the 
present inefficiency of the oil glands as a means of protecting the 
cotton from the boll weevil furnishes no argument against the 
adaptation of the glands nor their development through the selective 
agencies of the boll weevil itself. This fact is sufficiently obvious 
when once stated, but it is not commonly taken into account in con- 
sidering questions of this kind. We may be sure that the gradual 
development of a protective character like the oil gland would carry 
with it a corresponding increase in the power of the weevil to avoid 
or to endure the injury. The ultimate value of the device would de- 
pend on whether the glands were able to keep ahead of the weevils 
in quantity and distastefulness. The readiness with which the boll 
weevils attack the Egyptian cotton renders it obvious that oil is now 
no adequate protection, but the preference of the weevils for the un- 
protected strips of the bolls of the Kekchi cotton indicates that the 
weevils still dislike the oil, though they may have foiled the attempt 
of the plant to protect itself m this way. 

There are two attendant facts which under certain circumstances 
might readily obscure the immunity of the young bolls. Many such 
small bolls fall off, a particularly large number it seemed from our 
row of Parker cotton, but an examination of these failed to show 
anything in the way of weevil injuries, except such as had been in- 
flicted while the bud or flower was still in place, the style and a 
small apical cavity having been eaten away in numerous instances. 
Many small bolls were to all appearances quite uninjured. They 
may have been rejected by the plant as supernumerary, the plant 
being unable to furnish the food material needed to bring them to 
maturity, or they may have failed of fertilization as a result of 
weevil injuries to the bud or from other causes, such as the absence 
of bees, which were extremely scarce in the Guatemalan cotton fields. 
The frequency with which the boll weevils were found inside the 

a'The flowers of the Kekchi cotton are pure creamy white when young and as 
long as they remain open. When old and rolled together they-become a pinkish 
red. They are not yellow or bluish at any stage. The stamens and pistils are 
also nearly white, the latter with rows cf oil glands showing as small grayish 
dots. 


RAPID GROWTH OF YOUNG BOLLS. 55 


cotton flowers and well dusted over with pollen suggests the possi- 
bility that in this district at least they were a not unimportant 
agency of cross-fertilization. The performance of such a service by 
the boll weevil would be comparable to the famous case of the yucca 
and its moth, the plant being dependent for cross-fertilization upon 
its insect parasite. The weevils eat the pollen from the bud: that 
they visit the flowers for the same purpose seems highly probable. 
The investigations of Messrs. Hunter and Hinds have shown, indeed, 
that a pollen diet is a necessity for the complete sexual maturity and 
reproduction of the weevils; if without buds to feed upon they 
seldom copulated and never laid eggs.’ 


RAPID GROWTH OF YOUNG BOLLS. 


Mr. John H. Kinsler, who gave careful attention to the earlier 
stages of the Guatemalan experiment, gained an impression that the 
young bolls of the Kekchi cotton increased in size with a rapidity 
distinctly greater than that of the United States Upland varieties 
planted alongside. It was not practicable to establish the fact by 
carrying out a series of daily measurements, though it was possible 
to ascertain from dated tags used in connection with the hybridiza- 
tion experiments that the Kekchi cotton can grow bolls to full 
size in less than a month from the time the flower opens. Plate IX, 
figure 2, shows on the right two bolls of Kekchi cotton less than a 
month from flowering. On the left are the two largest bolls from an 
adjoining plant of King, the seed of both varieties having been sown 
the same day. 

Such an acceleration of the growth would be of very obvious utility 
in lessening the period in which the danger of infestation is greatest. 
A large proportion of the weevils found in adult bolls of Kekchi cot- 
ton were in “ locks” or compartments of diminutive size, showing 
that the infestation had taken place while the boll was less than half 
grown. Indeed, the weevils seldom seem to be able to affect lodg- 
ment in bolls more than half grown, although numerous attempts 
are made in fields where the weevils are numerous. The following 
field note describes such an instance: 

A boll showing many external marks of weevil punctures was found on being 
cut up with care to have been attacked at least fourteen times. In five cases 
the outer wall seemed not to have been penetrated, but in nine others there had 


been complete perforations. All of these had been closed, however, by prolifer- 
ation from the inner surface, and no living larvze were found. 


Such persistent attacks, however, may finally induce a diseased 
condition which interferes with the normal growth of the boll, even 


« Funter, W. D., and Hinds, W. E., 1905. The Mexican Cotton Boll Weevil, 
Bul. 51, Bureau of Entomology, U. S. Department of Agriculture, p. 113. 


56 WEEVIL-RESISTING ADAPTATIONS OF COTTON. 


though the weevils be successfully resisted. Such injured bolls often 
show a brownish discoloration of the interior tissues near the base 
and connecting with the nectaries, which may indicate a bacterial 
disease, to be discussed later. Sometimes this affects the walls only, 
sometimes one or more seeds and the surrounding lint. 


THICK-WALLED BOLLS. 


In the Kekchi cotton there are considerable variations in the thick- 
ness of the outer wall of the boll. Not infrequently the wall equals 
or exceeds the length of a weevil’s snout, so that only the largest or 
iongest snouted weevils would be able to make an opening into the 
interior cavity. It was noted, also, that on the inside such bolls are 
often quite free from these injuries or small larve, though numerous 
attempts may have been made. Large larve or pupz may be found, 
but these have come, obviously, from eggs laid while the boll was still 
young. On some plants the development of large thick walls takes 
place very promptly, so that a protective character of considerable 
value might be obtained if this feature could be increased and ren- 
dered constant. Early development of the thick walls was indicated 
by the fact that the young seeds and lint did not fill the cavity, and 
the seeds were still far from mature. Instances might be drawn 
from other plants where the growth of the pod or seed vessels far 
outruns the seeds at first, so that the development of such a character 
in cotton might reasonably be expected. 

Even when a wall thicker than usual has been bored through, the 
egg must be laid on the outside of the mass of lint which still inter- 
venes between it and the young seed, so that the larva’s chances of 
development are greatly lessened. As will be shown later in the dis- 
cussion of proliferation in the bolls, the instances are very numerous 
in which, although the wall is penetrated, no further damage results; 
either the egg is not laid or the development of the larva is pre- 
vented by proliferation. In any event the boll escapes further 
injury, and it is a very significant fact that in the dissection of a large 
number of such bolls of Kekchi cotton scarcely any young larve 
were found, in spite of the fact that most of them had been punctured 
not once only, but many times. 


TOUGH LININGS OF CHAMBERS OF BOLLS. 


The three, four, or five chambers which contain the locks of cotton 
in the unopened boll have each a complete membranous lining. In 
the Kekchi cotton, at least, this is extremely tough and parchment- 
like, even in bolls not yet full grown and in which the seeds are not 
yet fully formed. This membrane is readily separable from the 
more fleshy external layers of the boll, and though flexible, it is very 


TOUGH LININGS OF CHAMBERS OF BOLLS. 57 
firm and incompressible, and resists tearing unless considerable 
strength be exerted. 

A large percentage of attempted punctures of the larger bolls 
failed because the weevils are unable to penetrate this protective lin- 
ing. This fact is readily determined by the study of radial sections 
of the outer wall through the warts which mark the weevils’ points 
of attack. The different texture of the new tissue which has closed the 
wound shows, usually, that the cavity eaten out by the weevil extended 
down to the tough basal lining, even when no evidence of the injury 
has become apparent on the inside. In other instances, also very fre- 
quent, the new tissue, developed as a result of the irritation of the 
attempted puncture, exceeds the cavity and causes an inward swelling 
or prominence of the inner lining analogous to the projecting warts 
which are the usual external indication of weevil punctures. 

It occasionally happens, too, that the projection of the new tissue 
occurs almost entirely in the inside, the external wart being very 
slightly developed or not at all, though the new tissue and the inner 
swelling show that a puncture had been attempted. 

The utility of this lining as a means of excluding the boll weevil 
seems not to have been considered heretofore, and there has been no 
opportunity as yet to compare the Kekchi cotton with other varieties 
with regard to this feature.* Certain it is, however, that in the Kekchi 
cotton the parchment lining is almost as firm and tough as that which 
surrounds an adult coffee seed. And it is certain, also, that a very 
large proportion of the attempted punctures of the bolls failed to 
bore through this inner wall of defense. 

The examination of a large number of bolls, which were full size or 
nearly so, though still far from maturity, in most cases failed to find 
more than a very few instances, if any, of very recent perforation, 
though there were large numbers of instances where the weevils had 
gnawed their way down through the parchment and deposited an 
egg. In many such cases the proliferation or new growth induced by 
the injury causes the parchment to be raised up from the wall on the 
inside to form a blister-like, rounded protuberance. (Pl. VIII.) 
Eggs laid outside the parchment are firmly embedded in the new 


a Since this was written Mr. McLachlan has reported the existence of the same 
form of protection in Upland varieties in Texas. The following note describes 
the results of injuries inflicted upon the bolls of a plant of Parker cotton in four 
days from August 8 to August 12, 1905: 

“The 9 larger bolls, when opened, were found to have 28 weevil eggs deposited 
in them; 6 had struck the dissepiment ; 12 were not entirely through the shuck of 
the boll (either not more than half way there or else stuck in the tough inner tis- 
sue of the shuck) ; the others were embedded in the lint. In only two instances 
was there any proliferation apparent. The outer shuck had proliferated at the 
wound and in one case had encysted the egg. The other had merely forced the 
egg to one side, haying begun the development too late.” 


58 WEEVIL-RESISTING ADAPTATIONS OF COTTON. 


growth and do not appear to hatch, or if they do the larva are not 
able to do any damage, since they can not penetrate into the interior 
of the boll. It quite frequently happens that eggs are laid in the 
sinus or groove between the linings of two locks, but without penetrat- 
ing the parchment of either. The tissue is here somewhat looser 
than in other parts of the wall. In a few instances it was observed 
that the larve had hatched, but no case was found which indicated 
that larve hatched outside the parchment lining had been able to 
penetrate to the interior cavity. 


PROLIFERATION FROM THE WALL OF THE BOLL. 


The wall of the boll offers an active form of weevil resistance by 
proliferation, in a manner somewhat analogous to that of the pro- 
liferation of the square. The channel excavated by the weevil is 
closed by the new growth, which continues to push out on. the inner 
surface of the wall in the form of a rounded, blister-lke protuber- 
ance of loose tissue. This surrounds and encysts the weevil egg, and 
prevents its development. A section through the mass of new tissue 
shows the egg embedded in it or pressed against the lint. Prolifera- 
tion often takes place even when the tough lining of the chamber has 
not been penetrated, and then appears as a prominence underneath 
the membrane. 

It has been seen from the preceding paragraph describing the 
thick walls and tough hning that in the Kekchi cotton, at least, 
the weevil is practically excluded from the boll after the boll has 
reached about three-quarters of its full size; but even in its younger 
stages also there is a measure of defense through the formation of 
new tissue as a result of the irritation set up by the weevil’s injuries 
in a manner analogous to that which induces the formation of galls 
and other vegetable excrescences. 

The first result of the proliferation is to fill up and heal the 
wound bored out by the weevil. The cavity is not only completely 
filled, but in most cases a wartlike prominence is formed on the out- 
side, and if the parchment lining or the inner wall has been pene- 
trated the new proliferating tissue also grows through on the inside 
and often spreads out as a biscuit or button shaped protuberance 
of soft white or transparent tissue several millimeters in diameter 
and readily visible to the naked eye. (PI. VIII.) 

There are two alternatives in the fate of an egg destroyed by 
proliferation. Either it is completely surrounded in the proliferating 
tissue outside or inside of the parchment wall or it is carried on 
the apex of the proliferation down against the lint and flattened 
between the growing surfaces. After the egg has disintegrated 
and disappeared its position is frequently shown by a minute brown 


PROLIFERATION FROM WALL OF BOLL. 5Y 


stain. Such a discoloration often spreads back into the loose tissue 
and then gradually extends over the whole lock of cotton of that 
particular chamber. The seeds fail to develop and finally shrivel up. 

If the proliferation results, as usual, in the death of the weevil 
egg or young larva, the process of abnormal growth ceases with 
the formation of a knob or button of the new tissue on the inside 
of the wall of the boll. When, however, the young weevil escapes 
destruction and continues to eat and grow, the proliferating tissue 
also continues to increase, until in some instances the whole compart- 
ment is filled with a silvery-white cheesy material which seems to 
arise not only from about the original perforation of the outer wall, 
but also from other parts which have been injured and irritated 
by the presence of the weevil larva. This, with other facts already 
stated, seems to show that in some varieties of cotton, at least, the 
tendency to proliferation is very general, or, in other words, con- 
stitutional, which warrants a larger hope of increasing this character 
and making it uniform by selection. 

When proliferation, which results from the presence of the weevil 
larva, has become very extensive and fills the entire compartment, 
the weevil larva is sometimes found to have eaten through the dis- 
sepiment into the next chamber, perhaps to escape starvation. Such 
extensive proliferation, accompanied by the failure of the seeds to 
develop, means, of course, that the weevils gained entrance while the 
boll was still very young. Moreover, if the boll had been older 
there would have been plenty of food for the larva without the 
necessity of entering a second compartment. Finally, the dissepi- 
ment would have been too tough for the larva to penetrate easily. 

Further proof of the fact that the weevil larve are seldom or 
never able to gain a footing in the larger bolls is to be found in the 
fact, already stated, that the weevil larve found in them are nearly 
always in undersized compartments, much smaller than those which 
have remained uninjured, and have thus been able to continue their 
normal development. 

It is to be supposed, perhaps, that if the weevils could gain access 
to large bolls and feed upon the nearly adult seed they would be able 
to develop in less time than they usually spend in reaching maturity 
on the rather poor provender they secure among the abnormal tis- 
sues which arise after they have entered the young bolls. 

The exclusion of the weevil from the large bolls has been evidently 
not only an important measure of protection for the cotton, but it 
has probably compelled the weevil to accustom itself to a gradually 
longer and less prosperous development in the boll. The develop- 
ment of the weevil-resisting adaptations on the part of the cotton 
plant has left the insect aie two opposite alternatives. It must 
enter the boll early and submit to a very long period of development 


60 WEEVIL-RESISTING ADAPTATIONS OF COTTON. 


or enter the square late and develop very promptly. The insect has 
been able, as we know, to avail itself with a large measure of suecess 
of both these alternatives, but it is not without encouragement for 
future progress in weevil resistance to know that the plant has so 
successfully guarded itself in two parts of its life history. 

If additional evidence be needed to show that the food supply 
obtained by the weevil larve in the bolls is very different from that 
in the squares, it is to’be found in the large, firm-walled cells of com- 
pacted excrement with which they surround themselves in the bolls 
before reaching maturity. The food being of a much coarser nature 
and the period of development about three times as long, the amount 
of waste material is naturally very much greater. If feeding upon 
the boll is, as now appears probable, the ancestral habit of the wee- 
vil, it need not surprise us that the protective adaptations of the boll 
are more numerous and effective than those of the bud, which may 
have been attacked by the weevil in comparatively recent times. 


TIME REQUIRED FOR PROLIFERATION. 


In connection with the experiments in Texas, Mr. McLachlan at- 
tempted to ascertain the time required for proliferation to take place 
after the injury had been inflicted. The amount of proliferation 
and the time required for it to develop may be expected to depend 
much on external conditions. Squares of Parker cotton showed no 
development in six hours, but observation on bolls showed that pro- 
liferation was complete in twenty-four hours. Two of Mr. McLach- 
lan’s observations are described in the following notes: 


On August 14, at 9.15 a. m., a wire cage was placed over a plant of King 
cotton, and four weevils, of which at least two were females, were put inside. 
Later, three more were introduced. At the time there were 11 bolls, 3539 squares, 
and 1 flower on the plant. 

On August 17, at 1 p. m., 11 bolls and 18 squares were picked, a little more 
than three days being allowed for the weevils to work. There was no rain, and 
of the 18 squares examined only one revealed proliferated tissue, though the 
weevils had scarred the buds in more than 33 separate places and had deposited 15 
eggs. But the bolls showed better results. They had been scarred at 32 different 
points, and 23 eggs were discovered when the bolls were cut open. In 12 cases 
inward proliferation of the “ shuck”’ had destroyed the eggs. Several of the 
incited growths had caught the egg, encysted it, and carried it along, inclosed 
at the apex, as they pushed their way into the lint. As in the Parker cotton 
examined a short time ago, weevils seem to have some difficulty in getting 
the egg through the shuck of the boll. In dry weather it appears that the 
King cotton is as backward as the Parker in proliferation in the squares, but 
in bolls proliferation goes forward as well in dry as in wet weather. 

On the 30th of August, at 10.15 a. m., a boll (half grown and tender) was 
bagged with a weevil. At 6 p. m. of the same day an egg puncture was found 
on the fruit, but at 8 a. m. of the 31st no further injury had been inflicted. At 
12 m., September 1, four more egg punctures were discovered, and the boll was 


EFFICIENCY OF ADAPTIVE CHARACTERS OF BOLLS. 61 


pulled and examined. The first puncture was then forty-two hours old and ihe 
other four some twenty-four hours old. The examination revealed marked pro- 
liferation in every case, with no greater growth in that of forty-two hours’ 
duration than there was in that of twenty-four. Eggs had been laid inside the 
wall of the boll, since it was easy, in the case of young, tender fruit, for the 
weevil to cut an opening to the lint. But every one of the five eggs had been 
encysted by the proliferated tissue. It is quite possible that one or two of the 
punctures reckoned as twenty-four hours old were still more recent. 


EFFICIENCY OF ADAPTIVE CHARACTERS OF BOLLS. 


The amount of protection afforded in Guatemala by the weevil- 
resisting characters of the bolls might be greatly underestimated if 
it were to be supposed that the weevils make numerons attacks upon 
the bolls for the purpose of feeding upon them. 

In their accounts of the habits of the boll weevil in Texas, Messrs. 
Hunter and Hinds have devoted a chapter to * effects of feeding upon 
squares and bolls,”* but in Guatemala no indications were found that 
weevils punctured the larger bolls for any other purpose than egg 
laying. It is true that the outer surfaces of bolls are frequently 
marked with scars of weevil punctures from which no larve have 
developed and no internal injuries have resulted, but these failures 
can be explained in other ways than by the supposition that the wee- 
vils feed upon the tough and innutritious outer walls of the bolls. 
In Guatemala, at least, it appears that the weevil scars on large bolls 
mark attempts at egg laying, though for a variety of reasons already 
recited most of them are not effective. The only instance where wee- 
vils were found feeding in bolls in Guatemala was at Rabinal. Two 
weevils were together attacking a small boll, and had eaten out large 
superficial pits, quite unlike the punctures in which eggs are laid. 

Feeding punctures in bolls are referred to by Mr. McLachlan in a 
note dated at Victoria, Tex., August 31, 1905. Such injuries were 
not found, however, to lead to the formation of external warts which 
could be mistaken for egg punctures, doubtless for the reason which 
Mr. McLachlan gives: 

It has been noticed that in bolls no proliferation occurs following the injury 
from a feeding puncture, however serious that may be. Furthermore, from the 
above and other observations it is apparent that proliferation is not excited by 
the egg puncture or the egg, unless the puncture extends through the inside 
tissue and the egg is fixed in the tissue or has been pushed through it to the 
lint. In that case a dense knob of proliferation occurs on the inner side of the 
shuck, in the center of which the egg is often encysted. There must be a con- 
stant irritant like the egg, with an opening to give it access to the lint, in order 
to occasion the specialized growth. As a suggestion it might be noted that all 


the egg punctures are sealed by the adult weevil at the time of egg laying, 
while the feeding punctures are left open. 


“Vfunter, W. D., and Hinds, W. E.. 1905. The Mexican Cotton Boll Weevil. 
Bul. 51, Bureau of Entomology, U. S. Department of Agriculture, p. 59, Pl. VIII. 


62 WEEVIL-RESISTING ADAPTATIONS OF COTTON. 


The feeding experiment reported by Messrs. Hunter and Hinds “4 
shows that weevils fed exclusively upon bolls lived less than twenty 
days, while those fed upon the squares lived nearly seventy days. 
The bolls proved to be much less suitable for food than the leaves, on 
which the weevils were able to prolong life for thirty days and 
upward, though no eggs were laid on a leaf diet. It may be that in 
Texas, where the army worms sometimes destroy all the leaves, the 
weevils might be driven to gnawing the bolls for food, but in Guate- 
mala the plants remain in full leaf throughout the growing season. 


BACTERIAL DISEASES FOLLOWING WEEVIL INJURIES. 


In the study of the bolls of the Kekchi cotton three diseased con- 
ditions were observed, some or all of which may be of bacterial 
origin, the bacteria having been introduced, perhaps, by the weevils 
at the time of egg laying. None of these diseased conditions is fre- 
quent, and as they do not permit the fruit to reach normal maturity 
it seems very unlikely that they can be introduced into the United 
States with the seeds. It may be stated in addition that the seed 
obtained by Mr. Kinsler in the season of 1905 has been carefully 
selected in the field and comes from the earliest and most vigorous 
bolls. 

The first of the diseased conditions consists in a aide deliquescence 
of the immature seeds and lint as though the lock had been dipped in 
milk. There is also a distinct odor of fermentation. Another dis- 
ease turns the seed and lint brown. Though observed only in bolls 
which have been punctured by the weevil, there was often an 
apparent connection between the disease inside and the large extra- 
floral nectaries. A column of transparent or somewhat discolored 
{issue extends from each nectary obliquely upward to the cavity of 
the boll. This may be a symptom of the disease or it may indicate 
that bacteria find their way into the bolls by way of the nectaries. 

The third abnormal condition was also indicated by a brown dis- 
coloration of the wall and contents of the affected compartment of 
the boll. The seeds and lint soon die and shrivel. No special indi- 
‘ation of bacterial activity was noted, and it may be that the death 
of the weevil egg or larva has some prejudicial effect upon the sur- 
rounding cells, as suggested by the brown discoloration already 
noted in describing the effects of proliferation. Such a disturbance 
inight continue to spread and thus cause the death of the young seeds. 


BREEDING IN BUDS A DERIVED HABIT. 


The fact that the weevil larve are found in the young buds of the 
cotton plant and also in the full-grown bolls has been taken to mean 
that it affects all the intervening stages as well. This would imply 


4 Hunter, W. D., and Hinds, W. E., |. ¢., pp. 34-385. 


BREEDING IN BUDS A DERIVED HABIT. 63 


also that if the weevil fed originally upon the bolls it has followed 
back to earlier and earlier stages and finally to the bud. The facts 
already detailed seem to prove, however, that this is not the case. 
The weevil does not attack the very young bolls, nor does it operate 
while the flower is open or while it remains in place, though in a 
withered condition. The hatching of the weevil larva in the large 
buds is likewise ineffective because the larva is deprived of shelter 
when the flower opens. It seems necessary to believe, therefore, that 
the parasitism of the weevils upon the buds of the cotton is a habit 
quite distinct from that of its relations to the boll. The habit of 
breeding in the bud marked a new departure in the biological history 
of the insect and not a gradual change from the previous habit of 
infesting the bolls only. Nevertheless, the change of habits need not 
be thought of as anything very remarkable from the standpoint of the 
insect. A cotton bud is very much larger than a small boll. The 
peculiarity hes in the plant rather than in the insect, since very few 
plants afford a continuous and abundant succession of large, pollen- 
tilled buds.. It is this quality of the cotton plant which has enabled 
the weevil to develop its peculiar and highly destructive secondary 
habits of feeding upon the buds and using them as breeding places. 
If the boll weevil were restricted, like related beetles, to parasitism 
upon the fruit of the cotton, it would have remained a comparatively 
harmless and agriculturally insignificant enemy. These considera- 
tions may assist in a better appreciation of the extent to which the 
weevil’s power of injury would be diminished if we could obtain a 
variety of cotton with a fully determinate habit of growth, one which 
would cease producing buds as soon as a crop of cotton had been set. 

The much more rapid development of weevil larve in the bud is 
to be connected, doubtless, with the much richer food offered by the 
mass of pollen, but it may represent also a somewhat more definitely 
adaptive specialization of the life history of the weevil, for it is gen- 
erally a question of eating the pollen promptly or not at all. If the 
bud falls off on moist ground the pollen would be completely decom- 
posed long before the larva could develop, at the rate at which it 
grows in the boll, and if the bud did not drop off, but continued to 
grow, the flower would open and turn the larva out. It is obliged, 
therefore, to do damage fast enough to keep the flower from opening, 
and must then eat the remaining pollen before it spoils and leaves 
the larva too hungry and stunted to pass through the final meta- 
morphosis into the adult stage. In a cotton which has a highly 
developed habit of shedding the injured buds it would not be so neces- 
sary for the larva to attack the pistil. It may be that this policy 
on the part of the weevils in Guatemala has a use to the weevil as 
being necessary to prevent the opening of the flower and cause the 
falling of the bud. 


64 WEEVIL-RESISTING ADAPTATIONS OF COTTON. 


The diversity in size of the boll weevils, while not unprecedented 
among insects, is unusual, and not without biological significance in 
the present connection. An explanation of the variation in size is 
to be found, no doubt, in the varying amounts of food which the 
weevil larvee can obtain, but there is needed, none the less, a special 
adaptability on the part of the weevil to permit it to reach a normal 
reproductive maturity in spite of very unfavorable conditions. The 
smaller weevils probably have less than a quarter of the weight of 
the large ones, which means that they are able to develop with a cor- 
respondingly small proportion of the food required to raise a full- 
sized weevil. The weevils developed in the bolls have a much greater 
uniformity of size. The small weevils are at once a means and a 
result of the acquisition of the habit of living in the buds, and espe- 
cially in the small ones, where the supply of food is often very small. 


RELATION BETWEEN PROLIFERATION IN BUDS AND IN BOLLS. 


The analogy of the mucilaginous tissue found in the young fruits 
of okra and other relatives of the cotton would lead us to expect that 
proliferation could occur more readily in the boll than in the bud, 
which may mean that all the varieties which proliferate in the bud 
will do so in the bolls as well. 

It was at first supposed that if the buds proliferated but not the 
bolls the result would be merely a postponement of the breeding 
season of the weevil for two or three weeks, or until the bolls had time 
to develop. Such a delay would be of great practical importance in 
retarding for that length of time the effective breeding period of the 
weevils. Moreover, most of the eggs of the weevils which had passed 
through hibernation would be lost by being laid in the buds, which 
would further keep down numbers in the early part of the season. 
There is, however, the further and still more important consideration, 
that the period of development of the weevil in the boll is very much 
longer than required for it to mature and emerge from the square.” 


aDeterminations of the length of the life cycle in bolls have been made only 
in a few instances. In 7 cases between August 15 and November 11, 1903, 
the average time required from the deposition of the egg to the escape of the 
adult from the opening boll was sixty-one days. The average effective tempera- 
ture for the period was 31.7° F., and the average total effective temperature 
required for development in bolls was therefore 1,933.7° F., or nearly two and 
one-half times as much as in squares. Several larvee often develop within a 
single boll. ‘They appear to remain in the larval stage until the boll becomes 
sufficiently mature or so severely: injured: as to begin to dry and crack open. 
When this condition of the boll is reached, pupation takes place, and by the 
time the spreading of the carpels is sufficient to permit the escape of the weevils 
they have become adult.—Hunter, W. D., and Hinds, W. E., The Mexican Cotton 
soll Weevil, Bul. 45, Division of Entomology, U. S. Dept. of Agriculture, 1904, 


p. 75. 


PROTECTION OF SEEDS BY LINT. 65 


Moreover, it seems that the adult weevil does not come out through 
the wall of the boll, but waits to be liberated when the boll opens to 
maturity. This would mean that if proliferation can exclude the 
weevil from breeding in the squares it would afford a practical solu- 
tion of the problem, since instead of merely delaying the emergence 
of the first brood of weevils for two or three weeks, none of them 
would be able to set about the work of destruction until the crop had 
begun to ripen, and all danger of appreciable damage would have 
passed. It seems, therefore, that the proliferation in the squares is 
the much more valuable characteristic to be considered in seeking for 
a weevil-resistant cotton. Proliferation in the bolls is very desirable, 
but the absence of it should not be allowed to figure very largely 
against a variety which might have a pronounced tendency toward 
proliferation in the bud. Nevertheless, other factors must enter the 
calculation, for thin-walled bolls might allow the weevils to escape 
earlier. In moist weather the bolls might not crack open, but give the 
weevils comfortable shelter all winter, as would seem to have been the 
case in the spring of 1905, when various observers noted that some of 
the weevils seemed to have the appearance of having emerged only 
recently from the pupal condition, their very light color showing that 
their outer covering of scales was still in place. 

The probability is, however, that the proliferation in both places 
will be found to depend upon the same internal factor or quality, 
so that it will be safe to assume that a high degree of proliferation 
in the bud could be taken as an index of what might be expected 
from the bolls. This would simplify the problem of selection by 
permitting us to confine our attention to the buds. 


PROTECTION OF SEEDS BY LINT. 


Like the large leafy involucre, the lint is also a peculiar feature 
of the cotton plant which may prove to have a practical connection 
with the weevil. Cotton is the only food plant of the boll weevil, 
and only the cotton, of all the related plants, has an abundant pro- 
vision of lint. Some of the species of Hibiscus have the seeds 
shghtly silky, but the cotton stands quite alone in the length and 
abundance of the hairy covering which grows out from the seeds at 
the time the bolls are most subject to weevil injuries. 

From the standpoint of those who believe that all characters are 
useful to the organisms which possess them, the interpretation of the 
lint as a weevil-resisting adaptation will not appear unreasonable, 
since it can scarcely be claimed that there is any other use of the lint 
so important to the plant as protection of the seeds from the weevils. 
In other respects the lint seems rather a disadvantage than other- 


9962—No. SS—O5 M 


3) 


66 WEEVIL-RESISTING ADAPTATIONS OF COTTON. 


wise. In a humid tropical country the seeds, if left to themselves, 
remain inclosed in the tangled mass of lint and usually rot. Birds 
might carry the lint away to build nests, and in so doing might assist 
in scattering the seeds, but in most of the varieties the seeds are to 
be detached only with difficulty. 

Composed as it is of nearly pure cellulose, the lint can afford very 
little nourishment, even in the younger stages. Between the lint and 
the watery proliferating tissue the weevil larva must find the inside 
of a cotton boll a very inhospitable place unless it can penetrate to the 
seeds. Dead and moribund larve are occasionally found in these 
unfavorable situations. And even the seeds themselves do not pro- 
vide so favorable a food as the pollen, as shown by the much longer 
time required by the larve to develop in the boll than in the square. 


PROTECTIVE SEED ARRANGEMENT IN KIDNEY COTTON. 


Further intimation of the protective value of the lint is to be found 
in the very peculiar Kidney cottons, so called because the seeds are 
crowded together in the central angle of the chamber and adhere 
firmly to each other, thus forming a small, kidney-like mass. This 
unique arrangement brings all the lint to the outside of the seed, and 
may be the explanation of the fact that the Kidney cottons are the 
only representatives of the Sea Island type which have gained a wide 
distribution on the mainland. The separate-seeded Sea Island cot- 
tons came from Barbados, where the boll weevil did not exist and has 
not yet been introduced. (See Pl. X, fig. 2.) 

The outer wall of the boll of the Kidney cotton is notably thinner 
than that of Kekchi cotton, so that the beaks of the weevils could 
reach through without difficulty. But with the layer of lint to sup- 
plement it the wall becomes, for practical purposes, much thicker than 
in the free-seeded varieties. The inner parchment lining is rather 
tough, though apparently less so than in the Kekchi cotton. 

The Indians about Trece Aguas, Guatemala, are said to recognize 
the weevils as enemies of the dwarf cotton, but it is the local opinion 
that the Kidney cotton is proof against them. | 

No weevils were found on the two bushes of Kidney cotton exam- 
ined in that locality, but these were single plants growing near Indian 
houses several miles away from the nearest field culture. In a forest- 
covered country like this part of Guatemala the luxuriant and 
tangled vegetation may well impede the flight of such an insect as the 
weevil. And if it lives, as supposed, only on cotton, its chance of 
reaching a single bush of tree cotton would be very small. That the 
buds and young bolls of the Kidney cotton are able to offer any abso- 
lute resistance to the weevil seems very improbable, and the abundance 
of weevils found on the large tree of Kidney cotton at Tueuru last 
vear proved that the immunity, if any, is not general. 


ved 


NATURE AND CAUSES OF ADAPTATIONS. 67 


The Kidney cotton, though commonly treated as a distinct species 
under the name Gossypium peruvianum, agrees with the Sea Island 
type in all its characters except the peculiar arrangement of the seeds. 
If this should prove to be an adaptive feature the idea of specific 
distinctness would have little left to support it. 


CULTURAL VALUE OF KIDNEY COTTON. 


The possession by the Kidney cotton of a definite weevil-resisting 
adaptation would naturally raise a question regarding its cultural 
value. It belongs to the Sea Island series, and has the long, fine fiber 
and smooth seeds. The growing of the seeds together in masses would 
still further facilitate picking and ginning operations. The bolls, too, 
of this Guatemalan Kidney cotton, at least, are larger than those of 
any of the Sea Island varieties. 

It is not hkely, however, that any of the varieties of Kidney cotton 
thus far known will be found of use in the United States, for all are 
perennial “tree cottons,’ which have refused thus far to flower or 
fruit in the period of growth allowed by the shorter summers of our 
Temperate Zone. In tropical regions this objection would not hold, 
and there appears to be no reason why the Kidney cottons should be 
disregarded in the search for varieties suited to the various soils and 
climates. The Trece Aguas Kidney cotton, for example, seems to 
thrive well in a humid mountain climate considered by the natives to 
be unfavorable for the annual Kekchi cotton, which is planted several 
hundred feet lower down. 


THE NATURE AND CAUSES OF ADAPTATIONS. 


To explain how such characters as the weevil-resisting adaptations 
arise involves an interpretation of general evolutionary questions upon 
which the scientific world is still by no means agreed. Nevertheless, it 
is evident that students of such subjects should conduct and describe 
their investigations in accordance with some consistent plan or 
policy, if their writings are to be understood or their facts intelligibly 
recorded. Moreover, it would be scarcely reasonable to maintain that 
such characters can be further increased by selective influence unless 
it could be believed that they had been assisted in the past by the same 
agency. 

It seems necessary to state that in the present report it is not 
assumed that the weevil-resisting characters have arisen as direct pro- 
tective responses to the injuries, or that they are the results merely of 
stimulation or irritation caused by the weevils, as other writers on 
evolutionary subjects might hold. Nor have they been thought of as 
caused by selection in any strict sense of the word. Though consti- 
tuting a most striking instance of the results of selective influence, it 


63 WEEVIL-RESISTING ADAPTATIONS OF COTTON. 


is believed that the cotton plant must first have originated in some 
measure the protective characters before the external conditions (in 
this instance, the weevils) could make them of advantage to the 
plants and thus encourage their further development. 

The older theory that environment and natural selection are the 
efficient or actuating causes of evolutionary change has lost many 
adherents in the last decade, especially among those who found 
themselves unable to credit any longer the idea that all the characters 
and differences of plants and animals are, or have been, of use to 
them. It has been shown, too, by Professor Weissman and his fol- 
lowers, that direct adaptations or responses of individual organisms 
to the environment are seldom or never inherited by their offspring. 
To take the place of the doctrine of direct environmental influence 
in evolution it has been suggested that there may be an internal 
“hereditary mechanism,” as it has been called, which determines 
adult characters in advance, in the reproductive cells, so that modifi- 
cations of the specific or varietal type can arise suddenly. Selection 
would determine, of course, which of such new “ mutations” should 
survive, but it would be a mere accidental coincidence if the new 
character happened to fit the conditions better than the old. 

It is possible, however, to explain evolutionary progress and select- 
ive adaptations without ascribing them either to external causes or 
to theoretical internal mechanisms. The diversity which plants or 
animals of the same parentage often show under the same conditions 
makes it evident that there is no precise mechanism which determines 
their form in advance, and all attempts at securing any absolute uni- 
formity or “ fixity ” of form and color have failed. The fact is that 
organisms, even of the same species or variety, are normally diverse, 
and must have ancestry mixed by interbreeding if bodily vigor is to be 
maintained for any great number of generations. 

The generalized “ specific type,” which is a product, as it were, of 
this diversity and interbreeding, is constantly and gradually chang- 
ing, and in many ways at once, though in some characters more rap- 
idly than in others. Selection, while in no strict sense a cause of 
this vital motion of the species or variety, may profoundly influence 
the direction and rate of change. Selection, in other words, explains 
adaptation, but does not explain evolution.* 

The word adaptation is used in more than one sense by writers on 
‘biological subjects. Some treat as adaptations the changes of form 
or structure by which many plants and animals are able to conform 
to the needs of different conditions. There are several plants, for 
example, which have normal broad leaves when they grow on land, 
and very narrow and much-divided leaves when they grow submerged 


«Natural Selection in Kinetic Evolution, Science, N. S., 19: 549, 1904. 


NATURE AND CAUSES OF ADAPTATIONS. 69 


in water. Some plants are hairy in dry localities, but are nearly 
naked in humid districts. Others treat these direct responses to 
external conditions under the heading of accommodation, and reserve 
the word adaptation for characters which appear regularly in a spe- 
cies or variety, but which fit it for some special condition, such as 
that presented to the cotton plant by the boll weevil. It has seemed 
proper, therefore, to discuss as protective adaptations any characters 
which seem to give the Central American varieties an advantage in 
withstanding the attacks of the weevil, particularly if it can be shown 
also that the presence of the weevil would tend to the preservation 
and extension of the given character. 

In the strict sense of the words, the weevil-resisting adaptations 
of the cotton plant would include only those characters which have 
been increased by the selective influence of the boll weevil, but in the 
broader practical sense we may treat as a weevil-resisting adaptation 
any feature which tends to limit the destructiveness of the insect. 

The adaptive nature of some of the characters of the Central 
American.varieties discussed in the present paper is reasonably obvi- 
ous, but in other instances extended studies in developmental biology 
and primitive agriculture might be necessary to determine the origin 
and development of a varietal characteristic which may have signifi- 
cance in the weevil problem. 

It is easy to understand that so injurious an insect as the boll weevil 
has exerted a definite selective influence ever since its remote ances- 
tors turned their attention to the cotton. Perhaps its earlier food 
plants were completely exterminated. The nearest living relatives 
of the cotton are the species of Hibiscus, Paritium, and Thespesia, 
none of which is known to have any attractions for the weevil. It is 
evident, too, that in the presence of the weevil the cotton plant would 
have met long ago a like fate if it had not been able to take on its 
various adaptive characters. That so many of the features by which 
it differs from its nearest relatives have such obvious connection with 
the weevil would certainly justify the belief that strong adaptive 
influence had been at work, even if the other circumstances were 
unknown. , 

In thinking of the relation between two organisms like the weevil 
and the cotton we often fall into the error of too great humanizing, 
so to speak; that is, we ascribe too great intelligence or too complete 
a reaction to cause or conditions. Thus the weevil, although highly 
specialized in some of its instincts, has, of course, no equivalent for 
the human judgment. It will puncture, as already seen, buds much 
too small to raise a larva, and will lay its eggs in the rind of the boll, 
where the larvee can never develop. If the conditions are too favor- 
uble to the weevil. as in humid regions, it would undoubtedly exter- 


TO WEEVIL-RESISTING ADAPTATIONS OF COTTON. 


minate its own host plant by permitting the cotton to produce no 
seed. Paradoxical as it may at first seem, we may, nevertheless, 
believe that the best conditions for the perpetuation of the weevil are 
those which are not altogether favorable to its unlimited multi- 
plication. 


CONSCIOUS AND UNCONSCIOUS SELECTION. 


There are two principal ways in which improved varieties of cotton 
and other cultivated plants come into existence. The first is by sud- 
den or abrupt changes, or sports; also called mutations, saltations. 
and discontinuous variations. ‘These are represented in cotton by the 
occasional appearance of a plant with brown lint,’ deeply divided 
leaves” (okra cotton) or very short branches (cluster cotton). The 
Guatemalan varieties represent a second type of evolutionary history, 
in which improvement is accomplished by more gradual progressive 
change, fostered and accelerated by selection. 

Two forms of selection are commonly recognized, natural and arti- 
ficial, the latter effected by man, the former by circumstances of the 
environment. This distinction is of doubtful value in any case, and 
quite obscures the important point in the evolutionary history of 
cotton and other plants domesticated by primitive man. It would 
be much better to think of selection as either conscious or unconscious, 
and between these two a very practicable difference exists. Conscious 
selection implies the preservation of individuals having a desired 
quality in the highest degree, while unconscious selection, whether 
by man, animals, or inanimate conditions, means merely the rejec- 
tion of the most unfit, so that the improvement of the species or 
variety is gradual. Conscious selection acts, of course, much more 


a1In Guatemala several tribes of Indians prefer brown cotton, and for certain 
garments use brown cotton only. Separate plantings of brown cotton are not 
made in the neighborhood of Secanquim, where our experiment was located, but 
there were said to be such at Cajabon and Lanquin, only a few leagues away. 
The Cajabon people have a dark-brown cotton called “ canch nok,” and a lighter 
brown called * canni nok.” 

On the Pacific slope Mr. William R. Maxon found considerable culture of a 
brown cotton called * ixcaco.” At Antigua a similar brown variety is said to 
have been grown formerly in considerable quantities, the common name of 
which is “ cuyuseate.” It was not learned that any special religious use or 
significance is attached to brown cotton in Guatemala, as is said to be the case 
in Peru and in India. 

» Some may be inclined to interpret these as reversions and to argue that the 
deeply divided involucral leaves may be a reminiscence of an ancestral charac- 
ter of the cotton. Or it may be that the divisions attained by the involucral 
leaves represent a tendency of specialization which the remainder of the leaves 
sometimes share by mutation, in accordance with the principle of translocation 
of characters recently formulated by Dr. R. G. Leavitt (Contrib. Ames Bot. 
Lab. No. 8). 


CONSCIOUS AND UNCONSCIOUS SELECTION. (al 


speedily than unconscious, but is subject to the serious danger of 
weakening its protégés by inbreeding, if the selection be too rigid and 
persistent. 

The unconscious selection by which the development of the pro- 
tective characters of the Guatemalan types of cotton has been encour- 
aged differs in no respect from the progress by which adaptive 
evolution takes place in nature. The Indians have planted and har- 
vested the crop, it is true, instead of the birds or other natural agents, 
but they have been entirely unconscious of the struggle for existence 
to which the cotton plant was being subjected by the presence of the 
boll weevil. The Indians were only another factor, along with the 
dry and moist climates, the keleps, and the turkeys. The problem 
has been solved in a genuinely natural fashion, and affords an excel- 
lent illustration of the nature of selective influence in evolution. 

Instead of representing the final possibilities of improvement 
in characters which vive protection against the boll weevil, the 
Indian varieties of cotton may be looked upon rather as. affording 
materials which conscious selection can render still more valuable. 
The proliferation character, for example, might never be brought to 
uniform expression by unconscious selection, because the possession 
of it would give the individual plant no advantage over its neighbors 
in the production of seed. The proliferating plant might produce 
no weevils itself, but the free movement of the insects would keep the 
general average the same. Indeed, a plant might easily sacrifice all 
its buds, set no fruit at all, and thus fail to perpetuate itself. Pro- 
liferation can become a direct advantage to the individual plant only 
under conscious selection. The full value of the newly ascertained 
protective adaptations will not be known until they have had the 
direct selective encouragement now commonly accorded to desirable 
characters of other cultivated plants. 

It may appear remarkable that such definite and potentially valu- 
able characters as the weevil-resisting adaptations of the Kekchi cot- 
ton should have remained so completely unrecognized hitherto. The 
explanation of this doubtless lies in the fact that cotton culture is 
practiced in Central America largely by the Indians and very little 
by the foreigners or the more intelligent part of the native community, 
so that it had not received scientific study. Even the existence and 
utility of the keleps, though apparently known to the Indians from 
ancient times, had entirely escaped the attention of the European 
residents of the country. That the Indians should have come to 
recognize the keleps as beneficial and necessary to a full crop of cotton, 
although not knowing that the weevils injure the cotton or that the 
keleps eat the weevils, only shows in higher relief the completely 
unconscious character of the selection conducted in this system of 
primitive agriculture. The Indians of Alta Vera Paz are extremely 


(2 WEEVIL-RESISTING ADAPTATIONS OF COTTON. 


stolid, uncommunicative people, from whom little information is 
likely to be obtained except as replies to direct questions. Familiar 
from their earliest childhood with the agricultural lore of their own 
tribe, it does not occur to them that these everyday incidents can be 
of interest to the white stranger, or if they perceive his interest they 
learned long since to fear it as a danger of further intrusion. Even 
our own cotton experiments were misunderstood as a menace of addi- 
tional demands for lands from the white men who now own so large 
a part of the country. 


SUMMARY OF ADAPTATIONS. 


If the facts stated in the present report have been correctly observed 
and interpreted, we must admit that the cotton plant is in a high 
state of adaptive specialization in its relations with its now famous 
insect enemy, the boll weevil. Indeed, it may be that the most dis- 
tinctive and important characters of the plant, from both the botan- 
ical and the agricultural standpoints—such as the involucre, the 
nectaries, the oil glands, the large bolls, and the very lint itself— 
are adaptive features which the selective influence of the weevil has 
brought to their present degree of development. 


CLASSIFICATION OF ADAPTATIONS, 


The adaptations of the cotton plant might be summarized from 
three different standpoints. A historical treatment would proceed 
from the adaptations of the bolls to those of the buds. Breeding in 
the buds, for instance, was evidently a later adaptation on the part 
of the weevils which has called for a second set of the protective 
characters on the part of the plant. 

It may be better, however, to classify the adaptations as such, 
without special regard to their historical sequence of derivation. 
The more practical purposes are served by dividing the adaptations 
into four groups: (1) Those calculated to avoid the weevils by gen- 
eral habits of growth; (2) those which exclude the weevils, or at 
least hinder their operations in the buds and bolls; (3) those which 
attract insect enemies such as the weevil-eating kelep; (4) those 
which prevent the development of the weevil larve, even after the 
eggs have been laid. 

ADAPTATIONS TO AVOID WEEVILS. 


1. Determinate growth. 

2. Early bearing. 

3. Long basal branches. 

4. Early rejection of superfluous squares. 
5. Seasonal bearing of perennial varieties, 
6. Prompt bearing after cutting back. 

7. Hairy stalks and leaf stems. 

8. Pendent bolls. 

9. Rapid growth of young bolls. 


~] 
lets 
—_ 


SUMMARY OF. ADAPTATIONS. 


ADAPTATIONS TO EXCLUDE WEEVILS. 


1. Involucral bracts grown together at base. 

2. Closely appressed margins of involucral bracts. 

53. Margins of involucral bracts strongly laciniate and hairy. 
4. Unusual size and width of involucral bracts. 

5. Calyx produced into slender hairy laciniz. 

6. Persistent flowers. 

7. Oil glands (7) of very young bolls. 

8. Thick-walled bolls. 

9. Tough linings of boll chambers. 


ADAPTATIONS ATTRACTIVE TO THE KELEP. 


Nectaries of leaves. 

Large outer nectaries of involucre. 
Large inner nectaries of involucre. 
Bractlets subtending inner nectaries. 
Continued secretion of nectar. 
Hairy stalks and leaf stems. 

Dwarf. compact habits of growth. 


OR go ke 


~ 
we 


=I 


ADAPTATIONS TO PREVENT DEVELOPMENT OF WEEVIL LARV-®. 


Shedding of weevil-infested buds. 

Proliferation of internal tissues of buds. 

Proliferation from the walls of the bolls. 

Absence of oil glands over dissepiments. 

. Growth of lint on seed. 

Compacted seeds (Kidney cotton). 

Lint confined to outer end of seed (San Lucas Sea Island cotton). 


Dom NS 


aS 


ADAPTIVE CHARACTERS OF DIFFERENT TYPES OF COTTON. 


The third standpoint for viewing the adaptive characters is that 
of the different types of cotton. All varieties share, to some extent, 
the older adaptive features, but the special characters are accentuated 
in different degrees in the various types. Our study has been directed 
toward the Kekchi variety, both on account of its relation to the 
keleps and because it has seemed to possess by far the largest series 
of adaptive features. But now that the existence of adaptations of 
practical value has been ascertained it will be necessary to canvass 


the field thoroughly. 
ADAPTATIONS OF KEKCHI COTTON. 


An enumeration of the adaptations of the Kekchi cotton is scarcely necessary. 
because that variety has nearly the whole series and most of them in a more 
accentuated form than the other types thus far studied. The few exceptions 
are noted below. 


ADAPTATIONS OF RABINAL COTTON. 


1. Prompt bearing after cutting back. 

2. Very hairy stalks, leaf stems, and involucral bracts. 
3. Closely appressed margins of involucral bracts. 

4. Involucral bracts grown together at base. 


74 WEEVIL-RESISTING ADAPTATIONS OF COTTON. 


ADAPTATIONS OF PACHON COTTON, 


1. Involucral bracts margined with stiff lacinize and bristles. 
2. Calyx large, the divisions slender and hairy. 


ADAPTATIONS OF SAN LUCAS SEA ISLAND COTTON. 


Definite seasonal bearing. 

Lint confined to outer half of seed. 
Proliferation in buds. 

4. Proliferation in bolls. 


wb ee 


ADAPTATIONS OF KIDNEY COTTON. 


1. Definite seasonal bearing. 
2. Seeds compacted at center, covered with thick layer of lint. 


ADAPTATIONS OF UPLAND COTTON. 


1. Shedding of weevil-infested buds. 

This is the only weevil-resisting character in which the Upland varieties 
excel the Kekchi cotton, but, as already explained, the habit is of practical use 
only in dry climates. The Upland cottons share, however, a large number of 
the adaptations, though in a less degree than in the Kekchi. Thus there is pro- 
liferation both in buds and in bolls, the stems and petioles are somewhat hairy, 
the habit of growth is somewhat reduced from the tree-cotton stage, the nec- 
taries are often large and active, the involucral bracts are sometimes well 
tolded together, ete. 

And now that the possibility of weevil resistance has been shown, 
variations may be found in all probability among our United States 
varieties which will enable weevil-resisting strains of the Upland 
sorts to be developed. At this stage of the inquiry it is too much to 
hope that the Kekchi type will prove to be adapted to the wide 
diversity of conditions to be found in the cotton belt. Either the 
Kekchi or the native cottons, or both, are likely to require extensive 
modification before the full value of the weevil-resisting adaptations 
can be realized. 


CONCLUDING REMARKS. 


The protection afforded by the weevil-resisting adaptations is 
most effective at the two ends of the period of development, but con- 
tinues in varying degrees from the young bud to the ripe boll. Under 
favorable conditions an extremely small proportion of the weevil 
eggs develop to maturity. Instead of a single attack being fatal to 
a bud or boll, the same fruit at its different stages may resist numerous 
punctures and egg-layings. The young bud is protected for a time 
by the closed involucre. After the weevils have gained entrance the 
first egg, and often the second or third, may be rendered harmless 
through the proliferation of the bud in its younger stages. Pro- 
liferation becomes less certain as the bud increases in size, but if egg 
laying be delayed a few days too long the development of the larvee 


CONCLUDING REMARKS. 75 


is rendered impossible by the opening of the flower. Then ensues 
another period of immunity while the withered flower remains in 
place and while the bolls are still too small to be attacked. Between 
about the quarter and the three-quarter size the bolls can still be 
parasitized, though proliferation reduces the successful attempts to a 
very small percentage. But after the lint has grown out, the lining 
has hardened, and the walls have become thick, the boll is well-nigh 
impregnable, though the surface may be roughened by a dozen or 
a score of warts, which mark the location of as many persistent but 
ineffectual attempts to gain entrance. 

As an instance of adaptive specialization the cotton plant seems des- 
tined to a very high rank. The development of such a series of pro- 
tective characters can scarcely be explained except upon the suppo- 
sition that the culture of cotton in Guatemala is extremely ancient, 
and of this there are many other indications. 

The practical utilization of these protective characters in the cotton 
industry of the United States may require the solution of many pre- 
liminary problems of acclimatization and adaptation, as well as of 
physiology and cultural methods. The proliferation characters, for 
example, appear to be much more pronounced in some varieties than 
in others, but they are also affected, probably to a very considerable 
extent, by conditions of climate or soil which check the growth of 
the plant or cut down its water supply and thus reduce the normal 
turgidity of the tissues.’ 

The weevil-resisting characters are much more highly developed in 
the variety of cotton cultivated by the Kekchi Indians of eastern 
Guatemala than in any other type yet known, and it produces also 
large bolls and lint of good length and quality, so that it may be of 
yalue in the United States. But even though the Kekchi cotton in 
its present form should prove, for any reason, not to be adapted to 
cultural conditions in the United States, it demonstrates, at least, the 
fact that the Upland type of cotton is capable of assuming other 
characters which will render it far better adapted to cultivation in 
the presence of the boll weevil than the varieties hitherto grown in 
the United States. 


a@That the transfer to Texas will not destroy the proliferating habit of the 
Kekchi cotton is shown by the following report from Mr. McLachlan: 

“On the 23d of August Mr. Kinsler and I made a comparative examina- 
tion of four varieties of cotton at Mackay, Tex., to determine the nature of 
their proliferation. Rows of Kekchi cotton from Secanquim and Lanquin, and 
two of native Upland varieties (Parker and King) were compared. The results, 
in brief, are that in squares the Kekchi cotton proliferated much more readily 
than did the native varieties. In the bolls all four varieties were about equally 
active in this protective adaptation. The extent of proliferation in the Guate- 
malan bolls was, if different in any way, somewhat greater than in the native 
varieties.” 


76 WEEVIL-RESISTING ADAPTATIONS OF COTTON. 


No end is in sight of the new problems and adjustments of cotton 
culture occasioned by the invasion of the weevils, and no assurances 
‘an be given in advance regarding the utility of the weevil-resisting 
adaptations, any more than with the kelep, or so-called “ Guatemalan 
ant.” Both have a present value, however, in proving that the weevil 
is no invulnerable dragon which it is hopeless to resist. Instead of 
having no enemies, as long supposed, the weevil is regularly preyed 
upon by the active and efficient kelep. And instead of there being 
no remedies which can be used against the weevil, it is now found 
that the cotton plant itself has a whole series of weevil-resisting 
characters—a whole boll weevil armory, as it were, from which we 
may select and sharpen the weapons which prove best suited to our 
purposes. 

The weevil period of each year, that in which the damage is done, 
extends from the time when the squares are large enough for egg 
laying to the period when a full crop would normally be set. If the 
value of the cotton crop be divided by the number of days of this 
period, the result will show the value of each day of protection. It 
has been estimated by Mr. W. D. Hunter that the boll weevil damaged 
the cotton crop in 1904 to the extent of $20,000,000. It is therefore 
a very conservative estimate that when the pest shall have spread 
over the other cotton-growing States the damage will be well beyond 
a million dollars a day for the growing season—in unfavorable years 
probably two million dollars or more a day. Each day of protection 
which can be secured by the utilization of weevil-resisting adapta- 
tions will have, therefore, very definite and considerable value, so that 
the study and perfection of this group of characters are sure to be 
the objects not only of formal scientific study on the part of special- 
ists but of general interest and consideration on the part of the prac- 
tical cotton-growing public. 


> 


2 eon ries 


DESCRIPTION OF PLATES. 


PLATE I. (Frontispiece.) Valley at Secanquim, Alta Vera Paz, Guatemala, the 
scene of experiments with weevil-resisting cotton. 

PLATE II. Fig. 1.—Mature plant of Kekchi cotton, to show small size and 
determinate habits of growth, compact foliage, and long basal branches. 
Fig. 2.—Plant shown in figure 1, opened to show numerous large bolls and 
habit of fruiting on basal branches. 

PLATE III. Involucres of Kekchi cotton, opened to show external and internal 
nectaries, bracts, and bractlets. (Natural size.) 

PuaTE LY. Fig. 1.—Involucres of Rabinal cotton, showing connate and closely 
appressed involucral bracts. (Natural size.) Fig. 2.—Open involucres of 
Egyptian cotton. (Natural size.) 

PLATE VY. Fig. 1.—Young buds of Kekchi cotton, showing numerous weevil punc- 
tures. The buds were split in half so that the full number of punctures 
could be seen. (Natural size.) Fig. 2.—Buds of Kekchi cotton (same as 
fig. 1), showing successful proliferations. (Natural size.) 

PuaTE VI. Large buds of Kekchi cotton, the distortion indicating proliferation. 
(Natural size. ) 

PLATE VII. Weevil-infested bolls of Kekchi cotton, showing larger number of 
punctures along the middle line of the carpel, where the oil glands are 
absent. (Natural size.) 

PLATE VIII. Carpels of Kekchi cotton, showing method of proliferation. 
(Natural size. ) 

PLATE IX. Fig. 1.—Kekchi cotton, successive stages of the boll. Fig. 2.— 
Kekchi bolls (right); King bolls (left), to show comparative size. (Re- 
duced to about one-half natural size.) p 

Puate X. Fig. 1.—Rabinal cotton, showing foliage, connate bracts, and weevil- 
infested bolls. (Reduced.) Fig. 2.—Bolls and seeds of Kidney cotton, 
showing oil glands and _ protective arrangement of lint and _ seeds. 
(Reduced. ) 


78 


PLATE Il. 


Bul. 88, Bureau of Plant Industry, U. S. Dept. of Agriculture 


Fic. 1.—MATURE PLANT OF KEKCHI COTTON. 


Fig. 2.—KEKCHI COTTON PLANT WITH BOLLS. 


Bul. 88, Bureau of Plant Industry, U. S. Dept. of Agriculture. PLATE III. 


INVOLUCRES OF KEKCHI COTTON, SHOWING NECTARIES AND BRACTLETS. 


(Natural size.) 


+ 
| 


Bul. 88, Bureau of Plant Industry, U. S. Dept. of Agriculture. PLATE IV. 


Fia@. 1.—INVOLUCRES OF RABINAL COTTON, SHOWING CONNATE AND APPRESSED 
MARGINS. 


(Natural size.) 


Fia. 2.—OPEN INVOLUCRES OF EGYPTIAN COTTON. 


(Natural size.) 


Bul. 88, Bureau of Plant Industry, U. S. Dept of Agriculture. PLATE V. 


FIG. 1.—YOUNG BuDs OF KEKCHI CoT- Fia. 2.—BuDs OF KEKCH! COTTON 
TON WITH WEEVIL PUNCTURES. WITH PROLIFERATION. 


(Natural size.) (Natural size.) 


“= s 
5 = 
- 
& 
ay 
1. 
saneabe de 


Bul. 88, Bureau of Plant Industry. U. S. Dept. of Agriculture. PLATE VI. 


LARGE BuDs OF KEKCHI CoTTON WITH PROLIFERATION. 


(Natural size.) 


—o , ‘ , . 


4% 


Bul. 88, Bureau of Plant Industry, U. S. Dept. of Agriculture. PLATE VII. 


WEEVIL-INFESTED BOLLS OF KEKCHI COTTON. 


(Natural size ) 


Bul. 88, Bureau of Plant Industry, U. S. Dept. of Agriculture. PLATE VIII. 


CARPELS OF KEKCHI COTTON, SHOWING PROLIFERATION. 


(Natural size. ) 


PLATE IX, 


Bul. 88, Bureau of Plant Industry, U. S. Dept. of Agriculture. 


Se ear 6S ae NR a ta | 
Fig. 1.—KEKCHI COTTON, SUCCESSIVE STAGES OF Fic. 2.—KEKCH!I COTTON BOLLS (RIGHT) COMPARED 
THE BOLL. WITH KING BOLLS (LEFT). 


(Reduced. ) (Reduced. ) 


. ta = 
- * . ~ ® a 
( 5 5 “moe 
‘ H ¥ “i 
. / ‘ Sie * 
 /;, 
| 
. , r ; ——_ 7 ‘ 
' ‘ 
oF ea 


PLATE X. 


ericulture. 


Industry, U. S. Dept. of A 


i¢ 


88, Bureau of Plan 


Bul. 


Fic. 1.—RABINAL COTTON WITH BOLLS. 


(Reduced, ) 


(Reduced, ) 


Fia, 2.—BOLLS AND SEEDS OF KIDNEY COTTON. 


f 


24 


A 


4 
= Ar 
INDEX 
Page. 
PMEEnGORON, Harms Of SQUATCS.- ~~. 5.52... 2-2 Se bee eee eee 40 
MP mnie HPCClAlZatiOn —...- 695. S205. _ 22 LLU Th eel 29 
mamenmvation-of- Cekehi cotton. 2..-0222)7 252 020 9-2 Nee ete 17 
Accommodation, direct responses to external conditions. -......---.--------- 69 
EEE SE SY a ge ee, Bm A ae en 68, 69 
expiaimed by natural. selectione. 75-=. e222 eo Say See Feet oss: 68 
mammnawons, attractive to the kelep._...) 2.000... -0 0.22 lee ee---- 73 
CHES NUTCCHRS Oras epee oa Sree RL 0 hea a 72 
CS TUNT AUT CCA OG Geer see real Tory ce CE RI SL 10, 48, 44 
OMIMMTEr eral VANIEs tomes Sek ek nee eo eT, Be Ee ok 76 
Peete REM OMG teen hs. AAP ei oe et Sale LI ae 72 
PFE E TIE gL CALS CARRE ene ee een eer ed Pee a SNE So on 67 
periods: whenimostietect V6 <2. e-. ce Set dk ose ced 74 
OMARIAMECOLLONSUMKMO Wms 2922922). 2305. 8o 2 ULE eres tz, 32 
iKekechimandenabinatl: Coston sss 55550 Renee eee Sess sel 73 
Kidney, Pachon, San Lucas Sea Island, and Upland cottons - - 74 
summary...--- ee era ae ee taint micelle ate ne cee oe Haars om ae 72 
OPA OLA WEG Ville ee ae ea ea ta uetay Ae een cnck ares Se ole Rm oes 72 
CRAIC WCC MIIN Sti ted AN ots UUM Ee 2 See Soe: 73 
prevent development of weevil larve -..--....--.----------- 73 
PRE Mone AG LETS AOS FEN Lp sae oe acts Smee Ses ee. Soe Bee ya 69 
OTN EIS 2 eA REE ee SS Sens rea POM OeS SOME ee tc ege oc 67 
ERTL RCTECO TRG HA CLO Gy ets a1) SS age Set atorelctnrs SE ee eee = SG OE 53 
milearcotton, earlier than King in Guatemala... .....522..2..2-.-.--2-6--- 17 
Anthonomus, behavior of species related to boll weevil. ........--.---------- 48 
EMER EVEL NUR VaR IRGC GU OTIetem spe so 2, ee CE MAIS, OT Soe ee 19 
Aomeua, Guatemala, culture of brown cotton. ....- ....2.------.---+.--6 ee. 70 
PPC SRRPMAINE PARE MGCIs 21. ooo bole ot eekeh oe teen Sele es eS QL Nl 39 
Hid PRESSES ELORCO U6 Wier SS Syn TRA PR sar SES asi ponies ee 39 
neprOiPenion amainst WEeVils so. ow Sao Rea Le 39 
“GSS shire ue2 | Pets Ml 9) ge ee nee ey 8 8 a eg ee 29 
EMM HE IN COLON. 2. $2252 020 oe ec ion doe eee Oe 28 SSL 39 
Aphis gossypvi, infesting plants at Rabinal, Guatemala .........--..---.------ 39 
Arid climate, inactive nectaries an adaptive result .............----.----.--- 24 
KEM A mC AVI OM Oli CObtOM samy.csio.s weet wc Donne niecanwecss = 0 ESSE Lak 43, 44, 45 
Tue Orman aiapted.. 02. cake foots Beige toes wnt oe 20 
mrianeil selection, an indefinite term... .-=....--+..-.-....-------022. 52-641 70 
Abmnerde VV dlliamer. vanteidentifieds) 22222028 bk os eee es See Oe 39 
Asiatic cottons, comparison with other varieties .............-.-.--.....i..-- 53 
LEVEL DTANES| 2 ets SE ee on Rene SER ree oy eg hats Renney 28, 31 
Bacterial diseases following weevil injuries.................---....---.-.--. 62 
nna pe one Meriar MEOCUCeH = >. =--225--an0- 526+ e ee ee ope e+ see 36 
Mime Ueno as Weevil GESLLOVers .-. 2022-5 - 2222.2 2. oe lee cs een ee pe 42 
Boll weevil larve, adaptations to prevent development -...-.---..---------- 73 
behavior in Guatemala and Texas ...:..52.....--.--.-+.- 47 
destruction by drying out of squares...-.......----.-- : 43 
DLOULET AL ODN Reeser tear eee aren ee Se 46 
development in bolls - .--- Se, GAS aE NLS Re = a ee 46 
elfect or Opemino. Of HOWEISs 225-52. 2 - ~~ scencnt tenn ssea ae 47, 68 
rape development tn UNE. 26s so oe a acim hae eminent 63 
weevils, absence from Easterri Hemisphere ......................-.---- 32 
Mexicanipl atedmterigmies 524.5 aoe os eee a ee 1] 
SUSE ef UN COTOR NONE A OLE | Seek ee ee 0 a a ai al i ay <n ae ee 72 
Gx CIC M eee Pe ene tS ho pace mete oe ets 73 
WG ORCROSS-LEL tl ZatlOMs satan meen eae Sas. oboe ac See kinase cae e 55 


WEEVIL-RESISTING ADAPTATIONS OF COTTON. 


Boll weevils—Continued. 


bacterial diseases following injuries -.......--...-.------------ 62 
behavior in absence: oi keleps..._- J. . -..2 22222552 eee 24 
Onf00b 226 2. oe a see cE Se ee 25 
WithiiMbouls.> > 8a. o23. o-eeeee ee 59 
best conditions for perpetuation. - «_..-..- 24. ..+--5-sesseneee 70 
change of habia’. - == 2-22.04. . 0625-22-50 4g6~ nae eee 63 
destruction by chickens and turkeys -..+ 22.2.2... 62a 24 
determination of life cycle in bolls and squares .....-....--.-.-- 64 
develcpment of pollen-eating habit ..............----.....-.-. 63 
diverse habits of males and females ._...-..-----2-seeeseeeeee 27 
diversity: SIZ@=: 2.020 2 <'s nee ee ie se eee eee 64 
effect on unprotected cotton field ...........:--2-22.58==eeee 35 
under favorable conditions. -...........:/.55-5 esau 69 
eggs, fate. 20 cee ib saee se ccee es ce ee eee oe 58 
exclusion from large bolls.-.......2...- 2-2-5 =e ee eee 59 
feeding in bolls ..: 22...22 2.22 s2det li oe eee 61 
general effect on cotton plants .......:.2..:..../:celseue eee 10 
in fields cultivated by Indians -..2-.2--- 2. ..-22e22eee == 35 
injuries in Guatemala --......-...-------+--s¢255e5=-se =e 47 
injurious nature unknown to Indians. -.-.-...-----..-.---------- 71 
mortality In Spring: << o2n.<c0csccn 7s snoe 2a 2k ee 14 
not attracted by nearest relatives of cotton........ 00 ese 69 
pollen: diet. .-..22.5-.2--22-4-02s535-52 0) eer 55 
preference for upper portion of plant............-----.--------- 27 
selective influence ~~... -.-<220.2--002 4o6. so4= ls coon 10 
self-destruction’. ..2- «.0.5--2 seS0Ues..0560-40-.= eee “= 34 
short-season varieties of cotton for control .......---.---------- 22 
starvation .....- 2-006 -6-22+4--52455 55a ee 12, 14 
Bolls, diseases from superfluous nectar. .-_.-.-2..._---22..--e= eee 28 
efficiency of adaptive characters - -... ..-..2.22-5-2-25-Jsss= eee 61 
fed upon by boll weevils..-...-.--------2--.---.5as.55= =e 61 
feeding punctures ...-..--.2.2-------s-ss0-2-- 5005 61 
infestation by weevils... <2 -..-22- 2524202426 25.25) eee 59 
injured, discoloratiom.... .. - .- 2 ..2--. 26082246. 02 502 56, 59, 62 
Kekchi cotton, description... . =~ -=2S2:255-52)2ce25- 5-2 16 
IMMUNItY =... sac she cote wh ee eee = 56 
linings of chambers, toughness as protection from boll weevil ..-...-.-- 56 
normal, absence at Rabinal, Guatemala.._<:.___-_ 222-2323 a2eeeeeeee 24 
oil glands. ~~ - ..<<s-s<00.5+-2kenc2 etses oe ee 53 
outer walls, proliferation: .-....2.:2.2.-20..-->.2= ses 58 
variation in thickness ..2.......-..--54s.-356n eee 56 
pendent, discussion - .--.... 265.4 --5- Gees ene oe ee dee eee 27 
periods of immunity... -Jis52.6-. 22525260 = 52 see eee ee 74, 75 
Position ...-..-2.-0- 525 52 5see eee ews ote ae ees eee 27 
proliferation ........2- +s =2--0.2-5+.-+0—5-05 5S Spee eee 59 
in relation to that of buds'.---.2-20- 2322-5 seen 64 
protection... 26.2020. ose enn nee nace sc nso eos aaa ee en 51 
unconscious selection for uniform ripening ...-...........-----..----- 11 
very young, immunity... .. 2. o.-0-2.25<. 20053. see 52 
weevil punctures <.- .. 1... 025.2. ce cena e sc505 7-26 eee 61 
weevil-proof lining . ...... 2... --.....0ccs~o0.c-ss 55 55e ene 57 
young, rapid growth. .@.~..-/.22scei<--54- ‘ada clon oe oles en i) 
Bractlets of involucre......2-- 2-2... Ao ee so oe ee 
Bracts, appressed margins... .-...-.--.----------+---- 20-0222 2 senses enneanes 
dimensions’... 3p. 2022 oe sawn a ee ee eee oe toe ee 40 
flaring ..-..:...9R..~..---.-4.-------=250---6se=e0ae 40 
involucral, grown together.-......--.--.--++---- +22. saess=ee ee 37 
hairyamargines .... 2.2.6 3s 2s es ae eee 41 
lacinMe. .-.'(-2-.'- pe ene cana con cans ene eee eee en 43 
of Kekchi cotton. . 2.2. 52-020 s ne seta see toe ee ee 16 
protective feature of large'size. .....-..2... 52... 35-- <0 eee 39 
Branches, long basal, disadvantages of............... Joc... sceuen eee 20 
of Kekchi cottons... 26. 3).0. an. de oes oe ee 19 
of cotton, compared with those of other plants. ...............--- 19 
dimorphic . . 22.225 cease ens oceseseasks eee 19 


INDEX. 


Re gc aii gir lk op er 
TST vcs B10. Ur). ar cl ee es a 

pierre se TOUSEEVAMOUS! 6 - Ge stoe. Soon eo - fae a een eae 
fide. precaims in, 4 derived habit. -.....-.---:------2-------- 
SQV SER ETO Sal eel oT P50 1 0) 1 Seg ak an a ea a 
TELA LTTENMOUSS stent et oe SS ee See 

LLL SHLD OME (CIRM TEG ot era Ne eet i eas rene ae See 
Ovaries of, attacked by weevils. .2-...-=--2:-2£--.------- 
Ss ra a a 
perieds of immunity_.--....-..-.---- aS a Sie a Ue 
PEI RCTCRNITIULCR 2 3222 eo ne eke oe = =e oe 

0 UTE As i SS OT nO se 
enused and: condjtionss2 5-22 22254-2522. J. 


Dumcetom NY ABVOLUETS 0225-2222 - oe. Sent ss Jone = -- 
recovery from weevil punctures. ..---- See Pe Eve 
UES TINO GR ae te a Nicaea ag ti = Se 

CIE THU COTSV eT Wl hes oi Ss San a nee A a eee 
Ey rd Eva ETS eo VATS) 0 C50 (0 bc: agen ee ca ee a ee 


Gacso branches compared with cotton -.-...-:.--.----2------- 
Cajabon, Guatemala, cotton grown in vicinity ---....---.------ 

planting of brown cotton ----.-.-----.--- 
meatopia symbolic specialization, 2-2 -- 2.22522 5-22-.-24----- 
wammaun: Doll weevil destroyers....:..-.2:-22.---..---2----- 


Pueewon AUaApialiOns: 25. =.2252 522-22 L252 Oe one 
variations in cotton -..-- Bees See ee ee ere ee ae 
MinmSnercanet, (INCISION ~ 2-2 2e26c20 22s ma - be 22 ee 
ede ae Sk Ee LE ee ame TS Ge 

emmcaD Glee 5555 Stee eles eo ten ake 

Fejechion OL DUUS tare: Ae sabe ee Bese le he 

Reeeeee i I POLLO ooh on te ee cb aod os 

Coffee branches compared with cotton .-.....-.--------------- 


OES) See aie Tala eee er Ie) Se ae eee 
Seen ienGs Ny. \ONREEVALION =. 2.5.2 sotinobe Loosen ewes ke es 
BAIT IRIE PIC CII: oe Se Peer me Rey ES pe es oe 


mer UE. IMeSOe COMO... =. -=- Sx --ordpoe cnn ose nees. - - 
production, probable extension= 222.5: --5---=-2-.+.- 
SME COCO VATICMCS 222 2s 5. =. ofp eises meet fea p ein cues 


effect of simultaneous blossoming------------ 

BEchneGuiMrOuch Nectar =<. 24. eee = 

SEB ALI CCORONBY =e smth ake os ite elalnn esl et ect Beene ude 
Culture of cotton, methods, in eastern Guatemala..........---- 
on Pacific slope of Guatemala 

GTS = att Ne MR pe st | et eagle eh a 


tienes 


RUOrere IG: 43, 


Determinate growth of Kekchi cotton..........5..2....2----.--- £56 oss 


habit, disadvantages of early planting avoided 


Dry climate, region, etc. See Arid. 


Dwarf cotton, effect on cultural methods.............-.------- 
MAD UMCE Ie kehiiCObOHL <o)- s/c, cio.<ob mw oeeeweee ui 


Earliness, effect of development of primary branches ---------- 
Proiaolc COMMIMOUA . 7-4 L2 tots. 22s eo 

not shown by date of flowering...........---.------ 

Early bearing, facilitated by long basal branches........------- 
poosijane, Ceadvyantages . Ws scccs = deen oe ween -ss---- 53 


9962—No. 88—05 u 6 


es 


82 WEEVIL-RESISTING ADAPTATIONS OF COTTON. 


Page. 

Early planting, discussion 72.52 ss. occ cee ss aos od eke deen eee 2 13 

essential in southern Texas....----:-------------+-+eeceeeee 12 

Egg puncture sealed by weevil” -=..-./---. 2-2. -- 020) oben é 61 

Eggs of boll weevils, encystation _.............-.--2+-0.-u+2+-se eee 58, 61 

Egyptian cottons, immunity Tom DoOllworm -... - :..22ecene. eee ance 53 

involucral ‘bracts ...... ..- = -..2-22)-1222-2. 6-33 37 

less hairy than Upland: varieties. -.'»-....-.. 22245 25 

not immune to boll weevil. ......2.:-2:te-4..25- =e 42 

oll glands... 25 222. .2.biwi 2 seo ee eee 53 

of American Origin .......-2-.---seps +52 ee ) 

precocious in Guatemala. ..-....--.2-22.-225- Lig 

preference of weevils’......:2-2.-.-2- 2025. 46 ee Be 26 

proliferation . 2. .-...5 5-2-2. dos66e0-oae): = 1s 50 

susceptibility to injury .....-..-.-..--.52.55-45.5en 42 

Environment not actuating cause of evolutionary change.........-...------- 68 
Evolution, illustration of influence of natural selection ...........--.------- 3 ii 

1) volutionary origin. of external nectaries..-....2.-.2+2.:-/2+5.25 eee 31 

Fertilization, discussion .......0:5..2 s< 2.50.22 so0e 050-52 2 ee eee 52 
prevention by rain .2...--15sssss4-4-5--6- lees) 21 

Field cultures, failure in plateau region of Guatemala ...............--..--- f 25 

Flower bud, position: . 22.2252. ceise- scenic vets moe aie ae ae eee 20 

Flowering period, ehort:./..- ose ose hasten -cecs eo teee ae 23 

Flowers, failure of pollination ... 2.25.24... <-62--2202-00555 oo 51 

keleps imprisoned ... 2-2... ss). e0c0 code cos en eed ee fener 29 

of Kekchi cotton, color... -. 22.2222 -< sees ee se eee 54 

Kidney cotton, color. 22. 2252-5 42224-0422 oe 53 

opening,-effect on larvie...-...-.2... 2.6222 - see eee 47,63 
persistence... 55.55 ~- seed caso eae eee s wae ee Ee er 51 

POSITION - 2p bac eee ees ek oe a} eins ww ns «ee er 7 

Foliage, compact, effect on. keleps ........--5.22220c0s24e53- een a 26 

Food, change, advantage to plants and animals............-----4.--sseeeeee 26 

Fungi , growth in nectaries .......---+-22-.-cecc+ soe sebee 30 

Germination, cotton seed in Guatemala ..............--- 2.24) 5055.0 15 

Gossypium barbadense, origin ........02.222555.00<0 5 55050 8 

herbaceum, Asia Minor varieties =..5:-.....2-.0en. Jose eee ee 53 

confusion with G. hirsutum... 1.2... 22a 9 

nectaries': ... .95 325/226. ee eee ee 31 

peruvianum, specific validity ...---2.--2.2+25<-e.eusn==e=eee eee 67 

Growth, alternating periods in tropical plants ..........s.02) eee 23 

utility of acceleration: 222./'...is.02sagpee cs eee j Sate ace 5d 

Guatemala, central plateau. -.......-.2.5---.2 26. -dse ene see ee 24 

eastern, climate. 22... 2s2.<i.cciesscasccs eee eee Re 12 

methods of cotton cultures 65222-22202 eee eee 19, 70 

nature of the country ...:...--.. 2222.25 seneeee ee 15 

field experiments = i222. s2oesseees ‘soot ll ues ee 34 

importation of foreign thread ....:.........1 2.20 sa ee 24 

Indian methods of cultivation. _-..- Ee ee se ne 12 

western‘ cotton’: 2 -..2 2600.02. 0ssces Oe A Sem erny U 

Guatemalan conditions, effect on United States varieties .......2......--.--- iv 

cottons, preference of keleps -...2...2. Ji. St 22e oe eee er 25 

Hairs, cotton plant, assistance to keleps........-.....:.---- «22's Soe eee s 25 

Hairy Guatemalan varieties of cotton.....:.........-..--<.5=¢eeeen : 25 

stalks andeleaf st@ms of cotton ...........0.5...2-4. cena oe 25 

Heredity, mechanism. @ 2.2.22. e052 05) os eng ee owe ee 68 

Hibiscus, involucral appendages of species ...........-..-.-------------- ant 33 

Nectaries Of MMCCIES. .. n+ sn dese uncwetan ee RR ey i. 32 

species not atmractive to boll weevil. ...:...22.-.. soe eesee eee ise 69 

Hinds,.W.. c.; ObservaOns «ii .o. sce vesewo geese see 25, 42, 43, 44, 55, 61, 62, 64 

Howard, L. .O., observations . ...2c.00.. 2... sewn ce wee ce eben e ete enn ann 45 

Humid distric ts, behavior of cotton... -..<.0.5..<..6.~.5ene eee Se _ 48, 44 

taunter, -W .);, ODSELYStLONSs cesmewces ooU ee eee 14, 25, 42, 43, 44, 48, 55, 61, 64, 76 

Immunity of buds and bolls, periods. ............-- *, MSs. dO eae ie 74, 75 

Inbreeding of cotton by Indians... .'..6232.Jis:..u l/l Je tee eee 15 

Indeterminate varieties unsuited to early destruction of plants...... cee goee 13 


INDEX. 83 


. Page. 
Mertemenretuods OlMeonONGUliurGe*==.-2: 4255ss0.-52 20s. S22. ko eecases ste 11, 35 
PETIMC TOM COMO BAe oe ent ees ee ee co ee Sk EL eee 53, 70 
Detain AMCAiraraabites. > M6. 2 2 2 sisoeus cleo cell. bese dll. 11, 15, 35, 48 
CORROMACE GUC mee re Ame Lear ms die eee ee i SN Re Lo es ahiie tor ar/l| 

AML OM Om PepPenssa==>seaees otek oe. coe ko ase eee ce ccs 1] 
SsromevatkesDimal Cuatemalaest sa fo. 22S ee ele ke 24 
MAMACINOL NEWAVERICUCS © it baa 4 = + one eas oe ce oe eke eel - cence 15, 71 
Guatemalan, observations on boll weevils.............-..--.------- 66 

GwAlta Vera Paz, Guatemala, characteristics’. -- £22 225.f.2-.2.2 22. (Ble G- 
BEIECOUSOMEOQILOME a= Se kecaece Se Saws chee Sone Ue Ne Se ee sa 15, 16, 71 
PERLMAN Oi COULOIN- ee er eee eee Lee Ae ewe a Se acts nis wernioe 45 
Ee eCeRSary. 00. VICOr. .2.seS4oscot-<<5 $2 eles el os.e cease 68 
Involucral appendages of plants related to cotton --.-.-.-.--.--------------- 33 
DLAcionO mL AeChOM COUOR== er ease see ae ae ee ees otc 2 SSS. 20 

fomenere, aaa protective structure ....--.---.c.-2- lis. sete tee ee eee 37 
oRa ULC kane ome eee kee A ee eal Soe ee re na ee Ble LL oS 33 

external nectaries: GISCUSHION: 22 ssece ft ee a2 SELLS kk lisek cee 31 

LIME MCChneIsGINCUSSION:. ooh een oes ce ace Meee ee ne sen esas 31 

protection to PASS HOCIIPICN. Ae oe ae hee eee PO hs 28 

Perr ROBIN ta oe Er eh ee See CU ates oe 37 
nme ClOSPO ah Vanlacese.— <.. ates tes ee Ske a eee ee cee 38, 41 
BPG all COLON RSs be = aoe eee eee san | ano 8 ce oe 41 

PERE ace Fan ee iho Te pee rs nt le ce PO ue pet das DL OOM = LE 37, 39, 40 

LESLIE ENB OUR OL ANS SE MS are ee Ne a ae 32 

PT ESD og S2W a EEE ale Sg ee Cae as a ae age er 42 

PD Un ECO RLO UM a aes em Aen meee Mee ey. Uh aera Te ae 37 

Hearlislangd cottonmatiractive. toukeleps!. 2-3. 2-ca...5 eos sco ates 26 

ING ee oS re Bere ees eS 38 

AT SRIEE ROLE HUN UO See a) Sie cay me on eee ec le ee 39, 41 

oo SS Set a ee eee ee ee eee oe yest Span ee ae 45 
PRE Hat ChALeMAls < «= 1- cire Sooo Sens Shae wien ence sewer elec cnn 70 
Jamnovireh cotton compared with Allen... <2... 2s..200i22.2.25..22202002005- 17 
co Mietahinksn Sis Wo SS h OSes sos oo Soo59 bs aed ote S 50 

Jewett cotton, dimensions of buds and bracts.-.-..........--...-----.------- 40 
SE IMPea Gn UAC ALIONS 202 252 ny ae ee a oled Beck Sas wos | ne cee eens 73 
aAtmorthern limit ok¢otion culture s4* 22236. b eles. See 2. 18 

Dehayl Oren OxAd seks a eee Ss ce ete ated et Es Se 17 

bolls, variation in thickness of walls........---.....--.------ 56 

characteristic EGER U ULL Cages aes yee needs tees Re mS No ee 20 

comparison with other varieties, at Eanharn, Mido Se 18 

duisien planing in Guatemala .25 3252 5022 Pecks Lege 15 

developed in weevil-infested regions .......-.--.------------ 1 

distinct type of Gossypium hirsutum ....-..--------+-----+--- 8 

eHechroh crowdime on characher 25 4-50-2026 Saseb hee etek 20 

TITS ee Bezier Oo ee eee eerie ett eee es ee EVE Ee 17 

AICS OM KO Wwillesee asec a = Sa es eS 8 ae eT eS 15 

MES Chey ULM LCG SULCS Paes as Soe ns on See Ce ES Sey ily 

most promising short-season variety ....-.-.---------------- 22 

TIPE LEE CUP | Pe ee eo SO oer a ny ae 7 

Pp OLMeVElOPMOMi sso M50 Le be te ce 55 

BOUL NEHA an peasOli: PX PlANALGN, -<~ 2-2-2). 22 hs oot Sook 11 

stems petloles, leaves and bolls. --22 =... 2.<2.02-2. 2... << 16 

OLE TAN CETh: GOL mr serene serene a= Me te ee oR 15 

(AEN FOL ONS oe Re Ee ie REE Re ae ere ee eee 15 

Weevil cesistance,igh Vallee = 22. 22 2s nn kt 11 

ep sen. prom astern. Hemisphere ...:..--<---<--<2-----s---<+-+--- 32 
CPi GAIT ON EEN ELOY AST Pele Ch eA See es OR Bt Ras Sia ey Sa ae 73 
fenmiely attracted to: cotton plants: 222.29. 2h.0..2.. 22 -<-~25--6-- 35 
Sree MaL RCM OUADO: | e522. pe eta None ei cr Uaeaed me ce ea Shin 26 

UEDA: E22 Se 5 aaa Mea ee <i aie ee 36 

hairs of cotton plant, assistance in climbing --...--.-.-.----..------ 25 

Riiiat EXP IC I OWCYS => so 2 aks aot hone oe ong te So ncn en OSS 29 
preference for Egyptian and Sea Island cotton and Bidens pilosa.. 26, 36, 37 
EGKECHOURLOICOLUGI <j sa. sea er ete ee 8 PL sis 8, 34, 35 


unknown to European residents in Guatemala .......----..--------- 71 


84 WEEVIL-RESISTING ADAPTATIONS OF COTTON. 


Page. 

Keleps, usefulness in comparison with ants.......-.-.------.-----------ene 39 
value known. by Indians] -~ co22-<. .. 4-052. 522 ee ee f(s 
visitation of other nectar-bearing plants than cotton. ......-.-.....-- 36 

Kidney cotton, adaptations -.....-.5.----2. 0-20.55. eose 055s =e 74 
at Trece Aguas, Guatemala........-..-.02.0=-ee oe 32, 53 

Tucuru, Guatemala... . =... ..5 40.052 eee pee 24, 66 

Characters. 52202 caso’ 2 ons os oa a's Sal oe re eee 67 

cultural*value ... 652 Js2o42 22-5055 005%.55 eee yee 67 

of American origin. ......2.-<2 -s2s6--2+5.0 seep 9 

oil glands! 2.2 2.2032 3.222 scons Soe 53, 54 

origin ‘of names. o ci. ee os Sw eee ee 9 

protective features..-....-:- 2262... 2.cns: 2) eee 42 

seed arrangement... 2...- 2c. 4228s 2 ce eee 66 

thin-outer wall of boll. 22... 2c. - 22.2% <bean 66 

King cotton, behavior In:Guatemala....: 220... 2 =255seeeee ease 17 
cluster varlety..< 2.2.0.2 est bese goo. 2 eee ee 17 

dimensions of buds'and bracts:.. < -2-0 2-2. 2e.- ese 40 

flaring of squares . 2.2... 42. 20s ncec uses 2 Sone 40 

large néctaries..2.....'....--. cc cek else alee wee ee 31 

proliferation .........- needcceesscege coe. eiteee © ae ea! 

rapidity of development ...-.2.------.eceue ee rr 55 

Kinsler,.:J. EH. ODServatlons 20 faeces aos eee 17, 29, 37, 47, 50, 51, 55 
plants cultivated in Guatemala .........- 2. -2-5 -0=== eee 15 

Lacinis of involucral bracts... 2.2.0. 4.20202. -4-2 322-6 er 43 
Lanham, Md:, experiments'’...:~..2...2...2 32. 2. eee eee 18, 25, 43 
Lanquim, Guatemala, planting of brown cotton. ....--..--..---------Jsuesee 70 
Large varieties, disadvantaves* 12222 (225) oo ee eee 5 <-\sa% a 19 
Late planting in northern localities ............-.-..------. 02 13 
Leaves, Kekchi cotton... 2222.02 22 2h Soke cee 2 cee 16 
nectaries, general discussion -..--:......-.---.-¢ === eee 30 
Leavitt, R. G., principle of translocation of characters .......-..-.---...--< 70 
Limbless variety of King cotton ........22..2 44-2802 17 
Lint, disadvantages....-..2 2.0... 5-1-0. 0005565 side Geo co aoe rr 65 
protection :of seeds . ....--22.- 4 -cedene cece bende ns rr 65, 66 
Locks, number in Kekchi cotton -......-2- 22.2.2. 2456456 Se 16 
Mackay, Tex., examination of cotton -.2:....+-..5.-.-0s5.52s0s seen 75 
Maxon, W. R., cotton obtained... .-...5-<2-+J ses a00-- > ee 25 
’ obser vations. ooo ssc ccicke cov be eee eee 70 

McLachlan; Argyle, observations: 3.22. --2o- eee eee 18, 25, 41, 47, 57, 60, 61, 75 
Melanthera deltoidea visited by keleps.....-..+-2.45+-+4-5 40-255. —== rr 37 
Mexican plateau, absence of weevils ....---.--.5-.-<s.s.00ssss5= =e 11 
region, advantages... J 2.2,4:+02...0%.s466e eee 43 

Mexico, cotton culturé... 2-20. bo lee 8. enon 55 5s ee 45 
Mit Afifi cotton, oil glands... 02. b.) oc ee ces a see oe er ee 53 
Molds, growth in nectaries.<..-.. 22.05 622.- 452 eae oo ee ee eer 30 
Mogqui Indian cotton, character... -/...-.25-- 24. -5es.e0e ee 52 
Mutations in Kekchi cotton... 22-0022 222.5. 2. . ee chaee ete eee 16 
Natural selection by boll weevil ......-........¢.-2.d.-e6= = 055s ete 54, 69 
COULGM sz ow ae cee wlron etree eee eee os ureese eee eee 9 

conscious and unconscious: .-.-....-.-.- 2-25 =ne =e 70 

effect of oil glands’... ..2 2.2352 2c. 2.222 ee 54 

illustration of influence in evolution ........-.--...--..-.- 71 

in simultaneous flowering .....2... 15-565. e6 ee eee 23 

not actuating cause of evolutionary change ............-.-- 68 

the explanation of adaptation. . ..-.....3 5.0s0ss eee 68 

Nectar, continued secretion by plants.........2-5 2.50250 eee ae tee 32 
PUFPOSE ... 2. Bo. 2 ee eee re nner nese eee ee 28 
secretion in Kékchi and Upland cottons. ......... 2u@edes scene 32 

on. BOlls 2S. 2 cnc cts ce ob sd Seo ree 33 

Nectaries, external, discussion... .........026 25 edecssaee meee eee 31 
position in pendent bolls. ............0.t.sueseeeeee 27 

BECTOUION a. oo... e soe bes civics Deets «bee ha 27 

extrafloral, connection with bacterial diseases ...........----.---- 62 
GISCUSSION F. 2 PS ays ccc uisis 2 cow 2 we se eRe 28 


fUNCtIONS «2 .c. caw eae cc'esee cdacwcl ad es um pe ene 29 


INDEX. 85 

Page, 

Beenie ex tradorsl. ir Mmekeht COLON ----..--.-.< 222-2 dcen eee ee- een nee 34 
floral, no connection with weevil resistance .......-..------------ 29 

homology er eee RI oe eee er ee 32 

MMe Opie ITEMECOLLGD=s- Sst mete tase sce os ce te sen eae 24 

CaS SSS Sa Uae a eee ee 38 
Counececd: witty Uroopine. Dail: ©_- 22 ole eee ee cee 28 

of involucre, discussion ......--------------.-------------- 31 

MEHCRCI iE VAEICUICS = soe tnd tee CELE Son cee ee eee ae erate BIPRY 

CLOSES VEG111 0 ge hh a ee I Re 2 Ne RED 16 

fea eoricrAl GInCUbtOM-.2 2 WSs hoe ewe se owoeue ew enw sa 30 
2S ae ee a a 32 
ee eee ee ee eee 53 
: protective feature. -........------------------+----------------- 52, d4 

Coot. QoL NMSE OUR E Tao SE a ee eee 70 

Old World cottons. See Asiatic. 

OE ST a ae 21 
improved SpunteeeiO cede 2 oie Me oon 70 
DipmiEnWARteLICe OF COLON, - o.22sss2—-e-2 4 sees secs. -- =e 10, 44 

0 SEP SA Ua a ae 74 

SERS T eT DE RS ea ereeoe er s oe RP es e  Se e e S wis 2 34, 48 

comparison in -Lexas'and Maryland $s: 2) o.-5 2. . 222 -ces2--- 18 

[WEEE STOVES Ses ap cies ak RLS a See ee See a PO ee 25 

The G Ley eel Bh avire ote Abe eS Ree ee Se iy i tea pelt teeny eee ae 25 

Palmer, Dr. Edward, discovery of weev il Beet eaten eet oe Se ae ae eee 45 
SpscrqawuM on colon tec o he Tee 45 

eet CuPein OF Ol WeeVIS — 0.55 22.022 225-22 2-2--- sips eee ee eee eae 14 

Panam, species not.attractive to boll weevil. .-.......-----+-2.4---.-2----- 69 

Parker cotton, dimensions. ot pudsand: practswes =. Ol eS ee ee eee ie 40 

PRICED ES eet ee ea ooo Lesbo LSS Se aes 41,457, 60 
ACN Ol GEAR eae opens a ee ee re ema ep eee oi eae 40 
ie ne ea A a Sr eT? 60 

ene me CE ACISCUBSION a= 520 mies se. as ee See ae OL dee Soe ecick 27 

_ erminmetiance to indians). > 4.20. seca ooan ole eee es eee oe eee se 11 

Pea iinconouns annual eupune Pack oo 4 22255. 2 eee take e sc ole 24 

not likely to be of use in the United States_......-...---- 67 
Beanie upiearint ey se ce ae te ee Rs 23 

Pergande, Theodore, identification of PUES eee ene Se Se See NS cea, Sas 39 

PRC RPmnI ey ie RO KAR me. <0 ie ete se els Se aioe ee See Se 17 

anaes oounmness an Opstacle to. -keleps: <=. - 22.242 -cjra.Ne cheese ee sk sce 26 

eee eee COON {te ne se ee ee en tS Se tec ce 16 

eminem er RO DSEL Vth GUS." 20025 NM ager Peete ci aoe woe ab eed 26, 39, 46 

SPEnREC CCA Leitain se 2 2 a ee Sg ake apt bee eee ewe ne 19 
UC MMO TC OCAMTICRS = 4405 Sees as oa 35 SSSa SSS ie eso 13 

imc nh Ir OUS COrCOLLON soe os. So isin one ne en ess we SS tte oss elses 39 

Bie Oesiricnion wn the iall “dithiculty. 99 - A assoc ot abe eee dese ee 22 

Py lheie. GiGi GEE ete Gere Oe as See et ae ae ene ee 5d 

PLDERIN DAV nWVCe Villa bel EMCO eps eens nn Sees Ope bi SNS ers ieas SS o tale 47 
necessity fOr KOLA MALLE YT GiaweeVule! + S32 2= sean sets. ool ack 5d 

Paent-carine habit, development. -..- 2... 3. 3..0.s.322-5-2-2-----+------ 63 

Poneridie, Rami et Sie ee ea 30 

Proliferation, advantage only under conscious selection --.-.....------------ 71 

best weevil-resisting PETS oe oe OS ine Bee 9 
diminution of power in larger-buds-.......-..-.--...-.---.----- 51 
PaO eCy) WERINCT Senuert wee oe ecto a sy eee bet nss 2 60 
sREACRICSTT I Vem ee are pt Meee ermal Lobe Sica 3 46 
PES MR RATS Ltt DE ea ae sees eee ta) out cients mt gs Fre 49 
el Cey tal 0/0 tl eee eree ss ABU AY Su Oe ee ee eee ee eee i 58 

iis piemancspoOls. TelAtOne seein! 26 oo eg Sense books 64 
ECKG DIR GOLUOTES <5 = ee iar eget em bare i Sp TE ee 46, 49, 75 
ein eacOnlol =. ee ee ee ee ene yl 5d ba Betis Dose 60 
eG eT CO UUOI ~¢ 2 ee ai ae a ety ae 8 eet ee Set as ioe ak 60 
Vanleuessopner than, Wekchiecs. 5. este. 2 is aes lo oSek Sass 50 
OB ONN a. 5 5, Sere pee ee eee Ro ec: Ci sae Dee 55, 56, 59 
PIG GANISES ANG, CONGMIONSH essere! Loe te Se 49 
COUOM i ee 7-2 5 tayo eee reper gt MEN yes 60 ays SINS ae 48 
TREE NAL ASUes On DUAN Ana. eae: - Ws 22 el eoea. 46 


86 WEEVIL-RESISTING ADAPTATIONS OF COTTON. 


Page. 

Proliferation, probable effects of culture ......---.<<=42--5-s.024555e-n eee 50 

summary or resultsiot study. -.-.\..... 222-2 2a 49 

time required’ 2-22... 2.2... -22 52 6o ee eee ee 60 

SCN a a i are A Me - ek 65 

Protection afforded by adaptations, commercial value.....-..--------------- 76 

when most effectives: .-+--. 2-22 === 74 

the involucre -. ..'.--2--2-0=-+ s.5+4.-+-- eee 37 

tough linings of chambers of bolls ...-...-------.----- 56 

by keleps, efficiency .....-......---<0.5--c: -<<> see 34, 35 

of seeds by lint -. 2.0.2.2 2.2.2 seek - 2 cbse ce eer 65 

Protective characters, first originated by cotton plant ........-.-...-........ 68 

general ..22..-.¢.<2-ce20055005.2255 55 11 

Of bolls ....-.-..----see-ses22 6025 020 eee 51 

value of involucre «. =... -s..s.25--2- 2155225350565 37 

Quaintance, A. L.,, observations....-...-:--...2 5+ -.2- +2 ==> err 52 

Rabinal cotton, adaptations ............2----.2+5 2-5. 2. 2.6 er (B3 

bolls fed upon by boll weevils........2-..50020 0a 61 

characters -22....i...--5-40022¢2 2 2 2s4e ee er 41 

hairiness: ... 5.2. 2s2ee 525-6 -e ee eee eee eee 25 

involucres ......--- Sect cdees ee s22e sso 37, 38 

nectaries .. 225). c2..c0-522 6s cece ete eee ee 31 

Guateniala, ante....2.. 20.022. 5e-2 Jean- 2 oc ese eer 39 

cotton culttire =. 2.2.2.2... 265-2 e050 eee 9, 38 

customs of Indians: 2.2 .- .2--f2..se eee 24 

Rain; effect at time of flowering...-....--:------<<2. cn a= eee == 21 

Redshank cotton, characters .....:..2----2.0e2 26-542 se ee rr 53 

large nectaries........-.. .--<-5--- == = =a 31 

Retalhuleu district of Guatemala, cotton planted .......------.-----------=5 25, 34 

Rivers cotton, character in Guatemala ..---......-.-2255-2 4) == 17 

Rubber, Central American, branches compared with those of cotton......--- 19 

Sajal, plant often visited by keleps ....-.--- (oa beenesssuce eee ar 37 

Salama, Guatemala, cotton culture of Indians ........-...-.-----------+--+--- 24 

San Lucas, Guatemala, Sea Island. cottom.s25-2.- 4.2225. 4-2 eee ‘2s eee 24 
adaptations - ...2: .<-.t2s-2 4555s 7 

attacked by other insects -...-----..- 32 

discovery. ...2 5. Uo. ceeee 50 

nectaries = .=.:.-.552 -s-20e= eee 32 

Schwarz, E. A., observations:.--.-....-22--¢ 4-2-2. 26 see see 42,48 

on Cuban cottons .....2.-.s.2 sess se = 23 

Sea Island. cotton, Guatemalan, discovery ..-2...---.2-s5-=ouee= = =e 50 

nectaries. .-...£..-.+<. 254. 5een ee 32 

of San Lueas, Guatemala, annual flowering ....-.--------- 24 

cottons, comparison of petals with Upland varieties ..........---- 26 

flaring of squares ...--.-------. +--+ +202 200s e snes eeeaes 40 

involucral bracts ....... 2c. ....5< <0 dee eee 37, 38 

lacking in protective features ...--..-\.. ss sa=e. sae 4 

less hairy than Upland varieties ._-.......-2 52. s2=seeeee 25 

not immune to boll weevil...... 2. 2222025 es ee eee 42 

oil glands .........-...2+50-5- «2555+ osau ee 53 

OVIPIN gs oe oe ee eee oak ce oe ee 8 

precocity in. Guatemala... ....).25 3 9-26 ee 17 

smoothness a disadvantage to keleps ...............-..-- 25 

Secanquim, Guatemala, cotton grown in vicinity................--.sssssuees 15 

experiments... ........060ss00e sen cae ee ae 39, 41 

Seed, low germination in Guatemala. ............-.-----------+-++-++-++-++-+- 15 

Se eds, protective arrangement in Kidney cotton ........-...-.-------------- 66 
Selection, conscious, unconscious, natural, and artificial, discussion ......- da 7 0F7 1 

in cotton, time required wcdeueceedaeeeusccel oneuleenne Ente hE 10 

provided by boll weevil... ........5--eussesteuceuu Meee een 10 

Selective influence of boll weevil.......... 0. cscs. Jessuueus Sennen 10 

Short season varieties of cotton... .. 2505. suc... Seen 12, 14, 22 

Solenopsis picea, ant inhabiting Rabinal cotton.................222...eeseeee 39 

Specific type, generalized, the product of diversity and interbreeding. ......- 68 
Squares, boll weevil injuries -.... 42.2.2 0cc.c cen w ccs ou came ete 41 

flared and fallen, countings.<..<...0....5. S000 Oi Geen ee 45 


AATING. 5.6 cc deus swe ek bulsouwvicacemalsue eee ccs ee etn woe RO pe 


INDEX. = 87 


Squares, multiplication in cluster cottons. ..........--.--------------------- 28 
Auta DrCeOMmonDIACeS Ol WEGVIlS=a > -a5 2-255 So ce a Soa esos <5. 11 
PH CREP SCTORNION 8 sap oon ao ono se ee dene eee 20 
OO CS Ss eS 21 Ue ee a es pe ae a aes 43, 74 

EUS See ne 2 i a a a ee eee 49 

Siipules of outer bracts represented by bractlets.......-....---------------- 33 

SIRE OVATICHICO OMCOMUON@ 22. 22 ao Seats oe io Sees eso nss cao. see 27 

EEK SONDEMSSION = 26 20 Oo so ott ds an onde eee wee -- 21 

Symbiotic specializations between plants and animals ..-.....-....--.------ 29 

Tapinoma ramulorum, ant inhabiting Rabinal cotton ...............--------- 39 

Temperatures at localities where Kekchi cotton is grown.......---.--------- 15 

EE ER IOPUNENGD 2 2 = = eens son ans shane node e ues eso tess 52 

PET aeVeImh OF WEEVIIG 2.0 = eae eee eee ee eece one-one ee 9 

saupuern, method of checking weevil .-.:-..--.-=--. ----.------------ 12 

Thespesia, species not attractive to boll weevil ...-...-- St Meter tee ee 69 

DC MCSIRBEACGCVIL@ ©. 05. 252 o2 5 Soci s non 2ec eae ede enes carer ecss-<- 22 

le eR ROUT SO UIGe Siac oe ec ats SP ne eee See kee S's tela 23 

LAPSE TSS SA ee eS ee: fl a pe enc eee Ol eae 17 
BIpiniLy 10. Weevils, reported... -.2=2222222 --ass--es-~---- 23 
in Guatemala.......-- EE chad alte Sn DUAR Le, oe a ne |S, Se a 12 
EDTEST ENRON OA eo 1S ee SES Se ae ae een eee 8 
hel eel GaN OPE CN > Pe ge Se ee me Me Le a 3 
PRET RELMUOV A POiE ATG FoR ae ans es Ci ae AND eee Ree eee ia 21 
SERNA Le A OCU EER OUI Sree cf eee ce cea ae ars care eee ee aie lara 23 
(See also Perennial cottons. ) 

Meee acini ObsenyVatlOnS 228 32s ose a SD eben ces ole Leste 33 

aeomcae america, field for experiments.-__-_.---.----.---.-------.-------- 9 

Pon aeenaninprotected by: the: kelep >. :-=:-..22222.2-225-2. 22-22 ete tenes 36 

Sieirieenareinials Kidney cottom>. 22.22. .2255.5 22.22.7225 ohne ote 22k 24, 66 

MgmneuempOloweeyil GeStroyers: 0225 0. 22 ol S222 festa i so. tle elle. s ee 24 

Ea RO USENVALIONS 252202 ..c se eree ecko test foe oe bee Sacto ses 28 

United States cotton varieties, effect of Guatemalan conditions.............-- 17 

Meee HOALLARIONA 9 2 hoon) nee oot oe esos. eos ecko eae Se oe 74 

Speen Ohne’ OF Chae! oo toe o loo oe eee ll. 16 

indeterminate habit in United States varieties -............-. 13 

Matiwvedny CentralAmertea. - 5-2 hose. 5- 228222225 Sabot ese k 10 

“LE a SS ae RR apne ee EN Rene a eee 10, 44 

Sb sten OO Oh WelOpiee <5 one ae oe Sk See 25 

Si eR RIORL PTAC OLE ait ks 5 ey ee os Se Ok kN 32 
MEMnGnrClTMn Ae IM COULON! == aja0 25 se oe anne ee oe oe eee ee se 17 
een MENG IRE CO LCN IS Se te = ey ee es td 15 

Bred erg MORCIY COLON. > 55-2 525 yee S22 See Sods lees cose 16 

eso now. iormaton Dy Indians: ...:2...-1-- 252.22 sess lee eucbe.. 15 

Bemeaerle xe Se xUenmenis- ses 2- a yal oe oe a2 teed Se ee te ee ce 60, 61 

Volunteer cotton, absence in eastern Guatemala ...............2.22--....---- 12 

DLCCoIn Of weewWlsa. ss) sat sues aoe oe, A ree 12 

Ee MNER, (CLLGCE ON CLOP on = 1c fe eo eee AS ee et dene 13 

exclusion of boll weevil from Mexican plateau -...........--- 11 

erie AS ALeIMeMt Sao o-Ps i sae te ee es 13, 42 

Weaving by Guatemalan natives, use of foreign thread ..................--- 25 

Weevil. See Boll weevil. 

GENS 0 Se a ee 57 

eV eiraman, Professor, doctrine of inheritance. :.-.-.....-...---...---...---. 68 

Wy imwvers, Beyere, no protection from weevils. ...............-.-----.-..----- 12 


O 


“a 


DS Er Aki MENT OF AGRICULTURE. 
BUREAU OF PLANT INDUSTRY—BULLETIN NO. 89. 


B. T. GALLOWAY, Chief of Bureau. 


WILD MEDICINAL PLANTS OF THE 
| UNITED STATES. 


BY 


add dO OM oka s bel Oph il NO Wee 


ASSISTANT, DruG-PLANT INVESTIGATIONS. 


IssuED JANUARY 16, 1906. 


=== 


fe PA 
. Me 


nye? 


o 


FOR nn 


WASHINGTON: 
GOVERNMENT PRINTING OFFICE. 
L906. 


BUREAU OF PLANT INDUSTRY. 


B. T. GALLOWAY, 
Pathologist and Physiologist, and Chief of Bureau. 

VEGETABLE PATHOLOGICAL AND PHYSIOLOGICAL INVESTIGATIONS. 
ALBERT F. Woops, Pathologist and Physiologist in Charge, Acting Chief of Bureau in Absence of Chief. 
BOTANICAL INVESTIGATIONS. 

FREDERICK VY. COVILLE, Botanist in Charge. 

FARM MANAGEMENT. 

W. J. SPILLMAN, Agriculturist in Charge. 

POMOLOGICAL INVESTIGATIONS, 

G. B. BRACKETT, Pomologist in Charge. 

SEED AND PLANT INTRODUCTION AND DISTRIBUTION. 

A. J. PIETERS, Botanist in Charge. 

ARLINGTON EXPERIMENTAL FARM. 

L, C. CORBETT, Horticulturist in Charge. 


INVESTIGATIONS IN THE AGRICULTURAL ECONOMY OF TROPICAL AND SUBTROPICAL 
PLANTS. 


0. F. Cook, Bionomist in Charge. 
DRUG AND POIS*NOUS PLANT INVESTIGATIONS, AND TEA CULTURE INVESTIGATIONS. 
RODNEY H. TRUE, Physiologist in Charge. 
DRY LAND AGRICULTURE AND WESTERN AGRICULTURAL EXTENSION, 
CARL 8S. SCOFIELD, Agriculturist in Charge. 
EXPERIMENTAL GARDENS AND GROUNDS. 
EK. M. BYRNES, Superintendent. 
SEED LABORATORY. 


EpGar Brown, Botanist in Charge. 


J. KE. RoCKWELL, Kdilor. 


JAMES E. JONES, Chief Clerk. 


DRUG-PLANT INVESTIGATIONS. 
SCIENTIFIC STAFF, 
RODNEY H. TRUE, Physiologist in Charge. 
W. O. RICHTMANN, W. W. STOCKBERGER, Haxperts. 


ALICE HENKEL, G. FRED KLUGH, Assistants, 


LETTER OF TRANSMITTAL. 


U.S. DeparTMENT OF AGRICULTURE, 
BurEAU OF PLant INDUSTRY, 
OFFICE OF THE CHIEF, 
Washington, D. C., October 30, 1905. 
Str: I have the honor to transmit herewith and to recommend for pub- 
lication as Bulletin No. 89 of the series of this Bureau the accompany- 
ing manuscript entitled **‘ Wild Medicinal Plants of the United States.” 
This paper was prepared by Miss Alice Henkel, Assistant in Drug- 
Plant Investigations, and has been submitted by the Physiologist in 
Charge with a view to its publication. 
Respectfully, B. T. GatLoway, 
Chief of Bureau. 
Hon. JAMES WILSON, 
Secretary of Agriculture. 


PREPAC E: 


In connection with the work of Drug-Plant Investigations many 
inquiries are received from various parts of the country asking for a 
list of the drug-producing plants of the regions concerned and for 
information as to the parts of the plants used in medicine, etc. It 
being impossible to comply with requests of this nature in any satis- 
factory way, Miss Henkel was asked to compile a list of the drug 
plants of this country, using as a basis the catalogues of dealers in 
crude drugs and the standard works on systematic botany. It has 
seemed from an inspection of these lists and of much current phar- 
maceutical literature that the recent changes in botanical nomenclature 
have succeeded one another too rapidly to permit the drug dealer and 
the pharmacist to keep pace with them. This has resulted in consider- 
able confusion in regard to botanical names, and in some cases in the 
matter of the common names of drug-producing plants. In such a 
list as that herewith presented the opportunity for helping to clear up 
this situation has seemed worth improving. The recent appearance 
of the new Pharmacopeeia, in which the botanical nomenclature has 
been revised, has seemed to emphasize the desirability of making this 
attempt, since the names in the case of official plants will be fairly 
definitely fixed among pharmacists for the next ten years. In the 
accompanying list the pharmacopeeial names are given and a revision 
of the nomenclature of the unofficial drugs is also presented. Mr. 
Frederick V. Coville, Botanist, has kindly revised the botanical names 
used in this publication. 

It is hoped that this compilation will tend to unify usage among 
those who have to do with crude drugs and drug plants. 

Ropnry H. True, 
Physiologist in Charge 
Orrice oF DruGc-PLANT INVESTIGATIONS, 
Washington, D. U., October 12, 1905. 


B. P. I.—187. 


WILD MEDICINAL PLANTS OF THE UNITED 
STATES. 


In the preparation of this bulletin only such wild medicinal plants 
as have a commercial value were considered; that is, such as were 
usually mentioned in the trade lists of drug dealers throughout the 
country. Plants that were found listed by only one or two firms 
have been omitted. 

Both official and nonoflicial drugs are included in this list. A num- 
ber of drug plants that were official in the United States Pharmacopceia 
for 1890 have been dropped from the Eighth Decennial Revision 
(1900), which became official on September 1, 1905, and a few new 
ones have been added. In this bulletin the drugs that were official in 
the Pharmacopeeia for 1890 are so indicated, while those of the new 
edition are marked simply ‘* official.” 

In the following list the information on each species .is given under 
the accepted botanical name. This name and that of the family to 
which the plant belongs occupy the first line of the description. Botan- 
ical synonyms, if any, are mentioned, and these are followed in the 
next line by the most common names. A few words of information 
indicating the most important features of habit and stature, as well as 
the sort of situation in which found, together with the geographical 
distribution in the United States, are then given in each case. This 
information is too meager for the identification of the plants concerned 
in all cases, but it was impossible within the space limits of a publica- 
tion such as this to include more descriptive matter. The parts of the 
plants used and the official status of the product close the description. 
Unless otherwise indicated, the products mentioned are used in the 
dried state. 

Abies balsamea (L.) Mill. Pine family (Pinaceae). 


Balsam-fir; Canada balsam tree. 


Slender, evergreen, native tree, 50 to 60 feet high, occurring in damp woods from 
Newfoundland to the high mountains of southwestern Virginia, west to Min- 
nesota, and northward. 

Parts used.—Balsam, known as Canada turpentine, Canada balsam, or balsam of 
fir (official); also bark (nonofficial), 


11072—No, 89—0)6——2 7 


8 WILD MEDICINAL PLANTS OF THE UNITED STATES. 


Abies canadensis Michx. Same as Tsuga canadensis. 

Abies nigra Desf. Same as Picea mariana. 

Abscess-root. See Polemonium reptans. 

Absinth. See Artemisia absinthium. 

Absinthium. See Artemisia absinthium. 

Acacia, false. See Robinia pseudacacia. 

Acer rubrum L. Maple family (Aceraceae). 


Red maple; swamp-maple. 

Large, native tree, often 120 feet in height, growing in swamps and low grounds 
from Canada to Florida and Texas. 

Part used.—Bark (nonofficial. ) 


Achillea millefolium L. Aster family (Asteraceae). 
Yarrow; milfoil; thousandleaf. 

Perennial weed, 10 to 20 inches high, common in fields and waste places nearly 

throughout the United States, especially eastward; naturalized from Europe 


and Asia. 
Part used.—Herb (nonofficial). 
Acorus calamus L. Arum family (Araceae). 


Calamus; sweet-flag. 
Native, herbaceous perennial, about 2 feet high, found in wet and muddy places 
and along streams from Nova Scotia to Minnesota, southward to Florida and 


Texas. 
Part used.—Unpeeled, dried rhizome (official). 
Actaea alba (L.) Mill. Crowfoot family (Ranunculaceae). 


White cohosh; white baneberry; necklace-weed; rattlesnake-herb. 

Native, perennial herb, 1 to 2 feet high, found in rich woods from Nova Scotia 
to Georgia and Missouri, and northward; most common from Indiana and 
Kentucky to Pennsylvania and New York. 


Parts used. —Rhizome and rootlets (nonofficial ). 


Actaea racemosa 1. Same as Cimicifuga racemosa. 
Actaea rubra ( Ait.) Willd. Crowfoot family (Ranunculaceae). 

Synonyin.—Actaea spicata var. rubra Ait. 

Red cohosh; red baneberry; rattlesnake-herb. 

Native, perennial herb, 1 to 2 feet high, found in woods from Nova Scotia to the 
Middle States, west to the Rocky Mountains; most abundant from New 
England to Ontario. 

Parts used. —Rhizome and rootlets (nonofficial). 

Actaea spicata var. rubra Ait. Same as Actaea rubra. 
Adam-and-Eve. See Aplectrum spicatum. 
Adder’s-tongue, yellow. See Hrythronium americanum. 


Adiantum pedatum L. Fern family (Polypodiaceae ). 

Maidenhair-fern. 

Native fern, 9 to 15 inches high, growing in rich moist soil in woods in Canada 
and almost all parts of the United States. 

Part used.—Herb (nonofficial). 

Aesculus glabra Willd. Buckeye family (Aesculaceae). 

Ohio buckeye; fetid buckeye; smooth buckeye. 

Small, native tree, 20 to 40 feet in height, found in woods and on river banks 
from Pennsylvania south to Alabama, westward to Michigan and the Indian 
Territory. 

Parts used.—Bark and fruit (nonoflicial ), 


AESCULUS HIPPOCASTANUM—ALNUS RUGOSA. Cy 


Aesculus hippocastanum L. Buckeye family (Aesculaceae). 
Horse-chestnut. 


Large tree, 60 feet or more in height. Escaped from cultivation, southeastern 
New York and New Jersey. Native of Asia. 


Parts used.—Bark and fruit (nonofficial ). 
Aiterbirth-weed. See Stylosanthes biflora. 
Agrimonia eupatoria (of American authors, not L.). Same as Agrimonia hirsuta. 
Agrimonia hirsuta (Muhl.) Bicknell. Rose family (Rosaceae). 

Synonym.—Agrimonia eupatoria of most American authors, not L.4 

Agrimony; tall hairy agrimony. 

Perennial herb, 3 to 4 feet high, found in woods and thickets from New Bruns- 
wick to Minnesota and Nebraska, south to North Carolina; also in California. 
Native. 

Part used.—Herb (nonofficial). 

Agrimony. See Agrimonia hirsuta. 
Agrimony, tall hairy. See Agrimonia hirsuta. 
‘ Agropyron repens (L.) Beauv. Grass family (Poaceae). 

Synonym.—Triticum repens Beauy. 

Triticum; couch-grass; dog-grass; quack-grass. 

A troublesome grass in cultivated land from Maine to Maryland, west to Minne- 


sota and Missouri; sparingly distributed in the South. Introduced from 
Europe. 


Part used.—Rhizome, gathered in spring (official). 
Ague-tree. See Sassafras variifolium. 
Agueweed. See Eupatorium perfoliatum and Gentiana quinquefolia. 
Alder, black. See [lea verticillata. 
Alder, common. See Alnus rugosa. 
Alder, red. See Alnus rugosa. 
Alder, smooth. See A/nus rugosa. 
Alder, tag-. See Alnus rugosa. 


Aletris farinosa L. Lily family (Liliaceae). 
Star-grass; false (not true) unicorn-root;? colic-root. 


Native, perennial herb, 2 to 3 feet high; in dry, sandy soil from Maine to Minne- 
sota, south to Florida and Tennessee. 


Part used.—Rhizome (nonofficial) , gathered after the plant has flowered. 
Allspice, Carolina. See Butneria florida. 
Allspice, Florida. See Butneria florida. 
Allspice, wild. See Benzoin benzoin. 
Alnus rugosa (Du Roi) K. Koch. Birch family (Betulaceae). 
Synonym.—Alnus serrulata Willd. 
Tag-alder; common alder; red alder; smooth alder. 


Native shrub, or sometimes a small tree, occurring in swamps and marshy bor- 
ders of streams from the New Engiand States west to Minnesota and south- 
ward to Florida and Texas. 


Part used.—Bark (nonofficial ). 


a According to Bicknell (Bul. Torr. Bot. Club, 23 : 508-525, 1896), the name Agrimonia eupatoria L.., 
long used in local floras and text-books for the agrimony of the Eastern States, has been doing duty 
for a group of related species, of which at least five are now clearly recognized. Furthermore, Doctor 
Britton (Bul. Torr. Bot. Club, 18: 366, 1891) states that the true Agrimonia eupatoria is not known at 
all as an American plant. The native plant to which the name Agrimonia eupatoria has been most 
frequently applied by American authors is Agrimonia hirsuta (Muhl.) Bicknell. 

>The name ‘true unicorn-root’’ has long been applied to Aletris farinosa, but as ‘‘ unicorn-root”’ 
was the common name first given to Chamaelirium luteum (Helonias dioica), this should more prop- 
erly be called the true unicorn-root and-Aletris farinosa the false unicorn-root. 


10 WILD MEDICINAL PLANTS OF THE UNITED STATES. 


Alnus serrulata Willd. Same as Alnus rugosa. 


Alsine media L. Pink family (Silenaceae). 
Synonym.—Stellaria media Cyr. 
Common chickweed. 


Small, annual herb, probably introduced from Europe, and now common in 
fields and around dw ellings throughout the United States. 


Part used.—Herb (nonofficial) . 


Althaea. See Althaea officinalis. 
Althaea officinalis L. Mallow family (Malvaceae). 
Althaea; marshmallow; sweatweed; mortification-root. 
Perennial herb, 2 to 4 feet high, naturalized from Europe; occurs in salt marshes, 
coast of Massachusetts and New York, and in Pennsylvania. 
Parts used.—Root from plants of second year’s growth, deprived of the periderm 
(official); leaves and flowers (nonofficial) are also used. 
Alum-root. See Geraniwm maculatum and Heuchera americana. 


Ambrosia artemisiaefolia L. Ragweed family (Ambrosiaceae). 
Roman wormwood; ragweed; hogweed; stammerwort. 


Coarse, native weed, annual, 1 to 3 feet high; in waste places, eastern United 
States, west to British Columbia and Mexico. 


Part used.—Herb (nonofficial) . 
Ampelopsis quinquefolia Michx. Same as Parthenocissus quinquefolia. 
Amy-root. See Apocynum cannabinum. 


Anagallis arvensis L. Primrose family (Primulaceae). 
Red chickweed; red pimpernel; scarlet pimpernel; shepherd’s-weatherglass. 


Low, spreading, annual herb, naturalized from Europe, and growing along road- 
sides and in fields almost throughout the United States. 


Part used.—Herb (nonoflficial). 


Anaphalis margaritacea (L.) Benth. & Hook. Aster family (Asteraceae). 
Synonyins.—Gnaphalium margaritaceum L.; Antennaria margaritacea Hook. 
Everlasting; pearly everlasting; large-flowered everlasting; cottonweed. 
White-hairy or woolly perennial herb, native in dry soil from Newfoundland to 

Alaska, south to North Carolina and California. 


Part used.—Leaves (nonofficial) . 


Andromeda arborea L. Same as Oxydendruin arboreum. ) 
Anemone patens var. nuttalliana A. Gray. Same as Pulsatilla hirsutissima. 
Angelica, American. See Angelica atropurpurea. 
Angelica atropurpurea L. Parsley family (Apiaceae). 
Synonym.—Archangelica atropurpurea Hoffm. 
Purple-stemmed angelica; American angelica; masterwort. 
Tall, stout, perennial herb, 4 to 6 feet high; native in swamps and damp places 
from Labrador to Delaware and west to Minnesota. 
Parts used.—Root and seeds (nonofficial). 
Angelica, purple-stemmed. See Angelica atropurpurea. 
Anise-root. See Washingtonia longistylis. 
Anlennaria margaritacea Hook. Same as Anaphalis margaritacea. 
Anthemis cotula L. Aster family (Asteraceae). 
Synonym.—Maruta cotula DC. 
Mayweed; dog-fennel; fetid camomile (or chamomile). 
Strong-scented, annual herb, naturalized from Europe; occurs in dry soil, fields, 


waste places, and along roadsides almost throughout North America, with 
the exception of the extreme North, 


Part used.—Herb (nonoflicial). 


APLECTRUM HYEMALE—ARCHANGELICA ATROPURPUREA. 11 


Aplectrum hyemale Nutt. Same as Aplectrum spicatum. 


Aplectrum spicatum ( Walt.) B.S, P. Orchid family (Orchidaceae). 
Synonym.—Aplectrum hyemale Nutt. 
Adam-and-Eve; putty-root. 
Native herb, perennial, 1 to 2 feet high; in rich woods and swamps from Canada 
to Georgia and California. 
Part used.—Root (nonofiicial) . 
Apocynum. See Apocynum cannabinum. 


Apocynum androsaemifolium L. Dogbane family (Apocynaceae). 
Bitterroot; spreading dogbane; honeybloom. 


Perennial herb, 1 to 4 feet high, native in fields and thickets from Canada south 
to Georgia and Arizona. “The most common species in Canada and the North- 
eastern States. 


Part used.— Root (nonofficial). 


Apocynum cannabinum L. Dogbane family (Apocynaceae). 
Apocynum; ean hemp; black Indian hemp; amy-root. 


Perennial herb, 2 to 3 feet high, native in moist ground and borders of fields 
throughout fhe U nited States. 


Part used.—Rhizome of this or of closely allied species of Apocynum (official). 
Apple, custard-. See Asimina triloba. 
Apple, May-. See Podophyllum peltatum. 
Apple, thorn-. See Datura stramonium. 
Apple-of-Peru. See Datura stramonium. 
Aquilegia canadensis L. See under Aquilegia vulgaris. 
Aquilegia vulgaris L. Crowfoot family (Ranunculaceae). 


European columbine; garden-columbine. 


Perennial herb, with showy flowers. Naturalized from Europe, and well known 
in cultivation; escaped from gardens into woods and fields; frequent in the 
Eastern and Middle States. The wild columbine (Aquilegia canadensis L.), 
occurring in rocky woods throughout Canada and the eastern United States, 
is said to possess properties similar to those of the European columbine. 


Part used.—Herb (nonofficial). 


Aralia hispida Vent. Ginseng family (Araliaceae). 
Dwart elder; wild elder; bristly sarsaparilla. 


Ereci, leafy perennial, 1 to 3 feet high, native in sandy woods and fields from 
Labrador south to North Carolina, west to Minnesota and Indiana. 


Part used.—Root (nonofficial). 


Aralia nudicaulis L. Ginseng family (Araliaceae). 
American sarsaparilla; wild sarsaparilla; false sarsaparilla; Virginian sarsapa- 
rilla; small spikenard. 


Herbaceous perennial, native, growing in moist woods from Newfoundland west 
to Manitoba and south to North Carolina and Missouri. 


Part used. —Root (nonofficial). 
Aralia racemosa L. Ginseng family (Araliaceae). 
Indian-root; spikenard; American spikenard; spignet. 


Herbaceous perennial, native, 3 to 6 feet high, growing in rich woods and rocky 
places from Canada to Georgia, west to Minnesota and Missouri. 


Part used.—Root (nonofficial). 
Arbor-vitae. See Thuja occidentalis. 
Arbutus, trailing. See Epigaea repens. 
Archangelica atropurpurea Hoffm. Same as Angelica atropurpurea. 


13 WILD MEDICINAL PLANTS OF THE UNITED STATES. 


Arctium lappa L. Aster family (Asteraceae). 

Synonym.—Lappa major Gaertn. _ 

Lappa; burdock; cockle-button; beggars’-buttons; bardane. 

Coarse, biennial weed, 4 to 9 feet high, introduced from the Old World, and 
occurring along roadsides and in fields and waste places in the Eastern and 
Central States. 

‘arts used.—Root of this or of other species of Arctium collected from plants of 
first year’s growth (official). The fresh leaves and the seeds are also used 
(nonofficial ). 

Arctostaphylos glauca Lindl. Heath family (Ericaceae). 

Manzanita. 

A shrub-like tree, 9 to 25 feet high, growing in California, in dry, rocky dis- 
tricts on the western slopes of the Sierras. 

Part used.—Leaves (nonofficial ). 

Arctostaphylos uva-ursi (L.) Spreng. Heath family (Ericaceae). 

Uva-ursi; bearberry; upland-cranberry. 

Low, evergreen perennial, with trailing stems, native in rocky or dry, sandy 
soils from the Middle Atlantic States north to Labrador, westward to Califor- 
nia and Alaska. 

Part used.—Leaves (official). 

Arisaema triphyllum (L.) Torr. Arum family (Araceae). 

Synonym.—Arum triphyllum L. 

Indian turnip; wild turnip; wake-robin; Jack-in-the-pulpit. 

Native, perennial herb, 10 inches to 3 feet high, found in moist woods from 
Canada to Florida, west to Kansas and Minnesota. 

Part used.—Partially dried corm (nonofficial). 

Aristolochia reticulata Nutt. Birthwort family (Aristolochiaceae). 

Serpentaria; Texas serpentaria; Texas snakeroot; Red River snakeroot. 


Perennial herb, about 14 feet in height, native in the Southwestern States, occur- 
ring on river banks from Arkansas to Louisiana. 


Parts used.—Rhizome and roots (official). 


Aristolochia serpentaria L. Birthwort family (Aristolochiaceae ). 
Serpentaria; Virginia serpentaria; Virginia snakeroot. 
Native, perennial herb, 10 inches to 3 feet high, found in rich woods from Con- 
necticut to Michigan and southward. 
Parts used.—Rhizome and roots (official). 


Arrowwood. See Viburnum dentatum. 


Arrowwood, Indian. See Huonymus atropurpureus. 


. 


Artemisia abrotanum L. Aster family (Asteraceae). 
Southernwood. 


Shrubby, perennial herb, about 2 to 4 feet in height, occurring in waste places 
from Massachusetts to Nebraska. Adventive from Europe. 

Part used.—Herb (nonofficial ). 

Artemisia absinthium L. Aster family (Asteraceae). 

Absinthium; wormwood; absinth. 

Shrubby, perennial herb, 2 to 3 feet high, naturalized from Europe, and occur- 
ring in waste places and along roadsides from Newfoundland to New York and 
westward. 

Parts used.—Leaves and tops (official in U. 8. P. 1890). 

Artemisia vulgaris L. Aster family (Asteraceae). 

Common mugwort. 

Perennial herb, 1 to 34 feet high, naturalized from Europe; found in waste 
places, Nova Scotia to the Middle States and westward to Michigan. 

Part used.—Uerb (nonoflicial ). 


ARUM TRIPHYLLUM—ASTHMA-WEED, QUEENSLAND. 13 


Arum triphyllum L. Same as Arisaema triphyllum. 


Asarum canadense L. Birthwort family (Aristolochiaceae). 
Canada snakeroot; wild ginger; Indian ginger. 
‘Perennial herb, about 1 foot in height, native in rich woods from Canada to 
North Carolina and Kansas. 


Parts used.—Rhizome and rootlets (nonofficial). 

Asclepias. See Asclepias tuberosa. 

Asclepias cornuti Dec. Same as Asclepias syriaca. 

Asclepias incarnata L. Milkweed family (Asclepiadaceae). 
White Indian hemp; swamp-milkweed; swamp-silkweed; rose-colored silkweed. 


Perennial herb, 2 to 4 feet high, native in swamps from Canada to Tennessee and 
Kansas. 


Part used.—Root (nonoflicial) . 


Asclepias syriaca L. Milkweed family (Asclepiadaceae). 
Synonym.—Asclepias cornuti Dec. 
Common milkweed; silkweed. 


Perennial herb, 3 to 5 feet high, native in fields and waste places from Canada 
to North Carolina and Kansas. 


Part used.—Root (nonofficial ). 


Asclepias tuberosa L. Milkweed family (Asclepiadaceae). 
Asclepias; pleurisy-root; butterfly-weed; Canada-root; whiteroot. 
Native, perennial herb, 1 to 2 feet high, growing in dry fields from Canada to 
Florida and Arizona; most abundant southward. 


Part used.—Root (official in U. 8. P. 1890). 
Ash, American mountain-. See Sorbus americana. 
Ash, black. See Fraxinus nigra. 
Ash, cane-. See Fraxinus americana. 
Ash, hoop-. See Fraxinus nigra. 
Ash, prickly. See Fagara clava-herculis and Xanthoxrylum americanum. 
Ash, wafer-. See Ptelea trifoliata. 
Ash, white. See Fraxinus americana. 


Asimina triloba (L.) Dunal. Custard-apple family (Anonaceae). 
North American pawpaw; custard-apple. 


Small, native tree, growing in rich soil along the banks of streams from New 
York to Michigan and southward. Most common in the Ohio Valley. 


Part used.—Seed {nonofiicial ). 
Aspen, American. See Populus tremuloides. 
Aspen, quaking. See Populus tremuloides. 
Aspidium. See Dryopteris filix-mas and D. marginalis. 
Aspidium filix-mas Sw. Same as Dryopleris filix-mas. 
Aspidium marginale Sw. Same as Dryopteris marginalis. 
Asplenium filix-foemina (L.) Bernh. Same as Athyrium filix-foemina. 


Aster puniceus L. Aster family (Asteraceae). 
Red-stalked aster; cocash; meadow-scabish. 


Perennial herb, with stout, reddish stem, 3 to 8 feet high, native; in swamps 
and on banks of streams, Nova Scotia to Minnesota, south to North Carolina, 
Ohio, and Michigan. 


Part used.—Root (nonofficial). 
Aster, red-stalked. See Aster puniceus. 
Asthma-weed, Queensland. See Huphorbia pilulifera. 


14 


Athyrium filix-foemina (L.) Roth. 


WILD MEDICINAL PLANTS OF THE UNITED STATES. 


Synonym.—Asplenium filix-foemina (L.) Bernh. 
Backache-brake; female-fern; lady-fern. 


Fern family (Polypodiaceae). 


Native fern, with leayes 1 to 3 feet long; in woods and thickets, Canada to 


Alaska, southward to Florida and Arizona. 
Part used.—Rhizome (nonofficial). 


Avens, purple. See Geum rivale. 


Avens, water-. See Geum rivale. 


Backache-brake. See Athyrium filix-foemina. 


Backache-root. See Lacinaria spicata. 


Balm. See Melissa officinalis. 


Balm, bee-. See Monarda didyma. 


Balm, field-. See Glecoma hederacea. 


Balm, garden-. See Melissa officinalis. 


Balm, horse-. See Collinsonia canadensis. 


Balm, lemon-. See Melissa officinalis. 


Balm, mountain-. See Hriodictyon californicum. 


Balm, scarlet. See Monarda didyma. 


Balm, sweet. See Melissa officinalis. 


Balm-of-Gilead. See Populus candicans. 


Balmony. See Chelone glabra. 


Balsam, sweet. See Gnaphalium obtusifolium. 


Balsam tree, Canada. See Abies balsamea. 


Balsam, white. See Gnaphalium obtusifolium. 


Balsam-fir. See Abies balsamea. 


Bamboo-brier. See Smilax pseudo-china. 


Baneberry, red. See Actaea rubra. 


Baneberry, white. See Actaea alba. 


Baptisia tinctoria (L.) R. Br. 
Wild indigo; yellow indigo; American indigo; indigo-weed; horsefly-weed. 
Native, perennial herb, 2 to 3 feet high, growing in dry, poor soil from Maine 


to Minnesota, south to Florida and Louisiana. 
Parts used.—Root and leaves (nonofficial). 


Barberry, holly-leaved. See Berberis aquifolium. 


Bardane. See Arctium lappa. 


Basswood. See Tilia americana. 


Bay, rose-. See Rhododendron maximum. 


Bay 


, sweet. See Magnolia virginiana. 


3ay, white. See Magnolia virginiana. 


Bayberry. See Myrica cerifera. 


Bean, bog-. See Menyanthes trifoliata. 


Bean, buck-. See Menyanthes trifoliata. 


Bean, hog’s-. See /Tyoscyamus niger. 


Bearberry. See Arctostaphylos uva-ursi. 


Jearberry-tree. See Rhamnus purshiana. 


Bear’s-foot, yellow. See Polymnia uvedalia. 
y 


Pea family (Fabaceae). 


BEAR’S-WEED—BITTERBLOOM. 15 


Bear’s-weed. See Lriodictyon californicum. 

Beaver-poison. See Cicuta maculata. 

Beaverroot. See Nymphaea advena. 

Beaver-tree. See Magnolia virginiana. 

- Bedstraw. See Galiuvm aparine.~ 

Bee-balm. See Monarda didyma. 

Beech, American. See Fagus americana. 

Beechdrops. See Leptamnium virginianum. 

Beechnut-tree. See Fagus americana. 

Bee-plant. See Scrophularia marilandica. 

Beggars’-buttons. See Arctium lappa. 

Bellwort, perfoliate. -See Uvularia perfoliata. 

Benjamin-bush. See Benzoin benzoin. 

Bennet. See Pimpinella saxifraga. 

Benzoin benzoin (L.) Coulter. Laurel family (Lauraceae). 
Synonyms.—Laurus benzoin L.; Lindera benzoin Meissn.; Benzoin odoriferum Nees. 
Spicebush; feverbush; Benjamin-bush; wild allspice; spicewood. 

Indigenous shrub, 5 to 12 feet high; in damp, shady woods, and along streams, 
Massachusetts to Michigan, south to North Carolina and Kansas. 
Parts used.—Bark and berries (nonofficial). 

Benzoin odoriferum Nees. Same as Benzoin benzoin. 

Berberis. See Berberis aquifolium. 

Berberis aquifolium Pursh. Barberry family (Berberidaceae). 
Berberis; Oregon grape; holly-leaved barberry; Rocky Mountain grape. 


A shrub, native in woods from Colorado to the Pacific Ocean; especially abun- 
dant in Oregon and northern California. 


Parts used.—Rhizome and roots of this and of other species of Berberis (official). 
Bergamot, wild. See Monarda fistulosa. 
Bethroot, ill-scented. See Trillium erectum. 
Betony, Paul’s-. See Veronica officinalis. 
Betula lenta L. Birch family (Betulaceae). 


Sweet birch; black birch; cherry birch. 


Large, indigenous forest tree; Newfoundland to Ontario, south to Florida and 
Tennessee. 


Part used.—Bark (nonofficial). Oil of betula, obtained by maceration and dis- 
tillation from the bark, is official. 
Bikukulla canadensis (Goldie) Millsp. Poppy family (Papaveraceae). 


Synonyms.—Corydalis formosa Pursh; Corydalis canadensis Goldie; Dicentra 
canadensis Walp. 

Turkey-corn; squirrel-corn; turkey-pea; staggerweed. 

Native, perennial plant, 6 to 12 inches high; in rich woods from Nova Scotia 
south along the mountains to Kentucky, and westward to Missouri and 
Minnesota. 


Part used.—Tubers (nonofficial). 
Birch, black. See Betula lenta. 
Birch, cherry. See Betula lenta. 
Birch, sweet. See Betula lenta. 
Bird’s-foot violet. See Viola pedata. 
Birthroot. See Trilliwm erectum. 
Bitterbloom. See Sabbatia angularis. 


11072—No. 89—06——3 


16 WILD MEDICINAL PLANTS OF THE UNITED STATES. 


Bitter-buttons. See Tanacetum vulgare. 

Bitterroot. See Apocynum androsaemifolium. 

Bittersweet. See Solanum dulcamara. 

Bittersweet, false. See Celastrus scandens. 

Bitterweed. See Hrigeron canadensis. 

Blackberry, high-bush. See Rubus nigrobaccus. 

Blackberry, knee-high. See Rubus cuneifolius. ‘ 

Blackberry, low running. See Rubus procumbens. 

Blackberry, low-bush. See Rubus trivialis. 

Blackberry, sand-. See Rubus cuneifolius. 

Blackcap. See Rubus occidentalis. 

Blackroot. See Veronica virginica. 

Blackroot, Indian. See Pterocaulon undulatum. 

Blackwort. See Symphytum officinale. 

Bladderpod. See Lobelia inflata. 

Blazingstar. See Chamaelirium luteum. 

Blazingstar, blue. See Lacinaria scariosa, 

Blazingstar, scaly. See Lacinaria squarrosa. 

Bloodroot. See Sanguinaria canadensis. 

Bloodwort. See Hieracium venosum. 

Bloodwort, striped. See Hieracium venosum. 

Blowball. See Taraxacum officinale. . 

Blue-curls. See Prunella vulgaris. 

Bog-bean. See Menyanthes trifoliata. 

Bog-myrtle. See Myrica gale. 

Boneset. See Eupatorium perfoliatum. 

Boneset, deerwort-. See Eupatoriwin ageratoides. 

Boneset, purple. See Eupatorium purpureum. 

Bouncing-Bet. See Saponaria officinalis. 

Bowman’s-root. See Porteranthus trifoliatus and Veronica virginica. 

Boxwood. See Cornus florida. 

Brake, backache-. See Athyrium filix-foemina. 

Brake, buckhorn-. See Osmunda regalis. 

Brake, rock-. See Polypodium vulgare. 

Brassica nigra (L.) Koch. Mustard family (Brassicaceae) . 
Synonym.—Sinapis nigra L. 
Sinapis nigra; black mustard; brown mustard; red mustard. 
Annual herb, introduced from Europe; found in fields and waste places almost 

throughout the United States. 
Part used.—Seed (official); the volatile oil obtained from black mustard seed is 
also official. 

Brauneria angustifolia (DC.) Heller. Aster family (Asteraceae). 
Synonym.—Echinacea angustifolia DC. 
Echinacea; pale-purple coneflower; Sampson-root; niggerhead (in Kansas). 


Native, perennial, herbaceous plant, 2 to 3 feet high, occurring in rich prairie 
soil or sandy soil from Alabama to Texas and northwestward; most abundant 
in Kansas and Nebraska. 


Part used.—Root (nonofficial). 


BROOM—CALICO-BUSH. 17 


Broom. See Cytisus scoparius. 
Broom, green. See Cytisus scoparius. 
Broom, Scotch. See Cytisus scoparius. 
Brownwort. See Prunella vulgaris. 
Bruisewort. See Symphytuin officinale. 
Buck-bean. See Menyanthes trifoliata. 
Buckeye, fetid. See Aesculus glabra. 
Buckeye, Ohio. See Aesculus glabra. 
Buckeye, smooth. See Aesculus glabra. 
Buckhorn-brake. See Osmunda regalis. 
Buckthorn. See Rhamnus cathartica. 
Bugle, sweet. See Lycopus virginicus. 
Bugle, water-. See Lycopus virginicus. 
Bugleweed. See Lycopus virginicus. 
Bullbrier. See Smilax pseudo-china. 
Bull-nettle. See Solanum carolinense. 
Bulrush. See Typha latifolia. 
Burdock. See Arctium lappa. 
Burnet-saxifrage. See Pimpinella saxifraga. 
Burningbush. See Euonymus atropurpurea. 
Bursa bursa-pastoris (L.) Britton. Mustard family (Brassicaceae). 
Synonym.—Capsella bursa-pastoris Medic. 
Shepherd’s-purse; cocowort; toywort. 


Annual plant, about 1 foot in height, found in fields and waste places; widely 
distributed. Introduced from Europe. 


Part used.—Herb (nonofficial). 

Burseed, spiny. See Xanthiwm spinosum. 

Burweed, thorny. See Xanthium spinosum. 

Butneria florida (L.) Kearney. Strawberry-shrub family (Calycanthaceae). 
Synonym.—Calycanthus floridus L. 
Hairy strawberry-shrub; sweet-scented shrub; Carolina allspice; Florida allspice. 
Native shrub, 4 to 8 feet high; in rich soil, Virginia to Mississippi. 
Part used.—Bark (nonoflficial). 

Butterfly-weed. See Asclepias tuberosa. 

Butternut. See Juglans cinerea. 

Buttonbush. See Cephalanthus occidentalis. 

Button-snakeroot. See Eryngium yuccifolium. 

Button-snakeroot, dense. See Lacinaria spicata. 

Button-snakeroot, large. See Lacinaria scariosa. 

Button-tree. See Cephalanthus occidentalis. 

Buttonwood-shrub. See Cephalanthus occidentalis. 


Cabbage, skunk-. See Spathyema foetida. 
Cabbage, swamp-. See Spathyema foetida. 
Calamus. See Acorus calamus. 

Calfkill. See Kalmia angustifolia. 
Calico-bush. See Kalmia latifolia. 


15 WILD MEDICINAL PLANTS OF THE UNITED STATES. 


Calycanthus floridus L. Same as Butneria florida. 
Camomile, fetid. See Anthemis cotula. 

Canada balsam tree. See Abies balsamea. 

Canada-root. See Asclepias tuberosa. 

Cancerroot. See Leptamniwm virginianum. 

Candleberry. See Myrica cerifera. 

Cane-ash. See Fraxinus americana. 

Cankerroot. See Coptis trifolia and Limonium carolinianum. 
Canker-weed. See Nabalus serpentarius. 

Canker-weed, white. See Nabalus albus. 

Cankerwort. See Taraxacum officinale. 

Canoewood. See Liriwdendron tulipifera. 

Capsella bursa-pastoris Medic. Same. as Bursa bursa-pastoris. 
Cardinal, red. See Lobelia cardinalis. 

Cardinal-flower. See Lobelia cardinalis. 

Cardinal-flower, blue. See Lobelia siphilitica. 


Carduus arvensis (L.) Robs. Aster family (Asteraceae). 
Synonym.—Cirsium arvense Scop. 
Canada thistle; creeping thistle; cursed thistle. 


Perennial herb, 1 to 3 feet high; growing in cultivated fields, pastures, and waste 
places from Newfoundland to Virginia, west to Minnesota and Nebraska. A 
bad weed, introduced from Europe. 


Part used.—Root (nonofficial). 

Carduus benedictus Auct. Same as Cnicus benedictus. 

Carpenter’s-square. See Scrophularia marilandica. 

Carrion-flower. See Smilax herbacea. 

Carrot, wild. See Daucus carota. 

Carya alba Nutt. Same as Hicoria ovata. 

Cascara sagrada. See Rhamnus purshiana. 

Cassia marilandica L. Senna family (Caesalpiniaceae). 
American senna; wild senna; locust-plant. 


Native, perennial herb; in swamps and wet soil, New England to Florida, west 
to Louisiana and Nebraska. 


Part used.—Leaves (nonofficial). 


Castalia odorata (Dryand.) Woody. & Wood. 
Water-lily family (Nymphaeaceae). 
Synonym.—Nymphaea odorata Dryand. 
White pond-lily; water-lily; sweet-scented water-lily. 
Indigenous, aquatic herb; perennial; in ponds, marshes, and sluggish streams, 
from Canada to Florida and Louisiana. 
Part used.—Rhizome (nonofficial). 


Castanea. See Castanea dentata. € 
Castanea dentata (Marsh.) Borkh. Beech family (Fagaceae). 


Castanea; chestnut; American chestnut. 


A large, spreading tree, occurring in rich woods from Maine to Michigan, south 
to Tennessee. Especially abundant in the Allegheny region. Native. 


Part used.—Leaves (official in U. 8. P. 1890). 
Catchweed. See Galium aparine. 
Catfoot. See Glecoma hederacea. 


- 


CATGUT—CHAMAEILIRIUM LUTEUM. 1 


Je) 


-Catgut. See Cracea virginiana. 


Catmint. See Nepeta cataria. 

Catnip. See Nepeta cataria. 

Cattail, broad-leaved. See Typha latifolia. 

Cattail-flag. See Typha latifolia. 

Caulophyllum. See Caulophyllum thalictroides. 

Caulophyllum thalictroides (L.) Michx. Barberry family (Berberidaceae). 


Caulophyllum; blue cohosh; squawroot; papoose-root. 


Native, perennial herb, 1 to 3 feet high; found in rich, shady woods from New 
Brunswick to South Carolina, westward to Nebraska; abundant throughout 
the Allegheny Mountain region. 


Parts used.—Rhizome and roots (official in U. S. P. 1890). 

Ceanothus americanus L. Buckthorn family (Rhamnaceae). 
Jersey tea; New Jersey tea; redroot. 
A native shrub, growing in dry, open woods from Canada to Florida and Texas. 
Parts used.—Root, root-bark, and leaves (nonofficial). 


Cedar, red. See Juniperus virginiana. 

Cedar, shrubby red. See Juniperus sabina. 

Cedar, white. See Thuja occidentalis. 

Cedar, yellow. See Thuja occidentalis. 

Celandine. See Chelidonium majus. 

Celandine, garden-. See Chelidonium inajus. 
Celandine, great. See Chelidonium mMajus. 

Celandine, wild. See Impatiens aurea. : 


Celastrus scandens L. Staff-tree family (Celastraceae). 
False bittersweet; staff-tree; waxwork; fevertwig. ; 
An indigenous, twining; woody vine; in rich, damp soil, woods, and thickets, 

Ontario to Manitoba, south to North Carolina and New Mexico. 
Part used.—Bark of plant and of root (nonofficial ). 


Centaurea benedicta L. Same as Chicus benedictus. 
Centaury, American. See Sabbatia angularis. 
Centaury, ground-. See Polygala nuttallii. 


Cephalanthus occidentalis L. Madder family (Rubiaceae). 
Buttonbush; button-tree; buttonwood-shrub; globeflower. 


Indigenous shrub, 6 to 12 feet high; in swamps and damp places, Canada to 
Florida and California. 


Part used.—Bark (nonofficial ). 


Cercis canadensis L. Senna family (Caesalpiniaceae). 
Judas-tree; redbud. 


Small, native tree, growing in rich soil from New Jersey to Minnesota, south to 
Florida and Texas. 


Part used.—Bark of root (nonofiicial). 


Chamaelirium luteum (L.) A. Gray. Bunchflower family (Melanthiaceae). 
Synonym.—Helonias dioica Pursh. 
True (not false) unicorn-root;@ blazingstar; starwort; drooping starwort. 
Slender, perennial herb, about 2 feet high; native in moist meadows and thickets 
from Massachusetts to Michigan, south to Florida and Arkansas. 
Part used.—Rhizome (nonofficial ). 


aThe name ‘ unicorn-root’”’ was first applied to Chamaelirium luteum, and the designation “true 
unicorn-root’”” would seem to belong more properly to that species than to Aletris farinosa, to which 
the name unicorn-root was given later, and which may thus be called ‘false unicorn-root.” 


20 WILD MEDICINAL PLANTS OF THE UNITED STATES. 


Chamaenerion angustifolium (L.) Scop. Evening-primrose family 
(Onagraceae). 
Synonym.—Epilobium angustifolium L. 
Great willow-herb; wickup. 
Native, perennial herb, 2 to 8 feet high, found in dry soil from Canada to Alaska, 


south to North Carolina, Arizona, and California. Very common from Penn- 
sylvania northward. 


Parts used.—Leaves and root (nonofficial ). 
Chamomile, fetid. See Anthemis cotula. 
Champion-oak. See Quercus rubra. 
Checkerberry. See Gaultheria procumbens and Mitchella repens. 
Cheeseflower. See Malva sylvestris. 
Cheeses. See Malva rotundifolia. 
Chelidonium. See Chelidonium majus. 
Chelidonium majus L. Poppy family (Papaveraceae). 


Chelidonium; celandine; garden-celandine; great celandine; tetterwort. 


Perennial herb, 1 to 2 feet high, growing along fences, roadsides, and in waste 
places; common in the East. Naturalized from Europe. 


Part used.—Entire plant (official in U. 8. P. 1890). 


Chelone glabra L. Figwort family (Scrophulariaceae). 
Balmony; turtle-head; shellflower; snakehead; salt-rheum weed. 


Native, perennial, herbaceous plant, 2 to 3 feet high; in swamps and along 
streams, Newfoundland to Manitoba, south to Florida and Kansas. 


Part used.—Herb, and especially the leaves (nonofficial ). 
Chenopodium. See Chenopodium ambrosioides and C. anthelminticum. 
Chenopodium ambrosioides L. Goosefoot family (Chenopodiaceae). 
Chenopodium; Mexican tea; American wormseed; Jerusalem tea; Spanish tea.’ 


Strong-scented herb, 2 to 3 feet high, annual; naturalized from tropical America, 
and occurring in waste places, meadows, and pastures from New England 
to Florida, west to California. 


Part used.—Fruit (official in U. 8. P. 1890). 


Chenopodium anthelminticum L. Goosefoot family (Chenopodiaceae). 
Chenopodium; wormseed; Jerusalem oak. 


Annual, sometimes perennial, herb, usually taller than C. ambrosioides, natural- 
ized from Europe, and found in waste places from southern New York to 
Wisconsin, south to Florida and Mexico. 


Parts used.—Fruit (official in U. 8. P. 1890). The oil of chenopodium, distilled 
from this plant, is official. 


Chenopodium botrys L. Goosefoot family (Chenopodiaceae). 
Jerusalem oak. 


Annual herb, about 2 feet high, introduced from Europe; found in waste places 
from Nova Scotia to New York and Kentucky, westward to Oregon. 


Parts used.—Herb and seeds (nonofficial). 
Cherry birch. See Betula lenta. 
Cherry, rum-. See Prunus serotina. 
Cherry, wild. See Prunus serotina. 
Chervil, sweet. See Washingtonia longistylis. 
Chestnut. See Castanea dentata. 
Chestnut, American. See Castanea dentata. 
Chestnut, horse-. See Aesculus hippocastanum, 
Chickentoe. See Corallorhiza odontorhiza. 
Chickweed, common. See Alsine media. 


CHICKWEED, RED—CLEMATIS. 21 


Chickweed, red. See Anagallis arvensis. 
Chicory. See Cichorium intybus. 
Chimaphila. * See Chimaphila umbellata. 


Chimaphila umbellata (L.) Nutt. Wintergreen family (Pyrolaceae). 
Chimaphila; pipsissewa; prince’s-pine; bitter wintergreen; rheumatism-weed. 
Small, perennial herb, native in dry, shady woods, especially in pine forests, from 

Nova Scotia to Georgia, west to California. 
Part used.—Leaves (official). 
China-root, American. See Smilax pseudo-china. 
China-root, false. See Smilax pseudo-china. 


Chionanthus virginica L. Olive family (Oleaceae). 
Fringe-tree; old-man’s-beard. 


A shrub or small tree, native in moist thickets from Delaware to Florida and 
Texas. 


Part used.—Bark of root (nonofficial) . 
Chittem-bark. See Rhamnus purshiana. 


Chrysanthemum leucanthemum L. Aster family (Asteraceae). 
Synonym.—Leucanthemum vulgare Lam. 
Oxeye daisy; white daisy. 
Perennial herb, 1 to 3 feet high, naturalized from Europe; occurring in pastures, 


meadows, and waste places in nearly every section of the country, but less 
abundantly in the South and rarely in the West. 


Part used.—Herb (nonofficial). 


Chrysanthemum parthenium (L.) Pers. Aster family (Asteraceae). 
Synonym.—Pyrethrum parthenium Smith. 
Common feverfew; featherfew; febrifuge-plant. 
Perennial herb, naturalized from Europe. Mostly escaped from cultivation; in 

waste places, New Brunswick to New Jersey, and locally in the interior. 

Part used.—Herb (nonofficial). 

Cichorium intybus L. Chicory family (Cichoriaceae). 
Chicory; succory. 
Perennial herb, 1 to 3 feet high, growing in fields, waste places, and along road- 


sides from Nova Scotia to North Carolina, west to Nebraska. Abundant east- 
ward. Naturalized from Europe. 


Part used.—Root (nonofficial) . 
Cicuta maculata L. Parsley family (Apiaceae). 
Water-hemlock; musquash-root; beaver-poison. 


Native perennial, 3 to 6 feet high, stout, erect; poisonous. Found in swamps and 
low grounds from Canada south to Florida and New Mexico. 


Part used.—Leaves (nonofficial ). 

Cimicifuga. See Cimicifuga racemosa. 

Cimicifuga racemosa (L.) Nutt. Crowfoot family (Ranunculaceae). 
Synonym.—Actaea racemosa L. 
Cimicifuga; black snakeroot; black cohosh; squawroot; rattle-root. 


Native, perennial herb, 3 to 8 feet high; in rich soil in shady woods, Maine to 
Georgia, west to Wisconsin and Missouri. Most abundant in the Ohio Valley. 


Parts used.—Rhizome and roots (official). 
Cinquefoil. See Potentilla canadensis. 
Cirsium arvense Scop. Same as Carduus arvensis. 
Cleavers. See Galium aparine. 
Cleaverwort. See Galiwm aparine. 
Clematis. See Clematis virginiana: 


‘ ¥ — a 
29 WILD MEDICINAL PLANTS OF THE UNITED STATES. 


Clematis virginiana L. Crowfoot family (Ranunculaceae). 
Virgin’s-bower; clematis. 
Shrubby, perennial vine; native; found along river banks in hedges and thick- 
ets from Canada to Georgia and Kansas. 


Parts used.—Leaves and flowers (nonofficial). 
Clotbur, spiny. See Xanthium spinosum. 
Clotweed, thorny. See Xanthium spinoswin. 
Clover, bitter. See Sabbatia angularis. 

Clover, meadow-. See Trifolium pratense. 
Clover, purple. See Trifolium pratense. 
Clover, red. See Trifolium pratense. 

Clover, yellow sweet. See Melilotus officinalis. 
Club-moss. See Lycopodium clavatum. 


Cnicus benedictus L. Aster family (Asteraceae). 
Synonyms.—Carduus benedictus Auct.; Centaurea benedicta L. 
Blessed thistle; holy thistle; bitter thistle; spotted thistle; St. Benedict’s-thistle. 


Annual ee 1 to 2 feet high; in waste places, Southern States, and in Califor- 
nia and Utah; introduced from Europe. 


Part used.—Herb (nonofficial). 
Cocash. See Aster puniceus. 
Cocash-weed. See Senecio aureus. 
Cockle-but*on. See Arctiwm lappa. 
Cocowort. See Bursa bursa-pastoris. 
Cohosh, black. See Cimicifuga racemosa. 
Cohosh, blue. See Caulophyllum thalictroides. 
Cohosh, red. See Actaea rubra. 
Cohosh, white. See Actaea alba. 
Colic-root. See Aletris farinosa, Dioscorea villosa, Lacinaria spicata, and L. squarrosa. 


Collinsonia canadensis L. Mint family (Menthaceae). 
Stoneroot; richweed; knobroot; horse-balm. 


Native, perennial herb, about 2 feet high, occurring in rich, moist woods from 
Maine to Wisconsin, south to Florida and Kansas. 


Parts used.—Root and leaves (nonoflicial). 

Colt’s-foot. See Tussilago farfara. 

Colt’s-tail. See Lrigeron canadensis. 

Columbine, European. See Aquilegia vulgaris. 

Columbine, garden-. See Aquilegia vulgaris. 

Columbine, wild. See under Aquilegia vulgaris. 

Columbo, American. See Frasera carolinensis. 

Comfrey. See Symphytum officinale. 

Compass-plant. See Silphium laciniatum. 

Comptonia asplenifolia Gaertn. Same as Comptonia peregrina. 

Comptonia peregrina (L.) Coulter. Bayberry family (Myricaceae). 
Synonyms.—Comptonia asplenifolia Gaertn. ; Myrica asplenifolia L. 
Sweet fern; spleenwortbush; meadow-fern. 


Shrubby plant, about 2} feet high, native; in thin sandy or stony woods and on 
hillsides, Canada to North Carolina, Indiana, and Michigan. 


Parts used.—Leaves and tops (nonofficial). 


Coneflower, pale-purple. See Brauneria angustifolia. 


et ie all 


CONEFLOWER, TALL——CORNUS FLORIDA. 23 


Coneflower, tall. See Rudbeckia laciniata. 

Congo-root. See Psoralea pedunculata. 

Conium. See Conium maculatum. 

Conium maculatum L. Parsley family (Apiaceae). 
Conium; poison-hemlock; spotted parsley; spotted cowbane. 


Biennial herb, 2 to 6 feet high, naturalized from Europe; common in waste 
places, especially in the Eastern and Middle States. Poisonous. 


Parts used.—Full-grown, but unripe, fruit, carefully dried and: preserved (offi- 
cial); leaves (nonofficial). 
Consumptive’s-weed. See Hriodictyon californicum. 
Convyallaria. See Convallaria majalis. , 
Convallaria biflora Walt. Same as Polygonatum biflorum. 
Convallaria majalis L. Lily-of-the-valley family (Convallariaceae). 
Convallaria; lily-of-the-valley. 


A low, perennial herb; indigenous; on the higher mountains from Virginia to 
the Carolinas. 


Parts used.—Rhizome and roots (official); herb and flowers (nonoftiicial ). 
Convallaria racemosa L. Same as Vagnera racemosa. 
Convolvulus panduratus L. Same as Ipomoea pandurata. 
Coolwort. See Tiarella cordifolia. 


Coptis trifolia (L.) Salisb. - Crowfoot family (Ranunculaceae). 
Goldthread; cankerroot; mouthroot; yellowroot. 


Low, native, perennial herb, growing in damp mossy woods and bogs from Can- 
ada and Alaska south to Maryland and Minnesota; most common in the New 
England States, northern New York and Michigan, and in Canada. 


Parts used.—Rhizome and rootlets (nonofficial). 
Corallorhiza odontorhiza ( Willd.) Nutt. Orchid family (Orchidaceae). 
Crawley-root; coralroot; dragon’s-claw; chickentoe. 


Leafless plant, 6 to 15 inches high, found in rich woods from Maine to Florida, 
west to Michigan and Missouri. Native. 


Part used.—Rhizome (nonofficial). 

Coralroot. See Corallorhiza odontorhiza. 

Corn, squirrel-. See Bikukulla canadensis. 

Corn, turkey-. See Bikukulla canadensis. 

Cornel, silky. See Cornus amomum. 

Corn-snakeroot. See Eryngium yuccifolium and Lacinaria spicata. 

Cornus amomum Mill. Dogwood family (Cornaceae). 
Synonym.—Cornus sericea L. 
Red osier; swamp-dogwood; silky cornel; rose-willow. 


Native shrub, 3 to 10 feet high; in low woods and along streams, Canada to 
Florida, west to Texas and the Dakotas. 


Part used.—Bark (nonofficial). 


Cornus circinata L’ Her. Dogwood family (Cornaceae). 
Green osier; round-leaved dogwood. 


Native shrub, 3 to 10 feet high; in shady places, Canada and the northeastern 
United States. , 


Part used.—Bark (nonofficial). 
Cornus florida L. Dogwood family (Cornaceae). 
Flowering dogwood; boxwood. 


Small, native tree or large shrub, growing in woods from Canada to Florida, 
Texas and Missouri. Most abundant in the Middle States. 


Parts used.—Bark of tree and of root, the latter preferred (nonofficial 1 
11072—No. 89—06——4 


24 WILD MEDICINAL PLANTS OF THE UNITED STATES. 


Cornus sericea L. Same as Cornus amomuim. 

Corydalis canadensis Goldie. Same as Bikukulla canadensis. 
Corydalis formosa Pursh. Same as Bikukulla canadensis. 
Cotton-gum. See Nyssa aquatica. 

Cottonweed. See Anaphalis margaritacea. 

Couch-grass. See Agropyron repens. 

Coughweed. See Senecio aureus. 

Coughwort. See Tussilago farfara. 

Cowbane, spotted. See Conium maculatum. 

Cow-lily. See Nymphaea advena. 

Cow-parsnip. See Heracleum lanatum. 


Cracca virginiana L. Pea family (Fabaceae). 
Synonym.— Tephrosia virginiana Pers. 
Devil’s-shoestring; hoary pea; goat’s-rue; catgut. 
Hoary, perennial herb, 1 to 2 feet high, native; occurring in dry, sandy soil 
from New England to Florida, west to Texas and Minnesota. 


Part used.— Root (nonofficial). 

Cramp-bark. See Viburnum opulus. 

Cranberry, high-bush. See Viburnum opulus. 

Cranberry, upland-. See Arctostaphylos uva-ursi. 

Crane’s-bill, spotted. See Geraniwm maculatum. 

Crane’s-bill, wild. See Geranium maculatum. 

Crataegus oxyacantha L. Apple family (Malaceae). 
Hawthorn; hedgethorn; whitethorn; maythorn. 
Shrub or tree, introduced from Europe, and sparingly escaped from cultivation. 
Part used.—Berries (nonofficial). 4 

Crawley-root. See Corallorhiza odontorhiza. 

Crosswort.. See Eupatorium perfoliatum. 

Cucumber-tree. See Magnolia acuminata and M. tripetala. 

Cudweed, low. See Gnaphalium uliginosum. 

Cudweed, marsh-. See Gnaphalium uliginosum. 

Culver’s-physic. See Veronica virginica. 

Culver’s-root. See Veronica virginica. 

Cunila mariana L. Same as Cunila origanoides. 


Cunila origanoides (L.) Britton. Mint family (Menthaceae). 
Synonym.—Cunila mariana L. 
American dittany; stonemint. 


Indigenous, perennial plant, found on dry hills and in dry woods from New York 
to Florida, west to Ohio. 


Part used.—Herb (nonoflicial ). 

Cup-plant. See Silphium perfoliatum. 

Custard-apple. See Asimina triloba. 

Cynoglossum officinale L. Borage family (Boraginaceae). 
Hound’s-tongue; gypsy-flower. 


Biennial herb, about 3 feet high, naturalized from Europe, and occurring in 
waste places from Canada to North Carolina, west to Kansas and Minnesota. 


Parts used.—Leaves and root (nonofficial). 
Cypripedium. See Cypripedium hirsutum and C. parviflorum. 


ioe 


CYPRIPEDIUM HIRSUTUM—DELPHINIUM CONSOLIDA. 25 


Cypripedium hirsutum Mill. Orchid family (Orchidaceae). 
Synonym.— Cypripedium pubescens Willd. 


Cypripedium; large yellow ladies-slipper; yellow moccasin-flower; American 
valerian. 


Herb, 1 to 2 feet high, native in woods and thickets from Nova Scotia south to 
Alabama and west to Nebraska and Missouri. 


Parts used.—Rhizome and roots (official). 


Cypripedium parviflorum Salisb. Orchid family (Orchidaceae). 
Cypripedium; small yellow ladies-slipper. 
Herb, 1 to 2 feet high; native in woods and thickets from British America to 
Georgia, Missouri, and Oregon. 
Parts used.—Rhizome and roots (official). 
Cypripedium pubescens Willd. Same as Cypripedium hirsutum. 


Cytisus scoparius (L.) Link. Pea family (Fabaceae). 
Synonym.—Sarothamnus scoparius Wimm. 
Scoparius; broom; green broom; Scotch broom. 


Stiff, wiry plant, 3 to 5 feet high; naturalized from Europe; growing in dry, sandy 
soil from Massachusetts to Virginia and becoming common in many places in 
the northwestern United States. 


Part used.—Tops (official). 


Daisy, oxeye. See Chrysanthemum leucanthemum. 
Daisy, white. See Chrysanthemum leucanthemum. 
Daisy-fleabane. See Erigeron philadelphicus. 
Damiana. See Turnera microphylla. 

Dandelion. See Taraxacum officinale. 


Daphne mezereum L. Mezereon family (Daphnaceae). 
Synonym.—Mezereum officinarum C. A. Mey. 
Mezereum; mezereon; spurge-laurel; paradise-plant; spurge-olive. 
b ? too} 5 
A very hardy shrub, introduced from Europe and escaped from cultivation in 
Canada and New England. 


Part used.—Bark of this and of other European species of Daphne (official). 


Datura stramonium L. Potato family (Solanaceae). 
Stramonium; jimson-weed; Jamestown-weed; thorn-apple; apple-of-Peru. 


Poisonous weed; annual, 2 to 5 feet high; introduced from the Tropics, and 
occurring in fields and waste places throughout the United States, with the 
exception of the North and West. 


Pavis used.—Leaves (official); seeds (official in U. 8. P. 1890). 
Daucus carota L. Parsley family (Apiaceae). 
Wild carrot; Queen-Anne’s-lace. 


Biennial herb, 2 to 3 feet high; naturalized from Europe; common almost - 
throughout the United States, growing in old fields and along roadsides. 


Parts used.—Root, fruit, and leaves (nonofficial). 
Deerberry. See Gaultheria procumbens and Mitchella repens. 
Deer-laurel. See Rhododendron maximum. 

Deer’s-tongue. See Trilisa odoratissima. 
Deerwood. See Ostrya virginiana. 
Deerwort-boneset. See Hupatorium ageratoides. 


Delphinium consolida L. Crowfoot family (Ranunculaceae). 
Field-larkspur; knight’s-spur: lark-heel. 

An annual herb, about 2 feet high; naturalized from Europe, and found in waste 

places from southern New Jersey and Pennsylvania southward. The indig- 


26 WILD MEDICINAL PLANTS OF THE UNITED STATES. 


Delphinium consolida—Continued. 


enous tall larkspur, Delphinium urceolatuin Jacq. (D. exaliatum Ait.), is used 
for similar purposes. This is found in woods from Pennsylvania to Minnesota, 
south to Alabama and Nebraska. 


Parts used.—Herb and seeds (nonoflicial). 
Delphinium exaltatum Ait. See under Delphinium consolida. 
Delphinium urceolatum Jacq. See under Delphiniwm consolida. 
Devil’s-bit. See Lacinaria scariosa. : 
Devil’s-shoestring. See Cracca virginiana. 
Dewberry. See Rubus procumbens. 
Dewberry, one-flowered. See Rubus villosus. 
Dewberry, southern. See Rubus trivialis. 
Dicentra canadensis Walp. Same as Bikukulla canadensis. 
Digitalis. See Digitalis purpurea. P 
Digitalis purpurea L. Figwort family (Scrophulariaceae). 

Digitalis; foxglove; fairy-fingers; thimbles; lady’s-glove. 

Very handsome biennial plant, 3 to 4 feet high; introduced from Europe as a 
garden plant, and now escaped from cultivation in parts of Oregon, Washing- 
ton, and West Virginia. 

Parts used.—Leaves from plants of second year’s growth, gathered at commence- 
ment of flowering (official). 


Dioscorea villosa L. Yam family (Dioscoreaceae). 
Wild yam; colic-root; rheumatism-root. 


Slender, herbaceous, native vine, growing in moist thickets from Rhode Island 
to Minnesota, south to Floridaand Texas; more common in central and south- 
ern parts of the United States. 


Part used.—Rhizome (nonofficial). 


/ 


Diospyros virginiana L. Ebony family (Diospyraceae). 
Persimmon. 


Indigenous tree, 15 to 50 feet in height; in fields and woods, Rhode Island to 
Kansas, Florida, and Texas. 


Parts used.—Bark and unripe fruit (nonofficial). 


Dirca palustris L. Mezereon family (Daphnaceae). 

Leatherwood; moosewood; American mezereon; wickopy; rope-bark. 

A native shrub, occurring in woods and thickets, New Brunswick to Florida, 
west to Missouri and Minnesota; most common in the Northern and Eastern 
States. 

Part used.—Bark (nonofficial). 

Ditch-stonecrop. See Penthorum sedoides. 
Dittany, American. See Cunila origanoides. 
Dock, bitter. See Rumex obtusifolius. 

Dock, blunt-leaved. See Rumex obtusifolius. 
Dock, broad-leaved. See luwmea obtusifolius. 
Dock, curled. See Rumex crispus. 

Dock, narrow. See Rumer crispus. 

Dock, sour. See Rumex crispus. 

Dock, spatter-. See Nymphaea advena. 
Dock, velvet. See Verbascwm thapsus. 

Dock, yellow. See Rumex crispus. 

Dogbane, spreading. See Apocynum androsaemifolium. 


Dogberry. See Sorbus americana. 


DOG-FENNEL—EMETIC-ROOT. ae 


Dog-fennel. See Anthemis cotula. 

Dog-grass. See Agropyron repens. 

Dog’s-tooth violet. See Lrythronium americanum. 
Dogwood, flowering. See Cornus florida. 

Dogwood, round-leaved. See Cornus circinata. 
Dogwood, swamp-. See Cornus ainomum. 
Dooryard-plantain. See Plantago major. 
Dracontium foetidum L. Same as Spathyema foetida, 
Dragon’s-claw. See Corallorhiza odontorhiza. 
Dropwort, western. See Porteranthus trifoliatus. 


Drosera rotundifolia L. Sundew family (Droseraceae). 
Round-leaved sundew; youthwort. 
Low, perennial herb, growing in bogs and muddy shores of rivers from Canada 
to Florida and California. 
Part used.—Herb (nonofficial). 


Dryopteris filix-mas (L.) Schott. Fern family (Polypodiaceae). 
Synonyms.—Aspidium filiz-mas Sw.; Polypodium filix-mas L. 
Aspidium; male-fern. 
Fern, with leaves 1 to 3 feet long; in rocky woods from Canada to northern 
Michigan, and in the Rocky Mountains to Arizona. 


Part used.—Rhizome (official). 

Dryopteris marginalis (L.) A. Gray. Fern family (Polypodiaceae). 
Synonyms.—Aspidium marginale Sw.; Polypodium marginale L. 
Aspidium; evergreen wood-fern; marginal-fruited shield-fern. 


Fern, with leaves 6 inches to 23 feet long; in rocky woods from Canada south to 
Alabama and Arkansas. 


Part used.—Rhizome (official). 
Duleamara. See Solanum dulcamara. 
Dysentery-weed. See Gnaphaliwm uliginosum. 


Earth-smoke. See Fumaria officinalis. 
Echinacea. See Brauneria angustifolia. 
Echinacea angustifolia DC. Same as Brauneria angustifolia. 
Elder. See Sambucus canadensis. 

Elder, American. See Sambucus canadensis. 
Elder, dwarf. See Aralia hispida. 

Elder, sweet. See Sambucus canadensis. 
Elder, wild. See Aralia hispida. 
Elecampane. See Inula helenium. 

Elk-tree. See Oxydendrum arboreum. 
Elkwood. See Magnolia tripetala. 
Elliott’s-sabbatia. See Sabbatia elliottii. 
Elm. See Ulmus fulva. 

Elm, Indian. See Ulmus fulva. 

Elm, moose-. See Ulmus fulva. 

Elm, red. See Ulmus fulva. 

Elm, slippery. See Ulmus fulva. 
Emetic-root. See Huphorbia corollata. 


28 WILD MEDICINAL PLANTS OF THE UNITED STATES. 


Epigaea repens L. Heath family (Ericaceae). 
Gravel-plant; trailing arbutus; mayflower. 


Small, shrubby, native plant, spreading on the ground in sandy soil, especially 
under ey ergreen trees, from Florida to Michigan and northward. 


Part used.—Leaves (aondleial). 
Epilobium angustifolium L. Same as Chamaenerion angustifolium. 


Epilobium palustre L. Evening-primrose family (Onagraceae). 
Swamp willow-herb; wickup. 
Slender, erect, native herb, 6 to 18 inches high, found in swamps and marshes 

from Canadaand the New England States west to Colorado and Washington. 

Parts used.—Leaves and root (nonoflicial). 

Epiphegus virginiana Bart. Sameas Leptamnium virginianuin. 

Equisetum hyemale L. Horsetail family (Equisetaceae). 
Common scouring-rush; horsetail; shave-grass. 


Rush-like perennial plant, growing in wet places along river banks and borders 
of woods throughout nearly the whole of North America. 


Part used.—Plant (nonofficial). 


Erechtites hieracifolia (L.) Raf. Aster family (Asteraceae). 
Fireweed; pilewort. 
Native, annual herb, 1 to 8 feet high, in woods, fields, and waste places, Canada 
to Florida, Louisiana, and Nebraska. 
Part used.—Herb (nonofficial). 


Erigeron canadensis L. Aster family (Asteraceae). 
Synonym.—Leptilon canadense (L.) Britton.¢ 
Canada fleabane; horseweed; colt’s-tail; prideweed; bitterweed. 


Native, annual weed, 3 inches to 10 feet in height; in fields and meadows, along 
roadsides, and in waste places, almost throughout North America. 


Part used.—Herb (nonofficial); the oil of erigeron, distilled from the fresh, 
flowering herb, is official. 
Erigeron philadelphicus L. Aster family (Asteraceae). 
Philadelphia fleabane; sweet scabious; daisy-fleabane. 


Native, perennial herb, 1 to 3 feet high, in fields and woods throughout North 
America, except extreme North. 


Part used.—Herb (nonoflicial). 
Eriodictyon. See Hriodictyon californicum. 
Eriodictyon californicum (H. & A.) Greene. Waterleaf family 
(Hydrophyllaceae). 
Synonym.—Eriodictyon glutinosum Benth. 
Eriodictyon; yerba santa; mountain-balm; consumptive’s-weed; bear’s-weed. 


Shrubby plant, 2 to 4 feet high, native; grows in clumps in dry situations and 
among rocks throughout California and northern Mexico. 


Part used.—Leaves (official). 
Friodictyon glutinosum Benth. Same as Friodictyon californicum. 
Eryngium yuccaefolium Michx. Same as Lryngium yuceifoliun. 
Eryngium yuccifolium Michx. Parsley family (Apiaceae). 
Synonym.— Eryngium yuccaefolium Michx. 
bib mec iasd Arg button-snakeroot; rattlesnake-weed; rattlesmake-master; corn- 
snakeroot. 


Native, perennial herb, 1 to 5 feet high, growing in swamps and low wet ground 
from the pine barrens of New Jersey west to Minnesota, and south to Texas 
and Florida. 

Part used.—Rhizome (1 smnpsyiis 


aSome authors hold that this stant belouia © to 5 the eae Leptilon and that its name should be 
Leptilon canadense (L.) Britton. The Pharmacopceia is here followed. 


ERYNGO, WATER—EUPHORBIA NUTANS. 29 


Eryngo, water-. See Eryngium yuccifolium. 
Erythronium americanum Ker. Lily family (Liliaceae). 


Yellow adder’s-tongue; dog’s-tooth violet; yellow snowdrop; rattlesnake-violet; 
yellow snakeleaf. 


Native, perennial herb, occurring in moist woods and thickets, Nova Scotia to 
Minnesota, south to Arkansas and Florida. 


Parts used.—Leaves and root (nonofficial). 
Euonymus. See Huonymus atropurpureus. 


Euonymus atropurpureus Jacq. Staff-tree family (Celastraceae). 
Euonymus; wahoo; burningbush; spindle-tree; Indian arrowwood. 


Native shrub or small tree, growing in woods and thickets from Ontario and 
eastern United States west to Montana. 


Part used.—Bark of root (official) . 
Eupatorium. See Hupatorium perfoliatum. 


Eupatorium ageratoides L. f. Aster family (Asteraceae). 


White snakeroot; white sanicle; Indian sanicle; deerwort-boneset; poolwort; 
poolroot; richweed; squaw-weed. 


Erect, perennial herb, 1 to 4 feet high, native; in rich woods from Canada to 
Georgia, west to Nebraska and Louisiana. 


Part used.—Root (nonofificial ). 


Eupatorium aromaticum L. Aster family (Asteraceae). 
Smaller white snakeroot; poolwort; poolroot; wild hoarhound. 


Native, perennial herb, 1 to 2 feet high; in dry soil from ees to 
F lorida, especially thr oughout the Middle States. 


Part used.—Root (nonofficial ). 


Eupatorium perfoliatum L. Aster family (Asteraceae). 
Eupatorium; boneset; thoroughwort; Indian sage; agueweed; crosswort. 


Native, perennial herb, 1 to 5 feet high; in low, wet places from Canada to 
Florida, west to Texas and Nebraska. 


Parts used.—Leaves and flowering tops (official). 


Eupatorium purpureum L. Aster family (Asteraceae). 
Queen-of-the-meadow; gravelroot; Joe-Pye-weed; purple boneset; kidneyroot. 


Native, perennial herb, 3 to 10 feet high; in low grounds from Canada to Florida 
and Texas--~ 


Parts used.—Root and herb (nonofficial). 


Euphorbia corollata L. Spurge family (Euphorbiaceae). 
Flowering spurge; emetic-root; milk-ipecac; snakemilk; purging-root. 
Native, perennial herb, about 3 feet in height, growing in dry fields and woods 
from Ontario to Florida and Minnesota to Texas. 
Part used.—Root (nonofficial). 


Euphorbia hypericifolia A. Gray. Same as Euphorbia nutans. 


Euphorbia ipecacuanhae L. Spurge family (Euphorbiaceae). 
Wild ipecac; ipecac-spurge; American ipecac; Carolina ipecac. 
Native, perennial herb, 4 to 10 inches high; in dry, sandy soil, mostly near the 
coast, from Connecticut to Florida. 
Part used.—Root (nonoflficial). 


Euphorbia nutans Lag. Spurge family (Euphorbiaceae). 
Synonym.—Euphorbia hypericifolia A. Gray. 
Large spotted spurge; black purslane; fluxweed; milk-purslane. 


Native, annual plant, from 3 to 2 feet in height; in rich soils, fields, and thickets 
throughout eastern North America, oxceps extreme north, and extending 
west to the Rocky Mountains. 


Part used.—Herb (nonofficial). 


30 WILD MEDICINAL PLANTS OF THE UNITED STATES. 


Euphorbia pilulifera L. Spurge family (Euphorbiaceae). 
Pill-bearing spurge; snakeweed; Queensland asthma-weed. 


Herbaceous annual, 10 to 15 inches high, occurring from the Gulf States through 
Texas to New Mexico. 


Part used.—Herb (nonofficial ). 
Evening-primrose. See Oenothera biennis. 
Everlasting. See Anaphalis margaritacea. 
Everlasting, large-flowered. See Anaphalis margaritacea. 
Everlasting, pearly.. See Anaphalis margaritacea. 


Eve’s-cup. See Sarracenia flava. 


Fagara clava-herculis (L.) Small. Rue family (Rutaceae). 
Synonym.—Xanthoxylum clava-herculis L. 


Xanthoxylum; southern prickly ash; toothache-tree; yellowthorn; yellow- 
wood; Hercules-club. 


Small, indigenous, very prickly tree, sometimes 45 feet in height, occurring 
along streams from southern Virginia to Florida, west to Texas and Arkansas. 


Parts used.—Bark official under the name ‘‘Xanthoxylum’”’; berries (non- 
official). 


Fagus americana Sweet. Beech family (Fagaceae). 
Synonym.— Fagus ferruginea Ait. 
American beech; beechnut-tree. 


Large, native forest tree, growing in rich soil from Nova Scotia to Florida, west 
to Wisconsin and Texas. 


Parts used.—Bark and leaves (nonofficial ). 

Fagus ferruginea Ait. Same as Fagus americana. 
Fairy-fingers. See Digitalis purpurea. 

Featherfew. See Chrysanthemum parthenium. 
Febrifuge-plant. See Chrysanthemum parthenium. 
Female-fern. See Athyrium filix-foemina and Polypodium vulgare. 
Fennel, dog-. See Anthemis cotula. 

Fern, evergreen wood-. See Dryopteris marginalis. 

Fern, female-. See Athyrium filix-foemina and Polypodium vulgare. 
Fern, lady-. See Athyrium filix-foemina. 

Fern, maidenhair-. See Adiantum pedatum. 

Fern, male-. See Dryopteris filix-mas. 

Fern, marginal-fruited shield-. See Dryopteris marginalis. 
Fern, meadow-. See Comptonia peregrina. 

Fern, parsley-. See Tanacetum vulgare. 

Fern, royal. See Osmunda regalis. 

Fern, sweet. See Comptonia peregrina. 

Fernroot. See Polypodium vulgare. 

Feverbush. See Benzoin benzoin and Ilex verticillata. 
Feverfew, common. See Chrysanthemum parthenium. 
Feverroot. See Triostewm perfoliatum. 

Fevertwig. See Celastrus scandens. 

Field-balm. See Glecoma hederacea. 

Field-larkspur. See Delphinium consolida, 

Field-sorrel. See Rumex acetosella. 


FIGWORT, MARYLAND—FRAXINUS NIGRA. 3] 


Figwort, Maryland. See Scrophularia marilandica. 
Fir, balsam-. See Abies balsumea. 

Fireweed. See Erechtites hieracifolia. 

Fit-plant. See Monotropa uniflora. 

Fitroot. See Monotropa uniflora. 

Fivefinger. See Potentilla canadensis. 

Flag, blue. See Iris versicolor. 

Flag, cattail-. See Typha latifolia. 

Flag, sweet-. See Acorus calamus. 

Flag, water-. See Tris versicolor. 

Flag-lily. See Iris versicolor. 

Flannel-leaf. See Verbascum thapsus. 

Fleabane, Canada. See Erigeron canadensis. 
Fleabane, daisy-. See Erigeron philadelphicus. 
Fleabane, Philadelphia. See Erigeron philadelphicus. 
Fluxweed. See Euphorbia nutans. 

Flytrap. See Sarracenia purpurea. 

Foamflower. See Tiarella cordifolia. 

Foxglove. See Digitalis purpurea. 

Fragaria virginiana Duchesne. Rose family (Rosaceae). 


Virginia strawberry; scarlet strawberry. 
y; 


Native, perennial herb, occurring in dry soil from Canada to Georgia, west to 
Indian Territory and Minnesota. 


Part used.—Leaves (nonofficial). 
Frankenia grandifolia Cham. &Schlecht. Frankenia family (Frankeniaceae). 
Yerba reuma. 


Native, perennial herb, 8 to 13 inches high, common in salt marshes and sandy 
localities near the coast in California. 


Part used.—Herb (nonofficial). 

Frasera carolinensis Walt. Gentian family (Gentianaceae). 
Synonym.—Frasera walteri Michx. 
American columbo; Indian lettuce; meadowpride; pyramid-flower. 


Smooth, perennia! herb, 3 to 8 feet high, found in dry soil from New York to 
Wisconsin, south to Georgia and Kentucky. 


Part used.—Root (nonofficial). 
Frasera walteri Michx. Same as Frasera carolinensis. 
Fraxinus acuminata Lam. Same as Fraxinus americana. 
Fraxinus alba Marsh. Same as Fraxinus americana. 


Fraxinus americana L. Olive family (Oleaceae). 
| Marsh; Fraxinus acuminata Lam. 


White ash; cane-ash. 


Large, native forest tree, in rich woods from Nova Scotia to Minnesota, south to 
Florida and Texas. Occurs chiefly in the Northern States and Canada. 


Part used.— Bark (nonofficial). 

Fraxinus nigra Marsh. Olive family (Oleaceae). 
Synonym.—Fraxinus sambucifolia Lam. 
Black ash; hoop-ash. 


Native tree, 40 to 70 feet in height, occurring in swamps and wet woods from 
Canada to Virginia and Arkansas. 


Part used.—Bark (nonofficial). 
11072—No. 89—06——5 


32 ‘WILD MEDICINAL PLANTS OF THE UNITED STATES. 


Fraxinus sambucifolia Lam. Same as Fraxinus nigra. 
Fringe-tree. See Chionanthus virginica. 

Frost-plant. See Helianthemum canadense. 
Frostweed. See Helianthemum canadense. 

Frostwort. See Helianthemum canadense. 
Fuller’s-herb. See Saponaria officinalis. 


Fumaria officinalis L. Poppy family (Papaveraceae). 
Fumitory; hedge-fumitory; earth-smoke. 
Annual plant, 10 to 15 inches high, adventive from Europe and found in waste 
places about dwellings, in cultivated land, and on ballast, Nova Scotia to the 
Gulf States. 


Part used.—Herb (nonofficial). 
Fumitory. See Fumaria officinalis. 
Fumitory, hedge-. See Fumaria officinalis. 


Gagroot. See Lobelia inflata. 
Gale, sweet. See Myrica gale. 
Galium aparine L. Madder family (Rubiaceae). 

Cleavers; goose-grass; cleaverwort; bedstraw; catchweed. 

Annual plant, with weak, procumbent stem, 2 to 6 feet long, growing in shady 
thickets and margins of woods, New Brunswick south to Florida and Texas. 
Naturalized from Europe. 

Part used.—Herb of this and of other species of Galium (nonoflficial). 


Gallweed. See Gentiana quinquefolia. 
Garden-balm. See Melissa officinalis. 
Garden-celandine. See Chelidonium majus. 
Garden-columbine. See Aquilegia vulgaris. 
Garden-valerian. See Valeriana officinalis. 
Garget. See Phytolacca decandra. 


Gaultheria procumbens L. Heath family (Ericaceae). 

Wintergreen; checkerberry; mountain-tea; teaberry; deerberry. 

Small, native perennial, with evergreen leaves, found in sandy soils in cool, 
damp woods, especially under evergreen trees, in Canada and the northeastern 
United States. ; 

Part used.—Leaves (nonoflicial) ; the oil of gaultheria, distilled from the leaves, 
is official. 

Gay-feather. See Lacinaria scariosa and L. spicata. 
Gelsemium. See Gelsemiuwm sempervirens. 
Gelsemium sempervirens (L.) Ait. f. Logania family (Loganiaceae). 

Gelsemium; yellow jasmine; Carolina jasmine; wild woodbine. 


Twining, shrubby perennial, native, growing on low ground in woods and 
thickets from eastern Virginia to Florida and Texas, mostly near the coast. 


Parts used.—Rhizome and roots (official). 
Gemfruit. See Tiarella cordifolia. 
Gentian, American. See Gentiana saponaria, 
Gentian, blue. See Gentiana saponaria. 
Gentian, five-flowered. See Gentiana quinquefolia. 
Gentian, horse-. See Triosteum perfoliatum. 
Gentian, marsh-. See Gentiana villosa. 
Gentian, snake-. See Nabalus serpentarius. 


GENTIAN, SOAPWORT—GINSENG. 33 


Gentian, soapwort-. See Gentiana saponaria. 
Gentian, stiff. See Gentiana quinquefolia. 
Gentian, straw-colored. See Gentiana villosa. 
Gentian, striped. See Gentiana villosa. 
Gentian, white. See Triosteum perfoliatum. 
Gentiana catesbaei Walt. Same as Gentiana saponaria. 
Gentiana ochroleuca Froel. Same as Gentiana villosa. 
Gentiana quinqueflora Lam. Same as Gentiana quinquefolia. 
Gentiana quinquefolia L. Gentian family (Gentianaceae). 
Synonym.—Gentiana quinqueflora Lam. 
Stiff gentian; five-flowered gentian; agueweed; gallweed. 


Native, annual plant, 1 to 2 feet in height, growing in pastures and other open 
situations from Maine to Michigan, south to Florida and Missouri. 


Parts used.—Root and herb (nonofficial). 


Gentiana saponaria L. Gentian family (Gentianaceae). 
Synonym.—Gentiana catesbaei Walt. 
American gentian; blue gentian; soapwort-gentian. 


Native, perennial herb, 1 to 23 feet high; in wet soil, Ontario to Minnesota, 
south to Louisiana and Florida. 


Part used.—Root (nonoflicial). 

Gentiana villosa L. Gentian family (Gentianaceae). 
Synonym.—Gentiana ochroleuca Froel. 
Striped gentian; straw-colored gentian; marsh-gentian; Sampson’s-snakeroot. 


Native, perennial herb, 6 to 18 inches high; in shaded places, Middle and 
Southern States. 


Part used.— Root (nonofficial ). 
Geranium. See Geranium maculatum. 


Geranium maculatum L. Geranium family (Geraniaceae). 
Geranium; wild crane’s-bill; spotted crane’s-bill; wild geranium; spotted gera- 
nium; alum-root. 


Native, perennial herb, 1 to 13 feet high; found in low grounds and open woods 
from Canada south to Georgia and Missouri. 


Part used.—Rhizome (official ). 
Geranium, spotted. See Geranium maculatum. 
Geranium, wild. See Geranium maculatum. 


Geum rivale L. Rose family (Rosaceae). 
Water-avens; purple avens. 
Native, perennial herb, 1 to 2 feet high, occurring in swamps and wet meadows 
from Canada to Pennsylvania and Colorado, especially in the Northern and 
Middle States. 


Parts used.—Rhizome and rootlets (nonofficial ). 
Ghostflower. See Monotropa uniflora. 
(rllenia trifoliata Moench. Same as Porteranthus trifoliatus. 
Gill-over-the-ground. See Glecoma hederacea, 
Ginger, Indian. See Asarum canadense. 
Ginger, wild. See Asarwm canadense. 
Gingerroot. See Tussilago farfara. 


Ginseng. See Panax quinquefolium. 


34 WILD MEDICINAL PLANTS OF THE UNITED STATES. 


Glecoma hederacea L. Mint family (Menthaceae). 

Synonym.—Nepeta glechoma Benth. 

Ground-ivy; gill-over-the-ground; catfoot; field-balm. 

Low, perennial herb, with creeping stem. Naturalized from Europe and found 
in waste places, woods, and thickets from Newfoundland to Minnesota, south 
to Georgia and Kansas. : 

Part used.—Herb (nonofiicial ). 

Globeflower. See Cephalanthus occidentalis. 
Gnaphalium margaritaceum L. Same as Anaphalis margaritacea. 
Gnaphalium obtusifolium L. Aster family (Asteraceae). 

Synonym.—Gnaphalium polycephalum Michx. 

Sweet balsam; life-everlasting; sweet life-everlasting; white balsam. 


Native, herbaceous annual, 1 to 2 feet high; in dry, open places and old fields 
from Nova Scotia and Manitoba south to Florida and Texas. 


Part used.—Herb (nonofficial). 
Gnaphalium polycephalum Michx. Same as Gnaphalium obtusifolium. 


Gnaphalium uliginosum L. Aster family (Asteraceae). 
Mouse-ear; low cudweed; marsh-cudweed; wartwort; dysentery-weed. 


Annual herb, 2 to 8 inches high, occurring in damp soil from Newfoundland to 
Minnesota, south to Indiana and Virginia; apparently naturalized from Europe. 


Part used.—Herb (nonofficial). 
Gnaphalium undulatum Walt. Same as Pterocaulon undulatum. 
Goat’s-rue. See Cracca virginiana. 
Goldenrod, anise-scented. See Solidago odora. 
Goldenrod, fragrant-leaved. Sée Solidago odora. 
Goldenrod, sweet. See Solidago odora. 
Goldenseal. See Hydrastis canadensis. 
Goldthread. See Coptis trifolia. 
Goodyera pubescens R. Br. Same as Peramium pubescens. 
Goodyera repens R. Br. Same as Peramium repens. 
Goose-grass. See Galiwm aparine. 
Grape, Oregon. See Berberis aquifolium. 
Grape, Rocky Mountain. See Berberis aquifolium. 
Gravel-plant. See Epigaea repens. 
Gravelroot. See Hupatorium purpureum. 
Gravel-weed. See Onosmodium virginianum. 
Greenbrier, long-stalked. See Smilax pseudo-china. 
Grindelia. See Grindelia robusta and G. squarrosa. 


Grindelia robusta Nutt. ; Aster family (Asteraceae ). 
Grindelia; gum-plant. 
Perennial herb, about 14 feet high, native in the States west of the Rocky 
Mountains. 
Parts used.—Leaves and flowering tops (official). 
Grindelia, scaly. See Grindelia squarrosa. 
Grindelia squarrosa (Pursh) Dunal. Aster family (Asteraceae). 


Grindelia; scaly grindelia; broad-leaved gum-plant. 


Perennial herb, 1 to 2 feet high, native; occurring on the plains and prairies 
from the Saskatchewan to Minnesota, Texas, and California. 


Parts used.—Leaves and flowering tops (official). 


GROMWELL, VIRGINIA FALSE—HELENIUM AUTUMNALE. 35 


Gromwell, Virginia false. See Onosmodium virginianum 
Ground-centaury. See Polygala nuttallii. 
Ground-ivy. See Glecoma hederacea. 
Ground-raspberry. See Hydrastis canadensis. 
Ground-squirrel pea. See Jeffersonia diphylla. 
Gum, cotton-. See Nyssa aquatica. 

Gum, red. See Liquidambar styraciflua. 

Gum, star-leaved. See Liquidambar styraciflua. 
Gum, sweet-. See Liquidambar styraciflua. 

Gum, tupelo. See Nyssa aquatica. 

Gum-plant. See Grindelia robusta. 

Gum-plant, broad-leaved. See Grindelia squarrosa. 
Gypsy-flower. See Cynoglossum officinale. 
Gypsy-weed. See Lycopus virginicus. 


Hackmatack. See Larix laricina. 

Haircap-moss. See Polytrichum juniperinum. 

Hamamelis. See Hamamelis virginiana. 

Hamamelis virginiana L. Witch-hazel family (Hamamelidaceae). 
Hamamelis; witch-hazel; winterbloom; snapping hazel. 
Indigenous shrub, found in low, damp woods from New Brunswick to Minne- 

sota, south to Florida and Texas. 

Parts used.—Leaves (collected in autumn), bark, and twigs (official). 


Hardhack. See Spiraea tomentosa. 
Hart’s-thorn. See Rhamnus cathartica. 
Haw, black. See Viburnum prunifolium. 
Hawkweed, early. See Hieracium venosum. 
Hawthorn. See Crataegus oxryacentha. 
Hazel, snapping. See Hamamelis virginiana. 
Heal-all. See Prunella vulgaris and Scrophularia marilandica. 
Healing-herb. See Symphytum officinale. 
Heart-liverleaf. See Hepatica acuta. 
Heartsease. See Viola tricolor. 

Hedeoma. See Hedeoma pulegioides. 


Hedeoma pulegioides (L.) Pers. Mint family (Menthaceae). 
Hedeoma; American pennyroyal; tickweed; squawmint. 


Low, native, annual plant, 6 to 12 inches high, growing in barren woods and dry 
fields, Nova Scotia to Minnesota, south to Nebraska and Florida. 


Parts used.—Leaves and flowering tops, and the volatile oil distilled from these, 
are official. 
Hedge-fumitory. See Fumaria officinalis. 
Hedgethorn. See Crataegus oxyacantha. 


Helenium autumnale L. Aster family (Asteraceae). 
Sneezeweed; sneezewort; swamp-sunflower. 


Native perennial, 2 to 3 feet high, growing in swamps, wet fields, and meadows, 
Canada to Florida and Arizona. 


Part used.—Herb (nonofiicial). 


36 WILD MEDICINAL PLANTS OF THE UNITED STATES. 


Helianthemum canadense (L.) Michx. Rock-rose family (Cistaceae). 
Frostweed; frostwort; frost-plant; Canadian rock-rose. 


Native, perennial herb, about one foot in height; in dry, sandy soil, Maine to 
Wisconsin, south to North Carolina and Kentucky. 


Part used.—Herb (nonofficial). 
‘ Hellebore, American. See Veratrum viride. 
Hellebore, green. See Veratrum viride. 
Hellebore, swamp-. See Veratrum viride. 
Helmetpod. See Jeffersonia diphylla. 
Helonias dioica Pursh. Same as Chamaelirium luteum. 
Hemlock. See Tsuga canadensis. 
Hemlock, poison-. See Conium maculautum. 
Hemlock, water-. See Cicuta maculata. 
Hemlock-spruce. See Tsuga canadensis. 
Hemp, black Indian. See Apocynum cannabinum. 
Hemp, Canadian. See Apocynum cannabinum. 
Hemp, white Indian. See Asclepias incarnata. 
Henbane. See Hyoscyamus niger. 
Hepatica acuta (Pursh) Britton. Crowfoot family (Ranunculaceae). 
Synonym.— Hepatica acutiloba DC. 
Heart-liverleaf; sharp-lobed liverleaf; liverwort. 


Perennial herb, 4 to 9 inches high, found in woods from Quebee and Ontario, 
south to Georgia (but rare near the coast), west to Iowa and Minnesota. 


Part used.—Leaves (nonofficial ). 

Hepatica acutiloba DC. Same as Hepatica acuta. 

Hepatica hepatica (L.) Karst. Crowfoot family (Ranunculaceae). 
Synonym.—Hepatica triloba Chaix. 
Round-lobed liverleaf; kidney-liverleaf; liverwort. 


Perennial herb, 4 to 6 inches high; in woods from Nova Scotia to northern 
Florida, west to Iowa and Missouri; less common than the heart-liverleaf. 


Part used.—Leaves (nonofficial ). 

Hepatica triloba Chaix. Same as Hepatica hepatica. 

Heracleum lanatum Michx. Parsley family (Apiaceae). 
Masterwort; cow-parsnip; youthwort. 


Native, perennial herb, 3 to 5 feet high, growing in moist meadows and culti- 
vated ground from C anada south to North ( ‘arolina, Utah, and California. 


Parts used.—Root, leaves, and seeds (nonoflficial). 
Hercules-club. See Fagara clava-herculis. 
Heuchera americana L. ' S$axifrage family (Saxifragaceae ). 


Alum-root; American sanicle. 


Native, perennial herb, 2 to 4 feet in height; in shady, rocky woodlands from 
Connecticut to Minnesota, south to Alabama and Louisiana. 


Part used.—Root (nonofficial ). 

Hickory, shellbark-. See /icoria ovata. 

Hicoria ovata ( Mill.) Britton. Walnut family (Juglandaceae). 
Synonym.—Carya alba Nutt. 
Shagbark, shellbark-hickory. 


Large, native tree, sometimes 120 feet in height; in rich soil from Quebee to 
southern Ontario and Minnesota, south to Florida and Texas. 


Parts used.—Bark and leaves (nonoflicial). 


HYDRASTIS CANADENSIS. 37 


HIERACIUM VENOSUM 


Hieracium venosum lL. Chicory family (Cichoriaceae). 

Early hawkweed; rattlesnake-weed; bloodwort; striped bloodwort. 

Perennial herb, 1 to 2 feet high, native; occurring in dry woods and thickets 
from Maine to Georgia, west to Nebraska; more common in the northern and 
eastern United States. 

Parts used.— Leaves and root (nonofficial ). 


Highbelia. See Lobelia siphilitica. 

Hive-vine. See Mitchella repens. 

Hoarhound. See Marrubium vulgare. 
Hoarhound, water-. See Lycopus virginicus. 
Hoarhound, wild: See Lupatorium aromaticum. 
Hog-potato. See Ipomoea pandurata. 
Hog’s-bean. See Hyoscyaimus niger. 

Hogweed. See Ambrosia artemisiaefolia, 

Holly, American. See Ilex opaca. 

Holly, white. See Ilex opaca. 

Honeybloom. See Apocynuin androsaemifolium. 
Hoodwort. See Scutellaria lateriflora. 
Hoop-ash. See Fraxinus nigra. 

Hop-hornbeam. See Ostrya virginiana. 
Hop-tree. See Ptelea trifoliata. 

Hornbeam, hop-. See Ostrya virginiana. 
Horse-balm. See Collinsonia canadensis. 
Horse-chestnut. See Aesculus h ippocastanum. 
Horsefly-weed. See Baptisia tinctoria. 
Horsefoot. See Tussilago farfara. 
Horse-gentian. See Triostewm perfoliatum. 
Horseheal. See Inula helenium. 

Horsemint. See Monarda fistulosa and M. punctata. 
Horse-nettle. See Solanum carolinense. 
Horsetail. See Equisetum.hyemale. 

Horseweed. See Erigeron canadensis. 
Hound’s-tongue. See Cynoglossum officinale. 
Hydrangea. See Hydrangea arborescens. 


Hydrangea arborescens L. Hydrangea family (Hydrangeaceae). 

Hydrangea; wild hydrangea; seven-barks. 

Indigenous shrub, 5 or 6 feet in height; on rocky river banks from southern 
New York to Florida, west to lowa and Missouri; very abundant in the val- 
ley of the Delaware. 

Part used.—Root (nonofificial) . 


Hydrangea, wild. See Hydrangea arborescens. 
Hydrastis. See Hydrastis canadensis. 


Hydrastis canadensis L. Crowfoot family (Ranunculaceae). 
Hydrastis; goldenseal; yellowroot; ground-raspberry; orangeroot; yellow 
puccoon. 


Perennial herb, about 1 foot in height, native in rich soil in shady woods, 
southern New York to Minnesota, south to Georgia and Missouri, but prin- 
cipally in Ohio, Indiana, Kentucky, and West Virginia. 

Parts used.—Rhizome and roots (official). 


38 WILD MEDICINAL PLANTS OF THE UNITED STATES. 


Hyoscyamus. See Hyoscyamus niger. 
Hyoscyamus niger L. Potato family (Solanaceae). 
Hyoscyamus; henbane; hog’s-bean; insane-root. 


Biennial herb, 6 inches to 2 feet high, sparingly naturalized from Europe, in 
waste places from Nova Scotia to Ontario, New York, and Michigan. 


Parts used.—Leaves and flowering tops from plants of second year’s growth 
(official); seeds are also used (nonofficial). 
Hypericum perforatum L. St. John’s-wort family (Hypericaceae). 
John’s-wort; common St. John’s-wort. 


Herbaceous perennial, 1 to 2 feet high, naturalized from Europe; common in 
fields and waste places throughout almost the entire United States, except the 
Southern States. 


Part used.—Herb (nonofiicial ). 
Hyssop. See Hyssopus officinalis. 
Hyssop, wild. See Verbena hastata. 
Hyssop-skulleap. See Scutellaria integrifolia. 
Hyssopus officinalis L. - Mint family (Menthaceae). 

Hyssop. 

Perennial herb, 1 to 3 feet high, naturalized from Europe, and found along road- 
sides and in waste places from Ontario and Maine to North Carolina, and on 
the Pacific coast. 

Part used.—Herb (nonofificial). 


Ilex opaca Ait. Holly family (Aquifoliaceae). 
American holly; white holly. 


Native tree, 20 to 40 feet in height, with evergreen leaves; in moist woodlands, 
Maine to Florida, and west to Missouri and Texas; most abundant in the 
Atlantic States. 


Parts used.—Leaves and bark (nonofficial). 
Ilex verticillata (L.) A. Gray. Holly family (Aquifoliaceae). 
Synonym.—Prinos verticillata L. 
Black alder; feverbush; Virginia winterberry. 
A native shrub, growing in moist woods and along banks of streams from Noya 
Scotia to Florida, west to Wisconsin and Missouri. 
Parts used.—Bark and berries (nonofficial ). 
Impatiens aurea Muhl. Jewelweed family (Impatientaceae). 
Synonym.—Impatiens pallida Nutt. 
Jewelweed; pale touch-me-not; snapweed; wild celandine. 


Native, annual plant, 2 to 4 feet high, found in rich soil in moist, shady places 
from Quebec to Oregon, south to Georgia and Kansas. 


Part used.—Herb (nonofificial). 
Impatiens biflora Walt. Jewelweed family (Impatientaceae ). 
Synonym.— Impatiens fulva Nutt. 
Jewelweed; spotted touch-me-not; snapweed; silverleaf. 
Native, annual plant, 2 to 5 feet high, growing in rich soil in moist, shady places 
from Canada to Alaska and Oregon, south to Florida and Missouri; more com- 
mon than the pale touch-me-not. 


Part used.—Herb (nonofficial ). 
Impatiens fulva Nutt. Same as Impatiens biflora. 
Impatiens pallida Nutt. Same as Impatiens aurea, 
Indian-cup. See Silphium perfoliatum. 
Indian-paint. See Sanguinaria canadensis. 


Indian-physic. See Porteranthus trifoliatus. 


INDIAN-PIPE—JASMINE, YELLOW. — 39 


Indian-pipe. See Monotropa uniflora. 
Indian-root. See Aralia racemosa. 
Indigo, American. See Baptisia tinctoria. 
Indigo, wild. See Baptisia tinctoria. 
Indigo, vellow. See Baptisia tinctoria. 
Indigo-weed. See Baptisia tinctoria. 
Inkberry. See Phytolacca decandra. 
Inkroot. See Limonium carolinianum. 
Insane-root. See Hyoscyamus niger. 
Inula. See Inula helenium. 


Inula helenium L. Aster family (Asteraceae). 
Inula; elecampane; horseheal; scabwort. 


Rough, perennial herb, 3 to 6 feet high, naturalized from Europe, and found 
along roadsides and in fields and pastures from Nova Scotia to North Carolina, 
westward to Missouri and Minnesota. 


Part used.—Root (official in U. 8. P. 1890). 
Ipecac, American. See Huphorbia ipecacuanhae. 
Ipecac, Carolina. See Euphorbia ipecacuanhae. 
Ipecac, false. See Porteranthus trifoliatus. 
Ipecac, milk-. See Huphorbia corollata. 
Ipecac, wild. See Euphorbia ipecacuanhae and Triostewm perfoliatum. 
Ipecac-spurge. See Huphorbia ipecacuanhae. 


Ipomoea pandurata (L.) Meyer. Morning-glory family (Convolvulaceae). 
Synonym.-—Convolvulus panduratus L. 
Manroot; man-of-the-earth; wild potato; hog-potato; wild jalap. 
Native perennial, with trailing stems 2 to 12 feet long; in dry fields or on hills 
from Connecticut to Michigan, south to Florida and Texas. 


Part used.—Root (nonofficial ). 

Iris. See Jris versicolor. 

Iris versicolor L. Iris family (Iridaceae). 
Tris; blue flag; flag-lily; liver-lily; water-flag; snake-lily. 


Native, perennial plant, 2 to 3 feet high, found in wet, marshy localities from 
Newfoundland to Manitoba, south to Florida and Arkansas. 


Parts used.—Rhizome and roots (official in U. 8. P. 1890). 
Ironwood. See Ostrya virginiana. 
Ivy, American. See Parthenocissus quinquefolia. 
Ivy, ground-. See Glecoma hederacea. 


Ivy, poison-. See Rhus radicans and R. toxicodendron. 


Jack-in-the-pulpit. See Arisaema triphyllum. 
Jacob’s-ladder. See Polemonium reptans. 
Jacob’s-ladder, American. See Sinilax herbacea. 
Jalap, wild. See Jpomoea pandurata. 
James-tea. See Ledum groenlandicuin. 
Jamestown-weed. See Datwra stramonium. 
Jasmine, Carolina. See Gelsemiwm sempervirens. 
oe Jasmine, yellow. See Gelsemium sempervirens. 


40 WILD MEDICINAL PLANTS OF THE UNITED STATES. 


Jeffersonia diphylla (L.) Pers. Barberry family (Berberidaceae). 
Twinleaf; rheumatism-root; helmetpod; yellowroot; ground-squirrel pea. 


Native, perennial plant, 8 to 14 inches in height, growing in woods and near 
streams from New York to Virginia, westward to Wisconsin. 


Part used.—Rhizome (nonofficial ). 
Jewelweed. See Impatiens aurea and I. biflora. 
Jimson-weed. See Datura stramonium. 
Job’s-tears, wild. See Onosmodium virginianum. 
Joe-Pye-weed. See Eupatorium purpureumn. 
John’s-wort. See Hypericum perforatum. 
Judas-tree. See Cercis canadensis. 

Juglans. See Juglans cinerea. 


Juglans cinerea L. Walnut family (Juglandaceae). 
Juglans; butternut; white walnut. 


Indigenous tree, 20 to 50 feet in height, common in rich woods from New Bruns- 
wick to North Dakota, south to Georgia, Mississippi, and Arkansas. 


Part used.—Bark of root, collected in autumn (official in U. 8. P. 1890). 
Juniper. See Juniperus communis. 


Juniperus communis L. Pine family (Pinaceae). 
Juniper. ; 


Evergreen shrub or low tree, common on dry, sterile hills from Canada south 
to New Jersey, west to Nebraska, and in the Rocky Mountains to New Mexico. 


Part used.—Fruit (nonofficial). The oil of juniper, distilled from the fruit, is 
official. 
Juniperus sabina L. Pine family (Pinaceae). 
Sabina; savin; shrubby red cedar. 
\ shrub, usually procumbent, seldom more than 4 feet in height, occurring in 
rocky places in the northern United States. 
Part used.—Tops, and the oil of savin, distilled from the fresh tops, are official. 


Juniperus virginiana L. Pine family (Pinaceae). 
Red cedar; red savin. 
A tree, sometimes 100 feet in height, common in dry soil from Canada to Florida 
and Arizona. 


Parts used.—Leaves and ‘‘cedar apples’’ (nonofficial). 


Kalmia angustifolia L. Heath family (Ericaceae). 
Sheep-laurel; lambkill; calfkill; narrow-leaved laurel. 
Native, evergreen shrub, about 3 feet high, growing in moist soil from Canada 
south to Georgia. 
Part used.—Leaves (nonofficial ). 
Kalmia latifolia L. Heath family (Ericaceae). 
Mountain-laurel; calico-bush; broad- leaved laurel; sheep-laurel. 


Native, evergreen shrub, 10 to 20 feet high, growing in sandy or rocky soil 
from New Brunswick to Ohio, Florida, and Louisiana. 


Part used.—Leaves (nonoflicial). 
Kidney-liverleaf. See Hepatica hepatica. 
Kidneyroot. See Lupatorium purpureum. 
Knight’s-spur. See Delphinium consolida, 
Knobroot. See Collinsonia canadensis. 


Knotweed, biting. See Polygonum hydropiper. 


* 


KOELLIA MONTANA—LARIX AMERICANA. 4] 


Koellia montana (Michx.) Kuntze. Mint family (Menthaceae). 
Synonym.—Pycnanthemum montanum Michx. 
Thin-leaved mountain-mint. 


Native perennial, 2 to 3 feet high, found in woods from southern Virginia to 
Georgia and Alabama. 


Part used.—Herb (nonofficial) . 


Koellia pilosa ( Nutt.) Britton. Mint family (Menthaceae). 
Synonym.—Pycnanthemum pilosum Nutt. 
Hairy mountain-mint. 


Native perennial, 1 to 23 feet high, occurring in prairies and dry woods from 
Ohio to Georgia, west to Missouri and Arkansas. 


Part used.—Herb (nonofficial). 


Lacinaria scariosa {L.) Hill. Aster family (Asteraceae). 
Synonym.—Liatris scariosa Willd. 
Blue blazingstar; large button-snakeroot; rattlesnake-master; gay-feather; 
devil’s-bit. 
Native, perennial herb, 4 to’5 feet high, found in dry woods and sandy fields 
from Maine to Florida, west to Texas and Nebraska. 
Part used.—Root (nonofficial). 
Lacinaria spicata (L.) Kuntze. Aster family (Asteraceae). 
Synonym.—Liatris spicata Willd. 


Dense button-snakeroot; colic-root; prairie-pine; gay-feather; rattlesnake- 
master; corn-snakeroot; backache-root. 


Native, perennial herb, 2 to 5 feet high, in moist places from Massachusetts to 
Florida, west to Wisconsin and Arkansas. 


Part used.—Root (nonofficial ). 


Lacinaria squarrosa (L.) Hill. Aster family (Asteraceae). 
Synonym.—Liatris squarrosa Willd. 
Scaly blazingstar; colic-root; rattlesnake-master (in the South). 


Native, perennial herb, 2 to 3 feet high, in dry soil, Ontario to Florida, west to 
Nebraska and Texas. 


Part used.—Root (nonofiicial). 


Lactuca canadensis L. Chicory family (Cichoriaceae) . 
Synonym.—Lactuca elongata Muhl. 
Wild lettuce; tall lettuce; wild opium; trumpet-milkweed. 


Annual or biennial plant, 3 to 10 feet in height, native in moist, open places, 
British America south to Georgia and Louisiana. 


Part used.—Herb (nonofficial). 
Lactuca elongata Muhl. Same as Lactuca canadensis. 
Ladies-slipper, large yellow. See Cypripedium hirsutum. 
Ladies-slipper, small yellow. See Cypripedium parviflorum. 
Lady-fern. See Athyrium filix-foemina. 
Lady’s-glove. See Digitalis purpurea. 
Lambkill. See Kalmia angustifolia. 
Lappa. See Arctium lappa. 
Lappa major Gaertn. Same as Arctium lappa. 
Lareh, American. See Larix laricina. 
Larch, black. See Larix laricina. 
Larix americana Michx. Same as Larix laricina. 


42 WILD MEDICINAL PLANTS OF THE UNITED STATES. 


Larix laricina (Du Roi) Koch. Pine family (Pinaceae). 
Synonym.—Larix americana Michx. 
Tamarack; American larch; hackmatack; black larch. 


A tall, slender tree, native in swampy woods and moist places from Canada 
south to New Jersey, Indiana, and Minnesota. 


Part used.—Bark (nonofficial). 
Lark-heel. See Delphiniwm consolida. 
Larkspur, field-. See Delphiniwm consolida. 
Larkspur, tall. See under Delphinium consolida. 
Laurel, broad-leaved. See Kalmia latifolia. 
Laurel, deer-. See Rhododendron maximum. 
Laurel, great. See Rhododendron maximum. 
Laurel, mountain-. See Kalmia latifolia. 
Laurel, narrow-leaved. See Kalmia angustifolia. 
Laurel, rose-. See Rhododendron maximum. 
Laurel, sheep-. See Kalmia angustifolia and K. latifolia. 
Laurel, spurge-. See Daphne mezereum. 
Laurel, swamp-. See Magnolia virginiana. 
Laurus benzoin L. Same as Benzoin benzoin. 
Lavender, sea-. See Limonium carolinianum. 
Leafcup, yellow. See Polymnia uvedalia. 
Leatherwood. See Dirca palustris. 


Ledum groenlandicum Oeder. Heath family (Ericaceae). 
Synonym,.—Ledum latifolium Ait. 
. € 
Labrador tea; continental tea; James-tea. 


Evergreen shrub, 1 to 4 feet high, native in cold bogs and damp mountain 
woods, northern part of the United States and in Canada. 


Part used.—Leaves (nonofficial). 

Ledum latifolium Ait. Same as Ledum groenlandicum. 

Lemon, wild. See Podophyllum peltatum. 

Lemon-balm. See Melissa officinalis. 

Leonurus cardiaca L. Mint family (Menthaceae). 
Motherwort; lion’s-tail; throwwort. 


Perennial plant, 2 to 5 feet high, naturalized from Europe, and occurring in 
fields and waste places from Nova Scotia to North Carolina westward to 
Nebraska. 


Part used.—Herb (nonoflicial ). 


Leptamnium virginianum (L.) Raf. Broomrape family (Orobanchaceae). 
Synonyms.—LEpiphegus virginiana Bart.; Orobanche virginiana L. 
Beechdrops; cancerroot. 


Plant 6 inches to 2 feet in height, parasitic upon the roots of beech trees from 
New Brunswick to Florida, west to Michigan and Louisiana. 


Part used.—W hole plant (nonofficial). 
Leptandra. See Veronica virginica. 
Leptandra virginica (L.) Nutt. Same as Veronica virginica. 
Leptilon canadense (L.) Britton. Same as EFrigeron canadensis. 
Lettuce, Indian. See Frasera carolinensis. 
Lettuce, tall. See Lactuca canadensis. 


Lettuce, white. See Nabalus albus and N. serpentarius. 


LETTUCE, WILD—LIRIODENDRON TULIPIFERA. 43 


Lettuce, wild. See Lactuca canadensis. 

Leucanthemum vulgare Lam. Same as Chrysanthemum leucanthemum. 
Leverwood. See Ostrya virginiana. 

Liatris odoratissima Michx. Same as Trilisa odoratissima. 

Liatris scariosa Willd. Same as Lacinaria scariosa. 

Liatris spicata Willd. Same as Lacinaria spicata. 

Tiatris squarrosa Willd. Same as Lacinaria squarrosa. 

Life-everlasting. See Anaphalis margaritacea and Gnaphalium obtusifolium. 
Life-everlasting, sweet. See Gnaphalium obtusifolium. 

Liferoot. See Senecio aureus. 


Ligustrum vulgare L. Olive family (Oleaceae). 
Privet; primwort; prim. 
A shrub, 5 or 6 feet high, introduced from Europe; escaped from cultivation and 
grows wild in woods and along roadsides from Ontario to Pennsylvania and 
North Carolina. 


Part used.—Leaves (nonofficial). 
Lily, cow-. See Nymphaea advena. 
Lily, flag-. See Iris versicolor. 
Lily, large yellow pond-. See Nymphaea advena. 
Lily, liver-. See Jris versicolor. 
Lily, snake-. See Iris versicolor. 
Lily, sweet-scented water-. See Castalia odorata. 
Lily, water-. See Castalia odorata. 
Lily, white pond-. See Castalia odorata. 
Lily-of-the-valley. See Convallaria majalis. 
Lime, Ogeechee. See Nyssa ogeche. 
Limonium carolinianum (Walt.) Britton. 
Plumbago family (Plumbaginaceae). 
Synonym.—Statice caroliniana Walt. 


. Marsh-rosemary; inkroot; sea-lavender; cankerroot. 


Native, perennial herb, 1 to 2 feet high, in salt meadows on the Atlantic and 
Gulf coasts. 


Part used.—Root (nonofficial). 
Linden, American. See Tilia americana. 
Lindera benzoin Meissn. Same as Benzoin benzoin. 
Lion’s-foot. See Nabalus albus and N. serpentarius. 
Lion’s-tail. See Leonurus cardiaca. 


Liquidambar styraciflua L. Witch-hazel family (Hamamelidaceae). 
Sweet-gum; star-leaved gum; red gum. 


Large, native tree, 80 to 140 feet high, in moist woods from Connecticut to 
Florida, Illinois, and Missouri. Most common near the coast in the Middle 
and Southern States. 


Parts used.—Bark and resin (nonofficial). 


Liriodendron tulipifera L. Magnolia family (Magnoliaceae). 
Tulip-tree; yellow poplar; whitewood; tulip-poplar; canoewood. 
An indigenous tree, 60 to 190 feet in height, growing in rich woods from New 


England to Florida, west to Michigan and Arkansas; reaches greatest size in 
the Middle and Southern States. 


Part used.—Bark of trunk and of root (nonofficial). 


44 WILD MEDICINAL PLANTS OF THE UNITED STATES. 


Lithospermum virginianum L. Same as Onosmodium virginianum. 

Liverleaf, heart-. See Hepatica acuta. 

Liverleaf, kidney-. See Hepatica hepatica. 

Liverleaf, round-lobed. See Hepatica hepatica. 

Liverleaf, sharp-lobed. See Hepatica acuta. 

Liver-lily. See Jris versicolor. 

Liverwort. See Hepatica acuta and H. hepatica. 

Lobelia. See Lobelia inflata. 

Lobelia, blue. See Lobelia siphilitica. 

Lobelia cardinalis L. Bellflower family (Campanulaceae). 
Cardinal-flower; red cardinal; red lobelia. 


Native, perennial herb, 2 to 4 feet high, with showy scarlet flowers; in moist 
soil from British America south to Florida and Texas. 


Part used.—Herb (nonofficial). 
Lobelia, great. See Lobelia siphilitica. 
Lobelia inflata L. Bellflower family (Campanulaceae). 


Lobelia; Indian tobacco; gagroot; vomitwort;. bladderpod. 

Native, annual, herbaceous plant, 1 to 3 feet high, poisonous; in dry soil, fields, 
old pastures, and along roadsides from Canada to Georgia, Nebraska, and 
Arkansas. 

Parts used.—Leaves and tops, collected after a portion of the capsules have 
become inflated (official). The seeds are also used (nonofficial). 

Lobelia, red. See Lobelia cardinalis. 
Lobelia siphilitica L. Bellflower family (Campanulaceae). 

Blue cardinal-flower; great lobelia; blue lobelia; highbelia. 


Native, perennial herb, about 1 to 3 feet high, growing in moist soil from Ontario 
to Ge rgia, west to Louisiana and the Dakotas. 


Part used.—Herb (nonofificial). 
Locust, black. See Robinia pseudacacia. 
Locust, yellow. See Robinia pseudacacia. 
Locust-plant. See Cassia marilandica. 
Locust-tree. See Robinia pseudacacia. 
Lycopodium. See Lycopodium clavatum. ; 
Lycopodium clavatum L. Club-moss family (Lycopodiaceae). 
Lycopodium; club-moss; stag’s-horn. 
Native perennial, with trailing stem, growing in dry situations in woods from 
Canada to North Carolina, Michigan, and Washington. 
Part used.—Spores of this or of other species of Lycopodium (official). 
Lycopus virginicus L. Mint family (Menthaceae). 
Bugleweed; sweet bugle; water-bugle; gypsy-weed; water-hoarhound. 


Indigenous, perennial herb, 10 to 20 inches in height; in wet, shady places from 
Canada to Florida, Missouri, and Nebraska. 


Part used.—Herb (nonoflicial ). 


Madweed. See Scutellaria lateriflora. 
Magnolia acuminata L. ~ Magnolia family (Magnoliaceae). 
Cucumber-tree; mountain-magnolia; blue magnolia. 


Native tree, 60 to 80 feet in height, occurring in the mountainous regions from 
New York to Georgia. More abundant in the Southern States. 


Part used.—Bark (nonofficial). 


Magnolia, blue. See Magnolia acuminata. 


a ie, 


adel 


MAGNOLIA GLAUCA—MARSH-CUDWEED. 45 


Magnolia glauca L. Same as Magnolia virginiana. 

Magnolia, mountain-. See Magnolia acuminata. 

Magnolia, sweet. See Magnolia virginiana. 

Magnolia tripetala L. Magnolia family (Magnoliaceae). 
Synonym.— Magnolia umbrella Lam. 


Cucumber-tree; umbrella-tree; elk wood. 


A small native tree, not more than 40 feet high, growing in rather moist, rich * 
soil; widely distributed in the Appalachian Mountain region, but nowhere 
very common. 


Part used.—Bark (nonofficial). 
Magnolia umbrella Lam. Same as Magnolia tripetala. 
Magnolia virginiana L. Magnolia family (Magnoliaceae). 
Synonym.—Magnolia glauca L. 
White bay; sweet bay; sweet magnolia; beaver-tree; swamp-sassafras; swamp- 
laurel. 
A native tree, averaging about 25 feet in height, growing in swamps and 
morasses, Massachusetts to the Gulf of Mexico. 
Part used.—Bark (nonofficial). 


Maidenhair-fern. See Adiantum pedatum. 
Male-fern. See Dryopteris filix-mas. 
Mallow, common. See Malva sylvestris. 
Mallow, dwarf. See Malva rotundifolia. 
Mallow, high. See Malva sylvestris. 
Mallow, low. See Malva rotundifolia. 
Mallow, running. See Valva rotundifolia. 


Malva rotundifolia L. Mallow family (Malvaceae). 
Low mallow; running mallow; cheeses; dwarf mallow. 


Annual or biennial procumbent plant, naturalized from Europe, and widely dis- 
tributed as a weed in waste places. 


Parts used.—Leaves and flowers (nonofficial ). 


Malva sylvestris L. Mallow family (Malvaceae). 
High mallow; common mallow; cheeseflower. 
Biennial herb, adventive from Europe; sparingly distributed in the United 


States and Canada, growing in waste places and along roadsides. 

Part used.—F lowers (nonofficial ). 
Mandrake, American. See Podophyllum peliatum. 
Mandrake, wild. See Podophyllum peltatum. 
Man-of-the-earth. See Ipomoea pandurata. 
Manroot. See Ipomoea pandurata. 
Manzanita. See Arctostaphylos glauca. 
Maple, red. See Acer rubrum. 
Maple, swamp-. See Acer rubrum. 
Maple, vine-. See Menispermum canadense. 
Marrubium. See Marrubium vulgare. 


Marrubium vulgare L. Mint family (Menthaceae). 
Marrubium; hoarhound. 


Bushy, perennial herb, | to 3 feet high, naturalized from Europe, and growing 
in dry, sandy soil, in fields and waste places, from Maine southward to Texas 
and westward to California and Oregon. 


Parts used.—Leaves and flowering tops (official). 
Marsh-cudweed. See Gnaphalium uliginosum. 


46 WILD MEDICINAL PLANTS OF THE UNITED STATES. 


Marsh-gentian. See Gentiana villosa. 

Marshmallow. See Althaea officinalis. 

Marsh-rosemary. See Limonium carolinianum. 

Marsh-trefoil. See Menyanthes trifoliata. 

Maruta cotula DC. Same as Anthemis cotula. 

Masterwort. See Angelica atropurpurea and Heracleum lanatum. 
May-apple. See Podophyllum peltatum. 

Mayflower. See Hpigaea repens. 

May-pops. See Passiflora incarnata. 

Maythorn. See Crataegus oxyacantha, 

Mayweed. See Anthemis cotula. 

Meadow-clover. See Trifolium pratense. 

Meadow-fern. See Comptonia peregrina. 

Meadowpride. See Frasera carolinensis. 

Meadow-scabish. See Aster puniceus. 

Meadowsweet, pink. See Spiraea tomentosa. 

Mealy-tree. See Viburnum dentatum. 

Melilot, yellow. See Melilotus officinalis. 

Melilotus officinalis (L.) Lam. Pea family (Fabaceae). 


Yellow melilot; yellow sweet clover. 


Annual or biennial herb, 1 to 3 feet high, introduced from Europe, and occurring 
in waste places throughout the eastern United States. 


Parts used.—Leaves and flowering tops (nonofficial). 
Melissa. See Melissa officinalis. 


Melissa officinalis L. Mint family (Menthaceae). 
Melissa; balm; lemon-balm; garden-balm; sweet balm. 
Perennial herb, 10 to 20 inches high, naturalized from Europe, and growing in 
waste places, fields, and woods from Maine to Georgia. 
Parts used.—Leaves and tops (official in U.S. P. 1890). 
Menispermum. See Menispermum canadense. 


Menispermum canadense L. Moonseed family (Menispermaceae). 
Menispermum; yellow parilla; Canada moonseed; Texas sarsaparilla; vine-maple. 


Native, perennial, woody climber, found in woods along streams from Canada 
to Georgia and Arkansas. 
Parts used.—Rhizome and roots (official in U. 8. P. 1890). 


Mentha piperita. See Mentha piperita L. 


Mentha piperita L. Mint family (Menthaceae) . 

Mentha piperita; peppermint. : 

Aromatic, perennial herb, 1 to 2 feet high, naturalized from Europe, and oecur- 
ring in damp places from Nova Scotia to Minnesota, south to Florida and Ten- 
nessee. Cultivated principally in Michigan and New York. 

Parts used.—Leaves and flowering tops, and the oil of peppermint distilled from 
these, are official. 

Mentha spicata L. Mint family (Menthaceae). 

Synonym.—Mentha viridis L. 

Mentha viridis; spearmint. 

Aromatic, perennial herb, 1 to 2 feet high, naturalized from Europe, and_grow- 
ing in moist fields and waste places from Nova Scotia to Utah, south to Florida 
and Kansas. Also cultivated. 

Parts used.—Leaves and flowering tops, and the oil of spearmint distilled from 
these, are official. 


MENTHA VIRIDIS—MONARDA FISTULOSA. 47 


Mentha viridis. See Mentha spicata. 
Mentha viridis L. Same as Mentha spicata. 


Menyanthes trifoliata L. Buck-bean family (Menyanthaceae). 
Buck-bean; bog-bean; marsh-trefoil; water-shamrock. 
Indigenous, perennial plant, about 1 foot in height, found in spongy, boggy 
soils and swamps from Canada and Alaska south to Pennsylvania, Minnesota, 
and California. 


Parts used.—Rhizome’and leaves (nonofticial ). 

Mezereon. See Daphne mezerewm. 

Mezereon, American. See Dirca palustris. 

Mezereum. See Daphne mezerewm. 

Mezereum officinarum C. A. Mey. Same as Daphne mezereum. 

Micromeria chamissonis ( Benth.) Greene. Mint family (Menthaceae). 
Synonym.—Micromeria douglasii Benth. 
Yerba buena. 


A trailing, perennial herb, common in woods along the Pacific coast of the United 
States. 


Part used.—Plant (nonofiicial ). 
Micromeria douglasii Benth. Same as Micromeria chamissonis. 
Milfoil. See Achillea millefolium. 
Milk-ipecac. See Euphorbia corollata. 
Milk-purslane. See Euphorbia nutans. 
Milkweed, common. See Asclepias syriaca. 
Milkweed, swamp-. See Asclepias incarnata. 
Milkweed, trumpet-. See Lactuca canadensis. 
Milkwort, Nuttall’s-. See Polygala nuttalli. 
Mint, hairy mountain-. See Koellia pilosa. 
Mint, mountain-. See Monarda didyma. 
Mint, thin-leaved mountain-. See Koellia montana. 
Mistletoe. See Phoradendron flavescens. 
Mistletoe, American. See Phoradendron flavescens. 


Mitchella repens L. Madder family (Rubiaceae). 
Squaw-vine; checkerberry; partridgeberry; deerberry; hive-vine; squawberry. 
Small, creeping, evergreen herb, common in moist woods from Nova Scotia to 

Minnesota, south to Florida and Arkansas. 
Part used.—Plant (nonofficial) . 


Miterwort, false. See Tiarella cordifolia. 

Moccasin-flower, yellow. See Cypripedium hirsutum. 

Mohawk-weed. See Uvularia perfoliata. 

Monarda didyma L. Mint family (Menthaceae). 


Bee-balm; Oswego tea; mountain-mint; scariet balm. 


Native perennial, 2 to 3 feet high, growing in moist soil, especially along streams, 
from New Brunswick to Michigan and_.south to Georgia. 
' Part used.—Herb (nonoflicial). 


Monarda fistulosa L. Mint family (Menthaceae). 
Wild bergamot; horsemint. 


Native perennial, 2 to 3 feet high, found on dry hills and in thickets from 
Ontario south to Florida and Louisiana. 


Part used.—Herb (nonofficial). 


48 WILD MEDICINAL PLANTS OF THE UNITED STATES. 


Monarda punctata L. Mint family (Menthaceae). 
Horsemint. 


Native, perennial herb, 2 to 3 feet high, found in dry, sandy fields from New 
York to Florida, west to Wisconsin and Texas. 


Part used.—Herb (nonofficial ). ’ 


Monotropa uniflora L. Indian-pipe family (Monotropaceae). 
Indian-pipe; fit-plant; fitroot; ghostflower; pipe-plant. 
A curious plant, white in all its parts, growing in rich, moist woods from Canada 
to Florida, westward to Washington and California. 


Part used.—Root (nonofficial). 
Moonseed, Canada. See Menispermum canadense. 
Moose-elm. See Ulmus fulva. 
Moosewood. See Dirca palustris. 
Mortification-root. See Althaea officinalis. 
Moss, club-. See Lycopodium clavatum. 
Moss, haircap-. See Polytrichum juniperinum. 
Motherwort. See Leonwrus cardiaca. 
Mountain-ash, American. See Sorbus americana. 
Mountain-balm. See Hriodictyon californicum. 
Mountain-laurel. See Kalmia latifolia. 
Mountain-magnolia. See Magnolia acuminata. 
Mountain-mint. See Monarda didyma. 
Mountain-mint, hairy. See Koellia pilosa. 
Mountain-mint, thin-leaved. See Koellia montana. 
Mountain-sumac. See Sorbus americana. 
Mountain-tea. See Gaultheria procumbens. 
Mouse-ear. See Gnaphalium uliginosum. 
Mouthroot. See Coptis trifolia. 
Mugwort, common. See Artemisia vulgaris. 
Mullein. See Verbascum thapsus. 
Musquash-root. See Cicuta maculata. 
Mustard, black. See Brassica nigra. 
Mustard, brown. See Brassica nigra. 
Mustard, red. See Brassica nigra. 
Mustard, white. See Sinapis alba. 
Mustard, yellow. See Sinapis alba. 
Myrica asplenifolia L. Same as Comptonia peregrina. 
Myrica cerifera L. Bayberry family (Myricaceae). 
Bayberry; wax-myrtle; candleberry; waxberry. 
Grows in sandy swamps or wet woods from Florida and Texas northward to 


Maryland. In the South it is a small evergreen tree, becoming in its north- 
ward range a tall, semi-deciduous shrub, or a dwarfed and deciduous shrub. 


Parts used.—Bark of root, leaves, and berries (nonofficial). 
Myrica gale L. Bayberry family (Myricaceae). 
Sweet gale; Dutch myrtle; bog-myrtle; golden osier. 


Indigenous shrub, growing in swamps and along streams from Canada and 
Alaska to Virginia and Washington. 


Parts used.—Leayes and buds (nonofficial). 


MYRTLE, BOG—NYSSA CAPITATA. 49 


Myrtle, bog-. See Myrica gale. 
Myrtle, Dutch. See Myrica gale. 
Myrtle, wax-. See Myrica cerifera. 


Nabalus albus (L.) Hook. Chicory family (Cichoriaceae). 
Synonym.—Prenanthes alba L. 
Lion’s-foot; rattlesnake-root; white lettuce; white canker-weed. 
Native, perennial herb, 2 to 4 feet high, common in rich, moist woods from 
Canada to Georgia and Kentucky. 
Part used.—Plant (nonoflicial ). 
Nabalus serpentarius (Pursh) Hook. Chicory family (Cichoriaceae). 
Synonym.—Prenanthes serpentaria Pursh. 
Lion’s-foot; canker-weed; white lettuce; rattlesnake-root; snake-gentian. 
Native, perennial herb, about 2 feet high, growing in dry, sandy soil in fields 
and thickets from Ontario to Florida and Alabama. 


Part wsed.—Plant (nonoflicial). 
Nannybush. See Viburnum lentago. 
Necklace-weed. See Actaea alba and Onosmodiuin virginianum. 


Nepeta cataria L. Mint family (Menthaceae). 
Catnip; catmint. 


Common, perennial weed, 2 to 3 feet high, naturalized from Europe; found in 
waste places and cultivated land from Canada to Minnesota, south to Virginia 
and Arkansas. 


Part used.—Herb (nonofticial). 
Nepeta glechoma Benth. Same as Glecoma hederacea. 
Netleaf-plantain. See Peramium pubescens. 
Netleaf-plantain, smaller. See Peramium repens. 
Nettle, bull-. See Solanum carolinense. 
Nettle, great. See Urtica dioica. 
Nettle, horse-. See Solanum carolinense. 
Nettle, stinging. See Urtica dioica. 


Niggerhead. See Brauneria angustifolia. 

Nightshade, woody. See Solanum duleamara. 

Nuphar advena R. Br. Same as Nymphaea advena. 

Nuttall’s-milkwort. See Polygala nuttallii. 

Nymphaea advena Soland. Water-lily family (Nymphaeaceae). 


Synonym.—Nuphar advena R. Br. ‘ 

Large yellow pond-lily; cow-lily; spatter-dock; beaverroot. 

An aquatic plant, found in ponds and slow streams from Canada to Florida, and 
westward to the Rocky Mountains. 


Part used.—Rhizome (nonoflicial) . 

Nymphaea odorata Dryand. Same as Castalia odorata. 

Nyssa aquatica L. Dogwood family (Cornaceae). 
Synonym.—Nyssa uniflora Wang. 
Large tupelo; cotton-gum; tupelo gum. 


A large, native tree, occurring in swamps from southern Virginia to Florida, 
west to Texas and Missouri. 


Part used.—Root wood (nonofficial). 
Nyssa capitata Walt. Same as Nyssa ogeche. 


; 


50 WILD MEDICINAL PLANTS OF THE UNITED STATES. 


Nyssa ogeche Marsh. Dogwood family (Cornaceae). 
Synonym.—Nyssa capitata Walt. 
Sour tupelo; Ogeechee lime. 


A small tree, growing in swamps near the seacoast from southern South Caro- 
lina to Florida. « 


Part used.—Root wood (nonofficial ). 


Nyssa uniflora Wang. Same as Nyssa aquatica. 


Oak, champion-. See Quercus rubra, 

Oak, Jerusalem. See Chenopodium anthelminticum and C. botrys. 

Oak, poison-. See Rhus radicans and fh. toxicodendron. 

Oak, red. See Quercus rubra. 

Oak, Spanish. See Quercus rubra. 

Oak, stone-. See Quercus alba. 

Oak, white. See Quercus alba. 

Oenothera biennis L. Evening-primrose family (Onagraceae). 
Synonym.—Onagra biennis (L.) Scop. 
ivening-primrose; tree-primrose; night willow-herb. 


Annual or biennial plant, 2 to 5 feet high, common in fields and waste places 
from Labrador to Florida, west to the Rocky Mountains. Native. 


Part used.—Plant (nonofficial). 

Old-man’s-beard. See Chionanthus virginica. 

Olive, spurge-. See Daphne mezereum. 

Onagra biennis (L.) Scop. Same as Oenothera biennis. 

Onosmodium virginianum (L.) DC. Borage family (Boraginaceae). 
Synonym.—Lithospermum virginianum LL. 
Virginia false gromwell; gravel-weed; necklace-weed; pearl-plant; wild Job’s- 

tears. 
Rough-hairy, native, perennial herb, 1 to 2 feet high; in dry, hilly grounds 
from the New England States to Florida, Kansas, and Texas. 

Parts used.—Root and seeds (nonoflficial ). 

Opium, wild. See Lactuca canadensis. 

Orangeroot. See Hydrastis canadensis. 

Orobanche virginiana L. Same as Leptamnium virginianum. 

Osier, golden. See Myrica gale. 

Osier, green. See Cornus circinata. 

Osier, red. See Cornus amomum. 

Osmorrhiza longistylis DC. Same as Washingtonia longistylis. 

Osmunda regalis L. -Royal fern family (Osmundaceae). 
Royal fern; buckhorn-brake. 


A tall, native fern, with fronds 3 to 4 feet high, occurring in swamps and marshes 
from Canada to Florida and Mississippi. 


Part used.—Rhizome (nonoflficial). 
Ostrya virginiana ( Mill.) Willd. Birch family (Betulaceae). 
Hop-hornbeam; ironwood; deerwood; leyerwood. 


Native tree, 25 to 30 feet in height, growing in rich woods, Canada and eastern 
United States. 


Part used.—Bark (nonoflicial). ; 


a 


=e 


OXALIS ACETOSELLA—-PENNYROYAL, AMERICAN. 51 


Oxalis acetosella L. Wood-sorrel family (Oxalidaceae). 
White wood-sorrel; shamrock; sour trefoil. 


Small, native, perennial herb, found in cold, damp woods, Canada south to 
*Michigan and North Carolina. 


Part used.—Herb (nonofficial). 
Oxeye daisy. See Chrysanthemum leucanthemum. 


Oxydendrum arboreum (L.) DC. Heath family (Ericaceae). 
Synonym.—Andromeda arborea L. 
Sourwood; sorrel-tree; elk-tree. 


Native tree, sometimes 40 to 50 feet in height, growing in rich woods from Ohio 
to Maryland, south to Alabama and Florida. 


Parts used.—Leaves and bark (nonoffcial) 


Palmetto, saw-. See Serenoa serrulata. 


Panax quinquefolium L. Ginseng family (Araliaceae). 
Ginseng. 
Native, perennial herb, about 1 foot in height, found in rich, shady woods 
from the Middle and Northern States south to Alabama and Georgia. 


Part used.—Root (nonofiicial ). 
Pansy. See Viola tricolor. 
Papoose-root. See Caulophyllum thalictroides. 
Paradise-plant. See Daphne mezerewmn. 
Parilla, yellow. See Menispermum canadense. 
Parsley, spotted. See Coniwm maculatum. 
Parsley-fern. See Tanacetum vulgare. 
Parsnip, cow-. See Heraclewm lanatum. 


Parthenocissus quinquefolia (L.) Planch. Grape family (Vitaceae). 
Synonym.—Ampelopsis quinquefolia Michx. 
American ivy; Virginia creeper. 
A common, woody vine, native in woods and thickets from Canada to Florida 
and Texas. 


Parts used.—Bark and young twigs (nonoflicial). 
Partridgeberry. See Mitchella repens. 
Pasqueflower, American. See Pulsatilla hirsutissima. 


Passiflora incarnata L. Passion-flower family (Passifloraceae). 
Passion-flower; passion-vine; may-pops. 
Climbing, perennial plant, native in dry soil from Virginia to Florida, westward 
to Missouri and Arkansas. 
Parts used.—Root and stem base (nonofficial). 


Passion-flower. See Passiflora incarnata. 
Passion-vine. See Passiflora incarnata. 
Paul’s-betony. See Veronica officinalis. 

Pawpaw, North American. See Asimina triloba. 
Pea, ground-squirrel. See Jeffersonia diphylla. 
Pea, hoary. See Cracca virginiana. 

Pea, turkey-. See Bikukulla canadensis. 
Pearl-plant. See Onosmodium virginianum. 
Pencil-flower. See Stylosanthes biflora. 
Pennyroyal, American. See Hedeom« pulegioides. 


52 WILD MEDICINAL PLANTS OF THE UNITED STATES. 


Penthorum sedoides L. Virginia stonecrop family (Penthoraceae). 
Virginia stonecrop; ditch-stonecrop. 
Native, perennial herb, about 1 foot in height, growing in ditches and swamps 

from New Brunswick to Minnesota, south to Florida and Texas, 

Part used.—Herb (nonofficial. ) 

Pepper, water-. See Polygonum hydropiper. 

Peppermint. See Mentha piperita. 

Pepper-plant. See Polygonum hydropiper. 

Peramium pubescens ( Willd.) MacM. Orchid family (Orchidaceae). 
Synonym.—Goodyera pubescens R. Br. 
Downy rattlesnake-plantain; rattlesnake-weed; netleaf-plantain; scrofula-weed. 


Native, perennial herb, 8 to 12 inches in height, occurring in rich woods from 
Newfoundland to Minnesota, south to Florida and Tennessee. Most common 
southward. 


Part used.—Plant (nonofficial). 


Peramium repens (L.) Salisb. Orchid family (Orchidaceae). 
Synonym.—Goodyera repens R. Br. 


White plantain; lesser rattlesnake-plantain; smaller netleaf-plantain; squirrel- 
ear. 


A smaller plant than P. pubescens, but very similar to it and more common 
northward. 


Part used.—Plant (nonofificial). 


Persimmon. See Diospyros virginiana. 

Phoradendron fiavescens (Pursh) Nutt. Mistletoe family (Loranthaceae). 
Synonym.— Viscum flavescens Pursh. 
Mistletoe; American mistletoe. 


Parasitic shrub, found on deciduous-leaved trees from New Jersey to Missouri, 
south to Florida and Texas. 


Parts used.—Leaves and branches (nonoflicial ). 

Phytolaeca. See Phytolacca decandra. 

Phytolacca americana L. Same as Phytolacca decandra. é 

Phytolacca decandra L.4 Pokeweed family (Phytolaccaceae). 
Synonym.—Phytolacca americana L.4 
Phytolacca; poke; pokeweed; garget; scoke; inkberry. : 


Native, perennial herb, with large and branching stem, 6 to 10 feet high; in rich, 
moist soil, Maine to Minnesota, south to Florida and Texas. 


Parts used.—Root collected in autumn (official); fruit (official in U. 8. P. 1890); 

leaves (nonofficial). 
Picea mariana (Mill.) B. 8. P. Pine family (Pinaceae). 

Synonym.—Abies nigra Dest. 

Black spruce; spruce-gum tree. 

Indigenous, evergreen tree, 40 to 80 feet in height, growing on elevated situa- 
tions and in cold bogs from Canada south along the mountains to North Car- 
olina, and to Minnesota. 


Parts used.—Branches, and the essence obtained from the same (nonoflicial). 
Pilewort. See Erechtites hieracifolia and Scrophularia marilandica. 
Pilotweed. See Silphium laciniatum. 
Pimpernel. See Pimpinella saxifraga. 
Pimpernel, red. See Anagallis arvensis. 


Pimpernel, scarlet. See Anagallis arvensis. 


a Phytolacea americana L. by right of priority’ should be accepted, but P. decandra L. is used in con- 
formity with the Pharmacopeeia. 


PIMPINELLA SAXIFRAGA—POKE. 53 


Pimpinella saxifraga L. Parsley family (Apiaceae). 
Burnet-saxifrage; bennet; pimpernel. 


Erect, perennial herb, 1 to 2 feet high, adventive from Europe, and found in 
waste places in eastern Pennsylvania, at several localities in the valley of the 
Delaware, and in Ohio. 


Part used.—Root (nonofficial ). 
Pine, northern. See Pinus strobus. 
Pine, prairie-. See Lacinaria spicata. 
Pine, prince’s-. See Chimaphila umbellata 
Pine, Weymouth. See Pinus strobus. 
Pine, white. See Pinus strobus. 
Pink, rose-. See Sabbatia angularis. 
Pinkroot. See Spigelia marilandica. 
Pinkroot, Indian. See Spigelia marilandica. 
Pinkroot, Maryland. See Spigelia marilandica. 


Pinus strobus L. Pine family (Pinaceae). 
White pine; northern pine; Weymouth pine. 
Large, indigenous forest tree, sometimes 175 feet in height, growing in woods 
from Canada south to Georgia and Iowa. 


Part used.—Bark (nonofficial). 
Pipe-plant. See Monotropa uniflora. 
Pipsissewa. See Chimaphila wmbellata. 
Pitcher-plant. See Sarracenia purpurea. 


Plantago major L. Plantain family (Plantaginaceae). 
Common plantain; dooryard-plantain; greater plantain. 


Perennial herb, 1 to3 feet high, naturalized from Europe; common in fields and 
waste places and along roadsides nearly throughout North America. 


Parts used.—Root and leaves (nonofficial). 
Plantain, common. See Plantago major. 
Plantain, dooryard-. See Plantago major. 
Plantain, downy rattlesnake-. See Peramium pubescens. 
Plantain, greater. See Plantago major. 
Plantain, lesser rattlesnake-. See Peramiwm repens. 
Plantain, netleaf-. See Peramium pubescens. 
Plantain, smaller netleaf-. See Peramium repens. 
Plantain, white. See Peramium repens. 
Pleurisy-root. See Asclepias tuberosa. 
Podophyllum. See Podophyllum peltatum. 


Podophyllum peltatum L. Barberry family (Berberidaceae). 
. Podophyllum; May-apple; wild mandrake; American mandrake; wild lemon. 


Native, perennial herb, | to 14 feet high, found in low, rich woods from Canada 
to Minnesota, south to Florida and Texas. 


Part used.—Rhizome (official). 
Poison-hemlock. See Conium maculatum. 
Poison-ivy. See Rhus radicans and R. toxicodendron. 
Poison-oak. See Rhus radicans and R. toxicodendron. 
Poison-vine. See Rhus radicans. 
Poke. See Phytolacca decandra. 


54 WILD MEDICINAL PLANTS OF THE UNITED STATES. 


Pokeweed. See Phytolacca decandra. 
Polar-plant. See Silphium laciniatum. 
Polecat-weed. See Spathyema foetida. 


Polemonium reptans L. Phlox family (Polemoniaceae). 
American Greek valerian; abscess-root; sweetroot; Jacob’s-ladder. 


Native, perennial herb, 12 to 20 inches high, growing in woods and damp ground 
from New York to Minnesota, south to. Georgia and Missouri. 


Part used.—Root (nonofficial). 


Polygala nuttallii T. & G. Milkwort family (Polygalaceae). 
Nuttall’s-milkwort; ground-centaury. 


Slender, erect, annual herb, 6 to 12 inches high, native in dry, sandy soil from 
Massachusetts to North Carolina, west to Alabama and Missouri. 


Part used.—Herb (nonofficial). 


Polygala senega L. Milkwort family (Polygalaceae). 
Senega; Seneca snakeroot. 


Native, perennial herb, 8 to 12 inches high, found in rocky woods and on hillsides 
from New Brunswick and western New ‘England to Minnesota, south to North 
Carolina and Missouri. 


Part used.—Root (official). 
Polygonatum biflorum ( Walt.) EI. Lily-of-the-valley family 
(Convallariaceae). 
Synonyms.—Convallaria biflora Walt.; Salomonia biflora (Walt. ) Britton. 
Hairy Solomon’s-seal; smaller Solomon’s-seal. 


Native, perennial herb, 8 inches to 3 feet high, found in woods and thickets 
from Canada south to Florida and Michigan. 


Part used.—Rhizome (nonofficial). 
Polygonatum commutatum (Roem. & Schult.) Dietr. Lily-of-the-valley family 
(Convallariaceae). 


Synonyms.—Polygonatum giganteum Dietr.; Salomonia commutata (Roem. & 
Schult.) Britton. 


Giant Solomon’s-seal; great Solomon’s-seal; smooth Solomon’s-seal. 


Native, perennial herb, 1 to 8 feet high, occurring in moist woods and along 
streams from Canada to Georgia, W est to Louisiana and Utah. 


Part used.—Rhizome (nonofficial). 
Polygonatum giganteum Dietr. Same as Polygonatum commutatum. 


Polygonum hydropiper L. Buckwheat family (Polygonaceae). 
Smartweed; water-pepper; biting knotweed; pepper-plant. 
Smooth, annual plant, 8 inches to 2 feet high, naturalized from Europe; common 
in moist waste places almost throughout North America. 
Part used.—Herb (nonofficial ). 


Polygonum punctatum Ill. Buckwheat family (Polygonaceae). 
Dotted smartweed; water-smartweed. 


Native, annual or perennial herb, found in swamps and other wet places through- 
out most of North America. 


Part used.—Herb (nonofficial). 
Polymnia uvedalia L. Aster family (Asteraceae). 
Yellow bear’s-foot; yellow leafcup; uvedalia. 


Large, native, perennial plant, 3 to 6 feet high; in ravines and edges of woods 
from New York to Mic higan, south to Florida and Texas. 


Part used.—Root (nonofficial). 
Polypodium filiz-mas L. Same as Dryopteris filix-mas. 


Polypodium marginale L. Same as Dryopteris marginalis. 


POLYPODIUM VULGARE—POTENTILLA CANADENSIS. 55 


Polypodium vulgare L. Fern family (Polypodiaceae). 
Common polypody; fernroot; rock-brake; female-fern. 


Native fern, 3 to 10 inches in height, with a perennial, creeping rhizome; on 
shady, rocky banks, in woods and mountains almost throughout North 
America. 


Parts used.—Rhizome and tops (nonofificial). 
Polypody, common. See Polypodium vulgare. 


Polytrichum juniperinum Hedw. Haircap-moss family (Polytrichaceae). 
Haircap-moss; robin’s-rye. 
Native moss, 4 to 7 inches in height, growing along margins of dry woods and 
exposed places, mostly on poor, sandy soil. 
Part used.—Whole plant (nonofficial). 


Pond-lily, large yellow. See Nymphaea advena. 

Pond-lily, white. See Castalia odorata. 

Poolroot. See Eupatorium ageratoides, FE. aromaticum, and Sanicula marilandica. 
Poolwort. See Hupatorium ageratoides and E. aromaticum. 

Poplar, silver. See Populus alba. 

Poplar, silverleaf-. See Populus alba. 

Poplar, trembling. See Populus tremuloides. 

Poplar, tulip-. See Liriodendron tulipifera. 

Poplar, white. See Populus alba and P. tremuloides. 

Poplar, yellow. See Liriodendron tulipifera. 


Populus alba L. Willow family (Salicaceae). 
White poplar; silverleaf-poplar; silver poplar; white-bark. 
A large tree, sometimes 120 feet in height, naturalized in the United States; 
occurs along roadsides from New Brunswick to Virginia. 
Part used.—Bark, collected in spring (nonofficial). 


Populus balsamifera candicans A. Gray. Same as Populus candicans. 


Populus candicans Ait. Willow family (Salicaceae). 
Synonym.— Populus balsamifera candicans A. Gray. 
Balm-of-Gilead. 
A large tree, about 80 feet in height, mostly escaped from cultivation, New 
Brunswick to New Jersey, west to Minnesota. 
Parts used.—Leatbuds and bark (nonofficial) . 
Populus tremuloides Michx. Willow family (Salicaceae). 
Quaking aspen; American aspen; white poplar; trembling poplar; quiverleaf. 
A slender, indigenous tree, growing in dry or moist soil from lower Canada south 
to Kentucky and in the Rocky Mountains to Lower California. 
Part used.—Bark, collected in spring (nonofficial). 


oe enue trifoliatus (L.) Britton. Rose family (Rosaceae). 
Synonym.—Gillenia trifoliata Moench. 
Indian-physic; Bowman’s-root; false ipecac; western dropwort. 


Native, perennial herb, 2 to 3 feet high, found in moist, shady places in rich 
woods from New York to Michigan, south to Georgia and Missouri; more 
common in the Atlantic States than in the Western States. 


Part used.—Root (nonoflicial). 
Potato, hog-. See Ipomoea pandurata. 
Potato, wild. See Ipomoea pandurata. 
Potentilla canadensis L. Rose family (Rosaceae). 
Fivefinger; cinquefoil. 
A small, annual or biennial plant, with creeping stems, growing in dry soil from 
Quebec to Georgia, west to Minnesota and the Indian Territory. 
Part used.—Plant (nonofficial ). 


56 WILD MEDICINAL PLANTS OF THE UNITED STATES. 


Prairie-pine. See Lacinaria spicata. 

Prenanthes alba L. Same as Nabalus albus. 

Prenanthes serpentaria Pursh. Same as Nabalus serpentarius. 

Prickly ash, northern. See Xanthoxylum americanum. 

Prickly ash, southern. See Fagara clava-herculis. 

Prideweed. See Erigeron canadensis. 

Prim. See Ligustrum vulgare. 

Primrose, evening-. See Oenothera biennis. 

Primrose, tree-. See Oenothera biennis. 

Primwort. See Ligustrum vulgare. 

Prince’s-pine. See Chimaphila umbellata. 

Prinos verticillatus L. Same as Ilex verticillata. 

Privet. See Ligustrum vulgare. 

Prunella vulgaris L. Mint family (Menthaceae). 
Self-heal; heal-all; brownwort; sicklewort; blue-curls. 
Perennial plant, 2 inches to 2 feet high, naturalized from Europe, and found 


in fields, woods, and waste places throughout nearly the whole of North 
America. 


Part used.—Herb (nonoficial). 

Prunus serotina Ehrh. Plum family ‘Amygdalaceae). 

Synonym.—Prunus virginiana Mill., not of Linnaeus. 

Prunus virginiana; wild cherry; rum-cherry. 

A large, indigenous tree, 50 to 80 feet high, growing in woods or open places from 
Ontario to Florida, west to Texas and Dakota. Most abundant in the South- 
western States. 

Part. used.—Bark, which should be collected in autumn and carefully dried and 
preserved (official). 

Prunus virginiana. See Prunus serotina. 

Prunus virginiana Mill., not L. Same as Prunus serotina. 
Psoralea. See Psoralea pedunculata. 

Psoralea melilotoides Michx. Same as Psoralea pedunculata. 


Psoralea pedunculata ( Mill.) Vail. Pea family (Fabaceae). 
Synonym.—Psoralea melilotoides Michx. 
Psoralea; Samson’s-snakeroot; Congo-root. 
Slender, herbaceous perennial, 1 to 23 feet high, native in dry soil in open woods 
from Ohio and Kentucky southward. 


Parts used.—Root and leaves (nonofficial ). 


Ptelea trifoliata L. Rue family (Rutaceae). 
Wafer-ash; wingseed; hop-tree; shrubby trefoil. 
Native shrub, 6 to.8 feet high; in shady woods from New York to Florida, west 
to Minnesota and Texas; grows more abundantly west of the Alleghenies. 


Parts used. —Bark of root, fruit, and leaves (nonofficial). 
Pterocaulon undulatum ( Walt.) Mohr. Aster family (Asteraceae). 
Synonym.—Gnaphalium undulatum Walt. 
Indian blackroot. 
Native, perennial herb, growing in sandy pine lands from North Carolina to 
Florida and Mississippi. 
Part used.—Root (nonofticial). 
Puccoon, red. See Sanguinaria canadensis. 
Puccoon, yellow. See Hydrastis canadensis. 


Pulsatilla, American. See Pulsatilla hirsutissima. 


PULSATILLA HIRSUTISSIMA——-RATTLESN AKE-HERB. 57 


Pulsatilla hirsutissima (Pursh) Britton. Crowfoot family (Ranunculaceae). 
Synonym.—Anemone patens var. nuttalliana A. Gray. 
American pasqueflower; American pulsatilla. 


Native, perennial herb, 6 to 16 inches high, found in the prairie regions of Illi- 
nois, west to the Rocky Mountains and the Northwest. 
Part used.—F lowering herb (nonofficial ). 


Purging-root. See Euphorbia corollata. 

Purslane, black. See Euphorbia nutans. 

Purslane, milk-. See Euphorbia nutans. 

Pussy-willow. See Salix nigra. 

Putty-root. See Aplectrum spicatum. 

Pycnanthemum montanum Michx. Same as Koellia montana. 
Pycnanthemum pilosum Nutt. Same as Koellia pilosa. 
Pyramid-flower. See Frasera carolinensis. 

Pyrethrum parthenium Smith. Same as Chrysanthemum parthenium. 


Pyrus americana DC. Same as Sorbus americana. 


Quack-grass. See Agropyron repens. 
Queen-Anne’s-lace. See Daucus carota. 
Queen-of-the-meadow. See Lupatorium purpureum. 
Queen’s-delight. See Stillingia sylvatica. 
Queensland asthma-weed. See Huphorbia pilulifera. 
Queen’s-root. See Stillingia sylvatica. 

Quercus. See Quercus alba. 


Quercus alba L. Beech family (Fagaceae). 
Quercus; white oak; stone-oak. 


Large, indigenous forest tree, 50 to 100 feet in height, in woods from Maine to 
Minnesota, south to Florida and Texas. More abundant in the Middle States. 


Part used. —Bark, ‘“‘collected from trunks or branches 10 to 25 years of age, and 
deprived of the periderm”’ (official). 


Quercus rubra L. Beech family (Fagaceae). 
Red oak; champion-oak; Spanish oak. 


Large, wide-spreading, indigenous forest tree, about 70 feet in height, from Nova 
Scotia to Minnesota, south to Florida and Texas. More common in the 
Northern States and in Canada. 


Part used.—Bark (nonofiicial). 
Quinine-flower. See Sabbatia elliottii. 
Quinine-herb. See Sabbatia elliottii. 
Quinine-plant. See Sabbatia elliottii. 
Quiverleaf. See Populus tremuloides. 


Ragged-cup. See Silphium perfoliatum. 
Ragweed. See Ambrosia artemisiaefolia. 
Ragwort, golden. See Senecio aureus. 
Raspberry, black. See Rubus occidentalis. 
Raspberry, ground-. See Hydrastis canadensis. 
Raspberry, wild red. See Rubus strigosus. 
Rattle-root. See Cimicifuga racemosa. 
Rattlesnake-herb. See Actaea alba and A. rubra. 


58 WILD MEDICINAL PLANTS OF THE UNITED STATES. | iat» 


Rattlesnake-master. See Eryngium yuccifolium, Lacinaria scariosa, L. spicata, and 
L. squarrosa. 

Rattlesnake-plantain, downy. See Peramium pubescens. 

Rattlesnake-plantain, lesser. See Peramiwimn repens. 

Rattlesnake-root. See Nabalus albus and N. serpentarius. 

Rattlesnake-violet. See Hrythronium americanum. 

Rattlesnake-weed. See Eryngium yuccifolium, Hieracium venosum, and Peramium 
pubescens. 

Redbud. See Cercis canadensis. 


Redroot. See Ceanothus americanus. 
* 


Rhamuus cathartica L. Buckthorn family (Rhamnaceae). 
Buckthorn; hart’s-thorn; waythorn. 


A shrub 6 to 15 feet high, introduced from Europe; escaped from hedges and 
growing in dry soil in ‘the New England and Middle States. 


Part used.—Berries (nonofficial ). ’ 
Rhamnus purshiana. See Rhamnus purshiana DC. 


Rhamnus purshiana DC. Buckthorn family (Rhamnaceae). 
Rhamnus purshiana; cascara sagrada; chittem-bark; sacred-bark; bearberry-tree. 
Small, indigenous tree, 15 to 20 feet in height, found on the sides and bottoms 

of canyons, Rocky Mountains west to the Pacifie Ocean, and extending north 
into British America. 
Part used.—Bark, collected at least one year before being used (official). 


Rheumatism-root. See Dioscorea villosa and Jeffersonia diphylla, 
Rheumatism-weed. See Chimaphila umbellata. 


Rhododendron maximum L. Heath family (Ericaceae). 

Great laurel; rose-bay; deer-laurel; rose-laurel. 

Tall, native, evergreen shrub or small tree, found in low woods and along streams 
from Canada to Georgia. 

Part used.—Leaves (nonofficial ). 

Rhus aromatica Ait. Sumac family (Anacardiaceae). 

Fragrant sumac; sweet-scented sumac. 

Indigenous shrub, 2 to 6 feet high, growing in woods and rocky situations, 
Canada to Florida, especially along. the mountains, west to Minnesota and 
Arkansas. 

Part used.—Bark of root (nonoflicial). 


Rhus glabra. See Rhus glabra L. 


Rhus glabra L. Sumac family (Anacardiaceae ). 
thus glabra; smooth sumac; scarlet sumac. 


Indigenous, branching shrub, from 4 to 12 feet high; in dry soil, thickets, and 
waste grounds nearly throughout the United States and Canada. 


Parts used. —Fruit (official) ; bark and leaves (nonofficial). 
Rhus radicans L, “ Sumac family (Anacardiaceae). 
Rhus toxicodendron (pharmacopeial name, 1890); poison-ivy; poison-oak; 
poison-vine. 
Native, woody vine, clinging to trees and fence rows; Canada to Florida, west ; 
to Nebraska and Arkansa Very poisonous to the touch. 
Part used.—Fresh leaves (oficial in U. 8. P. 1890). 


Rhus toxicodendron. See Rhus radicans. 


a Rhus radicans L. was formerly believed to be a variety of Rhus toxicodendron L., but the two are 
now regarded as distinet species, and the leaves from both have been used under the pharmacopoeial 
name (U.S. P. 1890) Rhus toxicodendron. 


RHUS TOXICODENDRON—RUBUS OCCIDENTALIS. 59 


Rhus toxicodendron L. Sumac family (Anacardiaceae). 

Poison-ivy; poison-oak. 

Low, erect, and finely pubescent plant, more shrubby than Rhus radicans, and 
found in dry soil in more southern localities from Virginia to Georgia. Very 
poisonous to the touch. 

Part used.—Fresh leaves, collected with those of Rhus radicans. 


Richweed. See Collinsonia canadensis and Eupatorium ageratoides. 
Robinia pseudacacia L. Pea family (Fabaceae). 


Locust-tree; black locust; yellow locust; false acacia. 


A large, indigenous tree, sometimes 80 feet in height, growing in woods from 
Pennsylvania south along the western slope of the Allegheny Mountains to 
Georgia, west to the Indian Terr itory. Most abundant in the Middle and 
Eastern States. 


Part uwsed.—Bark of root (nonoflicial). 
Robin’s-rye. See Polytrichum juniperinumn. 
Rock-brake. See Polypodium vulgare. 

Rock-rose, Canadian. See Helianthemuim canadense. 
Rope-bark. See Dirca palustris. 

Rose, Canadian rock-. See Helianthemum canadense. 
Rose-bay. See Rhododendron maximum. 
Rose-laurel. See Rhododendron maximum. 
Rosemary, marsh-. See Limoniwm carolinianum. 
Rose-pink. See Sabbatia angularis. 
Rose-willow. See Cornus amomum. 
Rosinweed. See Silphiwm laciniatuin., 
Roundwood. See Sorbus americana. : 


Rubus. See Rubus cuneifolius, R. nigrobaccus, R. procumbens, R. trivialis, and R. 
villosus. 


Rubus canadensis T. & G., not L. Same as Rubus procumbens. 


Rubus cuneifolius Pursh. Rose family (Rosaceae). 
Rubus; sand-blackberry; knee-high blackberry. 


Shrubby plant, 1 to 3 feet high; in sandy soil from Connecticut to Florida, 
west to Missouri and Louisiana. 


Part used.—Bark of rhizome (official). 
Rubus idaeus var. americanus Torr. Same as Rubus occidentalis. 


Rubus nigrobaccus Bailey. Rose family (Rosaceae). 
Synonym.—Rubus villosus A. Gray, not Ait. 
Rubus; high-bush blackberry. 


Slender shrub, 3 to 7 feet high, growing in dry fields and along roadsides, New 
England States to Flor ida, and west to Arkansas. 


Part used.—Bark of rhizome (official). 


Rubus occidentalis L. Rose family (Rosaceae). 
Synonym.—Rubus idaeus var. americanus Torr. 
Black raspberry; thimbleberry; blackcap. 


A straggling shrub, growing along the borders of woods and in rocky thickets 
from Canada south to Georgia and Missouri. 


Parts used.—Fruit and leaves (nonofficial) . 


60 WILD MEDICINAL PLANTS OF THE UNITED STATES. 
Rose family (Rosaceae). 


Rubus procumbens Muhl. 
Synonym.—Rubus canadensis T. & G., not L. 


Rubus; low running blackberry; dewberry. 
Shrubby, trailing plant, found in dry soil from Newfoundland to Lake Superior, 


south to Virginia and the Indian Territory. 
Part used.—Bark of root (official in U. 8. P. 1890). 


Rubus strigosus Michx. Rose family (Rosaceae). 
Wild red raspberry. 
Shrubby plant, found in dry or rocky situations from Canada to North Carolina 


and New Mexico. 
Parts used.—Fruit and leaves (nonoflicial) . 
Rose family (Rosaceae). 


Rubus trivialis Michx. 
Rubus; southern dewberry; low-bush blackberry. 
Shrubby, procumbent plant, found in sandy soils, Virginia to Florida, west to 


Missouri and Texas. 
Part used.—Bark of root (official in U. 8. P. 1890). 
Rubus villosus A. Gray, not Ait. Same as Rubus nigrobaccus. 
Rose family (Rosaceae). 


Rubus villosus Ait. 


Rubus; one-flowered dewberry. 
Trailing plant, with slender branches, growing in sandy or dry soil near the 


coast from Maine to South Carolina. 


Part used.—Bark of rhizome (official) 
Aster family (Asteraceae). 


Rudbeckia laciniata L. 


Thimbleweed; tall coneflower. : 
Much-branched, native perennial, 3 to 12 feet high; in moist thickets, Canada 


and Montana, south to Florida and New Mexico. 
Part used.—Herb (nonofiicial ). 


Rum-cherry. See Prunus serotina. 


Rumex. See Rumex crispus. 
Buckwheat family (Polygonaceae). 


Rumex acetosella L. 
Sheep-sorrel; field-sorrel; sour-grass; common sorrel. 
Annual or perennial herb, abundant in dry fields, pastures, and waste ground 
throughout the United States. 
Leaves (nonofficial). 


Part used. 


Rumex crispus L. 
Rumex; yellow dock; curled dock; narrow dock; sour dock. 
A weed introduced from Europe, and common in cultivated and waste ground 
throughout the United States. Perennial plant, 2 to 4 feet high. 
Part used.—Root of this and some other species of Rumex (official in U, 8. P. 


Buckwheat family (Polygonaceae). 


1890). 
Buckwheat family (Polygonaceae). 


Rumex obtusifolius L. 
Bitter dock; blunt-leaved dock; broad-leaved dock. 
A perennial weed, 2 to 4 feet high, naturalized from Europe, and found in waste 


places from New England to Florida, west to Texas and Oregon. 


Part used.—Root, collected with that of Rumex crispus. 


Sabal. See Serenoa serrulata. 
Gentian family (Gentianaceae). 


Sabbatia angularis (L.) Pursh. 
American centaury; rose-pink; bitterbloom; bitter clover. 
2 feet high, growing in damp, rich soil, in meadows 


Native, biennial plant, 1 to 2 
and among high grass, from New York to Michigan, south to Florida and 


the Indian Territory. 
Part used.—Herb (nonofficial). 


SABBATIA ELLIOTTII—SANICLE, WHITE. 6) 


Sabbatia elliottii Steud. Gentian family (Gentianaceae). 
Synonym.—Sabbatia paniculata Ell. 
Quinine-flower; quinine-plant; quinine-herb; Elliott’s-sabbatia. 
An erect, native herb, about one foot in height, growing in pine barrens from 
North Carolina to Florida. 


Part used.—Herb (nonofficial). 

Sabbatia, Elliott’s-. See Sabhatia elliottii. 

Sabbatia paniculata Ell. Same as Sabbatia elliottii. 

Sabina. See Juniperus sabina. 

Sacred-bark. See Rhamnus purshiana. 

Sage, Indian. See Lupatorium perfoliatum. 

Saint-Benedict’s thistle. See Cnicus benedictus. 

Saint-John’s wort, common. See Hypericum perforatum. 

Salix alba L. Willow family (Salicaceae). 
White willow; European willow. 
A large tree, sometimes 90 feet in height, introduced from Europe; occurs in 


moist soil along streams from Pennsylvania northward to New Brunswick 
and Ontario, sparingly escaped from cultivation. 


Part used.—Bark (nonofificial). 


Salix nigra Marsh. Willow family (Salicaceae). 
Black willow; pussy-willow; swamp-willow. 
Tall, indigenous tree, growing on banks of rivers from Canada to Florida and 
California. 


Parts used.—Bark, and fresh aments gathered early in May (nonofficial). 
Salomonia biflora (Walt.) Britton. Same as Polygonatum biflorum. 
Salomonia commutata (Roem. & Schult.) Dietr. Same as Polygonatum commutatui. 
Salt-rheum weed. See Chelone glabra. 
Sambucus. See Sambucus canadensis. 


Sambucus canadensis L. Honeysuckle family (Caprifoliaceae). 
Sambucus; elder; American elder; sweet elder. 


s 5 ~ : P : A < / 
Indigenous shrub, 6 to 10 feet high, growing in low, damp ground from Canada 
to Florida and Arizona. 


Parts used.—Flowers (official in U. 8. P. 1890); bark and berries (nonofficial) . 
Sampson-root. See Brauneria angustifolia. 
Sampson’s-snakeroot. See Gentiana villosa. 
Samson’s-snakeroot. See Psoralea pedunculata. 
Sand-blackberry. See Rubus cuneifolius. 
Sandbrier. See Solanum carolinense. 
Sanguinaria. See Sanguinaria canadensis. 


Sanguinaria canadensis L. Poppy family (Papaveraceae). 
Sanguinaria; bloodroot; red puccoon; Indian-paint; tetterwort. 


Native, perennial herb, about 6 inches high, found in rich, open woods from 
Nova Scotia to Nebraska, south to Florida and Arkansas. 


Part used.—Rhizome, ‘‘ collected after the death of the foliage’’ (official). 
Sanicle, American. See Heuchera americana and Sanicula marilandica, 
Sanicle, black. See Sanicula marilandica. 

Sanicle, Indian. See Eupatoriwm ageratoides. 
Sanicle, white. See Eupatorium ageratoides. 


62 WILD MEDICINAL PLANTS OF THE UNITED STATES. 


Sanicula marilandica L. Parsley family (Apiaceae). 
Black sanicle; black snakeroot; American sanicle; poolroot. 
Native, perennial herb, 1 to 3 feet high; in rich woods, Canada to Georgia. 
Part used.—Root (nonofficial). 

Saponaria officinalis L. Pink family (Silenaceae). 
Soapwort; soaproot; bouncing-Bet; fuller’s-herb. 


Stout, perennial herb, 1 to 2 feet high, naturalized from Europe and found along 
roadsides and waste places; common almost everywhere. 


Parts used.—Root and herb (nonofficial). 

Sarothamnus scoparius Wimm. Same as Cytisus scoparius. 

Sarracenia flava L. Pitcher-plant family (Sarraceniaceae). 
Trumpetleaf; trumpets; Eve’s-cup; watercup; yellow-flowered watercup. 


Curious, indigenous perennial, about 1 to 3 feet high, found in low, wet pine 
barrens in the southeastern United States. 


Parts used.—Root and sometimes the leaves (nonoflcial). 
Sarracenia purpurea L. Pitcher-plant family (Sarraceniaceae). 
Pitcher-plant; flytrap; sidesaddle-flower; watercup; smallpox-plant. 


Indigenous perennial, 1 to 2 feet high, growing in wet, boggy places and marshes, 
from Canada to Minnesota and Florida. 


Parts used.—Root and sometimes the leaves (nonoflicial). 

Sarsaparilla, American. See Aralia nudicaulis. 

Sarsaparilla, bristly. See Aralia hispida. 

Sarsaparilla, false. See Aralia nudicaulis. 

Sarsaparilla, Texas. See Menispermum canadense. 

Sarsaparilla, Virginian. See Aralia nudicaulis. 

Sarsaparilla, wild. See Aralia nudicaulis. 

Sassafras. See Sassafras varufolium. 

Sassafras officinale Nees & Eberm. Same as Sassafras variifolium. 

Sassafras sassafras (L.) Karst. Same as Sassafras variifoliun. 

Sassafras, swamp-. See Magnolia virginiana. 

Sassafras variifolium (Salisb.) O. Kuntze. ¢ Laurel family (Lauraceae). 
Synonyms.—NSassafras officinale Nees & Eberm.; Sassafras sassafras (L.) Karst. 4 
Sassafras; ague-tree. 


Native tree, sometimes reaching a height of 125 feet; in rich woods, Massachu- 
setts to Ontario and Michigan, south to Florida and Texas. 


Parts used.—Bark of root, collected in early spring or autumn and deprived of 
the periderm (official); pith (official) ; and the oil of sassafras distilled from 
the root, especially the root bark (official). 


Satureia hortensis L. Mint family (Menthaceae). 
Summer-savory. 


Hairy, aromatic, annual herb, adventive from Europe and occurring in waste 
places from Canada to Pennsylvania and Nevada. 


Part used.—Herb (nonofficial). 
Savin. See Juniperus sabina. 
Savin, red. See Juniperus virginiana. 
Savory, summer-. See Satureia hortensis. 
Saw-palmetto. See Serenoa serrulata. 


Saxifrage, burnet-. See Pimpinella saxifraga. 


«Although the combination Sassafras a (L.) Karst. should be accepted by strict right of 
priority, the usage of the Pharmacopceia is followed. 


SCABIOUS, SWEET—SERVICE-TREE, AMERICAN, 63 


Scabious, sweet. See Hrigeron philadelphicus. 

Scabish, meadow-. See Aster puniceus. 

Scabwort. See Inula helenium. 

Scarletberry. See Solanum dulcamara. 

Scoke. See Phytolacca decandra. 

Scoparius. See Cytisus scoparius. 

“Scouring-rush, common. See Equisetum hyemale. 

Scrofula-plant. See Scrophularia marilandica. 

Scrofula-weed. See Peramium pubescens. 

Scrophularia marilandica L. Figwort family (Scrophulariaceae). 
Synonym.—Scrophularia nodosa var. marilandica A. Gray. 


Maryland figwort; scrofula-plant; carpenter’s-square; heal-all; bee-plant; pile- 
wort. 


Smooth, native perennial, 3 to 5 feet high; moist, shady ground in woods and 
thickets, New York to North Carolina and Kansas. 
Parts used.—Herb and root (nonofficial ). 
Scrophularia nodosa var. marilandica A. Gray. Same as Scrophularia marilandica. 
Scutellaria. See Scutellaria lateriflora. 
Scutellaria hyssopifolia L. Same as Scutellaria integrifolia. 
Scutellaria integrifolia L. Mint family (Menthaceae). 
Synonym.—Scutellaria hyssopifolia L. ‘ 
Larger skullcap; hyssop-skulleap. 
Native, perennial herb, 6 inches to 2 feet high, found in fields and woods from 
Connecticut south to Florida and Texas. 


Part used.—Herb (nonofficial). 
Scutellaria laterifiora L. Mint family (Menthaceae). 
Scutellaria; skullcap; madweed; hoodwort. 


Smooth, branching perennial, 1 to 2 feet high, native in damp places along 
banks of streams from Canada south to Florida, New Mexico, and Washington. 


Part used.—Plant (official). 

Sea-lavender. See Limoniwm carolinianum. 

Self-heal. See Prunella vulgaris. 

Senecio aureus L. Aster family (Asteraceae). 
Liferoot; swamp squaw-weed; golden ragwort; cocash-weed; coughweed. 


Indigenous, perennial herb, 1 to 23 feet high, growing in swamps and wet mead- 
ows, Newfoundland to Ontario, south to Florida, Missouri, and Texas. 


Parts used.—Root and herb (nonofficial). 
Senega. See Polygala senega. 
Senna, American. See Cassia marilandica. 
Senna, wild. See Cassia marilandica. 


Serenoa serrulata (Roem. & Schult.) Hook. f. Palm family (Phoenicaceae). 
Sabal; saw-palmetto. 


A palm, 3 to 7 feet in height, found in sandy soil from North Carolina and 
Arkansas to Florida and Texas. 


Part used.—Partially dried ripe fruit (official). 
Serpentaria. See Aristolochia reticulata and A. serpentaria 
Serpentaria, Texas. See Aristolochia reticulata. 
Serpentaria, Virginia. See Aristolochia serpentaria. 
Service-tree, American. See Sorbus americana. 


64 WILD MEDICINAL PLANTS OF THE UNITED STATES. 


Seven-barks. See Hydrangea arborescens. 

Shagbark. See /Ticoria ovata. 

Shamrock. See Ovalis acetosella. 

Shamrock, water-. See Menyanthes trifoliata. 

Shave-grass. See Hquisetum hyemale. 

Sheepberry. See Viburnum lentago. 

Sheep-laurel. See Kalmia angustifolia and Kk. latifolia, 

Sheep-sorrel. See Rumex acetosella. 

Shellbark-hickory. See Hicoria ovata. 

Shellflower. See Chelone glabra. 

Shepherd’s-purse. See Bursa bursa-pastoris. 

Shepherd’s-weatherglass. See Anagallis arvensis. 

Shield-fern, marginal-fruited. See Dryopteris marginalis. 

Shrub, sweet-scented. See Butneria florida. 

Shrub yellowroot. See Xanthorrhiza apiifolia. 

Sicklewort. See Prunella vulgaris. 

Sidesaddle-flower. See Sarracenia purpurea. 

Silkweed. See Asclepias syriaca. 

Silkweed, rose-colored. See Asclepias incarnata. 

Silkweed, swamp-. See Asclepias incarnata. 

Silphium laciniatum L. Aster family (Asteraceae). 
Rosinweed; compass-plant; pilotweed; polar-plant. 


Coarse, native perennial, 3 to 12 feet high, growing on prairies from Ohio to 
Alabama, west to Texas and South Dakota. 


Part used.—Herb (nonofficial ). 


Silphium perfoliatum L. Aster family (Asteraceae). 
Cup-plant; Indian-cup; ragged-cup. 
Stout, perennial herb, 4 to 8 feet high, native in moist soil and low ground from 
Ontario and the eastern United States west to Louisiana and Nebraska. 


Part used.—Root (nonofficial). 
Silverleaf. See Impatiens biflora, Spiraea tomentosa, and Stillingia sylvatica, 
Silverleaf-poplar. See Populus alba. 
Simpler’s-joy. See Verbena hastata. 
Sinapis alba. See Sinapis alba L. 


Sinapis alba L. Mustard family ( Brassicaceae). 
Sinapis alba; white mustard; yellow mustard. 


Annual herb, about 2 feet in height, naturalized from eer and found in 
fields and waste places, but not so widely distributed as the black mustard. 


Part used. —Seed (official ). 
Sinapis nigra. See Brassica nigra. 
Sinapis nigra L. Same as Brassica nigra. 
Skullcap. See Seutellaria lateriflora. 
Skullcap, hyssop-. See Seutellaria integrifolia. 
Skullcap, larger. See Seutellaria integrifolia. 
Skunk-cabbage. See Spathyema foetida. 
Skunkweed. See Spathyema foetida. 
Sloe. See Viburnum prunifolium. 


Smallpox-plant. See Sarracenia purpurea. 


SMARTWEED—SOLANUM CAROLINENSE. 65 


Smartweed. See Polygonum hydropiper. 

Smartweed, dotted. See Polygonum punctatum. 
Smartweed, water-. See Polygonum punctatum. 
Smilacina racemosa Desf. Same as Vagnera racemosa. 


Smilax herbacea L. Smilax family (Smilacaceae). 
Carrion-flower; American Jacob’s-ladder. 


Native, herbaceous perennial, occurring in woods and thickets in Canada and 
the eastern United States. 


Part used.—Herb (nonofficial ). 


Smilax pseudo-china L. Smilax family (Smilacaceae). 
Bamboo-brier; long-stalked greenbrier; American China-root; false China-root; 
bullbrier. 


Perennial vine, native, growing in dry or sandy thickets, Maryland to Florida, 
west to Texas and Nebraska. 


Part used.—Rhizome (nonofficial ). 
Snake-gentian. See Nabalus serpentarius. 
Snakehead. See Chelone glabra. 
Snakeleaf, yellow. See Erythronium americanum. 
Snake-lily. See Iris versicolor. 
Snakemilk. See Huphorbia corollata. 
Snakeroot, black. See Cimicifuga racemosa and Sanicula marilandica. 
Snakeroot, button-. See Hryngium yuccifolium. 
Snakeroot, Canada. See Asarum canadense. 
Snakeroot, corn-. See Hryngium yuccifolium and Lacinaria spicata. 
Snakeroot, dense button-. See Lacinaria spicata. 
Snakeroot, large button-. See Lacimaria scariosa. 
Snakeroot, Red River. See Aristolochia reticulata. 
Snakeroot, Sampson’s-. See Gentiana villosa. 
Snakeroot, Samson’s-. See Psoralea pedunculata. 
Snakeroot, Seneca. See Polygala senega. 
Snakeroot, smaller white. See EHupatorium aromaticum. 
Snakeroot, Texas. See Aristolochia reticulata. 
Snakeroot, Virginia. See Aristolochia serpentaria. 
Snakeroot, white. See Hupatoriwm ageratoides. 
Snake-violet. See Viola pedata. 
Snakeweed. See Euphorbia pilulifera. 
Snapweed. See Impatiens aurea and J. biflora. 
Sneezeweed. See Helenium autumnatle. 
Sneezewort. See Helenium autumnale. 
Snowdrop, yellow. See Hrythronium americanum. : 
Soaproot. See Saponaria officinalis. 
Soapwort. See Saponaria officinalis. 
Soapwort-gentian. See Gentiana saponaria. 


Solanum carolinense L. Potato family (Solanaceae). 
Horse-nettle; bull-nettle; sandbrier. 


Rough-hairy, native, perennial herb, common in dry fields and on sandy or 
gravelly banks from the eastern United States west to Texas and Nebraska. 


Parts used.—Root, leaves, and berries (nonofficial ). 


66 WILD MEDICINAL PLANTS OF THE UNITED STATES. 


Solanum dulcamara L. Potato family (Solanaceae). 
Duleamara; bittersweet; woody nightshade; violet-bloom; scarletberry. 


Climbing, shrubby perennial, naturalized from Europe; found in low, damp 
grounds and moist banks, New Brunswick to Minnesota, south to New Jersey 
and Kansas. 


Part used.—Young branches (official in U. 8. P. 1890). 


Solidago odora Ait. Aster family (Asteraceae). 
Sweet goldenrod; fragrant-leaved goldenrod; anise-scented goldenrod. 
Slender, perennial herb, 2 to 3 feet high, native; in dry soil from Maine to Texas. 
Parts used.—Leaves and tops (nonoflficial). 


Solomon’s-seal, false. See Vagnera racemosa. 
Solomon’s-seal, giant. See Polygonatum commutatum. 
Solomon’s-seal, great. See Polygonatum commutatum. 
Solomon’s-seal, hairy. See Polygonatum biflorum. 
Solomon’s-seal, small. See Vagnera racemosa. 
Solomon’s-seal, smaller. See Polygonatum biflorum. 
Solomon’s-seal, smooth. See Polygonatum commutatum. 


Sorbus americana Marsh. Apple family (Malaceae). 
Synonym.—Pyrus americana DC. 
American mountain-ash; roundwood; dogberry; mountain-sumac; American 
service-tree. 
Indigenous tree or tall shrub, growing in low woods or moist ground from New- 
foundland south along the mountains to North C ‘arolina, and to Michigan. 


Parts used.—Bark and berries (nonoflicial). 
Sorrel, common. See Rumex acetosella. 
Sorrel, field-. See Rumex acetosella. 
Sorrel, sheep-. See Rumesx acetosella. 
Sorrel, white wood-. See Oxalis acetosella. 
Sorrel-tree. See Oxydendrum arboreum. 
Sour-grass. See Rumex acetosella. 
Sourwood. See Oxydendrum arboreum. 
Southernwood. See Artemisia abrotanum. 


Spathyema foetida (L.) Raf. Arum family (Araceae). 
Synonyms.— Dracontium foetidum L.; Symplocarpus foetidus Nutt. 
Skunk-cabbage; skunkweed; polecat-weed; swamp-cabbage. 

Indigenous, perennial herb, about 1 to 2 feet high, found in swamps and wet 
soil from Canada south to Florida, Iowa, and Minnesota. Appears very early 
in spring. 

Parts used.—Rhizome and roots (nonofficial). 

Spatter-dock. See Nymphaea advena. 

Spearmint. See Mentha spicata. 

Speedwell, common. See Veronica officinalis. 

Speedwell, tall. See Veronica virginica. 

Spicebush. See Benzoin benzoin. 

Spicewood. See Benzoin benzoin. 


Spigelia. See Spigelia marilandica, 


SPIGELIA MARILANDICA—STARWORT. 67 


Spigelia marilandica L. Logania family (Loganiaceae). 
Spigelia; pinkroot; Maryland pinkroot; Indian pinkroot; worm-grass. 


Erect, native, perennial herb, 6 inches to 13 feet high, found in rich woods, New 
versey to Florida, west to Texas and Wisconsin. Occurs principally in the 
Southern States. 


Parts used.—Rhizome and roots (official). 
Spignet. See Aralia racemosa. 
Spikenard. See Aralia racemosa. 
Spikenard, American. See Aralia racemosa. 
Spikenard, false. See Vagnera racemosa. 
Spikenard, small. See Aralia nudicaulis. 
Spikenard, wild. See Vagnera racemosa. 
Spindle-tree. See Huonymus atropurpureus. 
Spiraea. See Spiraea tomentosa. 


Spiraea tomentosa L. Rose family (Rosaceae). 
Spiraea; hardhack; steeplebush; pink meadowsweet; silverleaf. 


Native shrub, occurring in low grounds and 1oist meadows from Nova Scotia 
south to Georgia, west to Kansas and Manitoba. 


Parts used.—Leaves and root (nonofficial). 
Spleenwortbush. See Comptonia peregrina. 
Spruce, black. See Picea mariana. 

Spruce, hemlock-. See Tsuga canadensis. 
Spruce, weeping. See Tsuga canadensis. 
Spruce-gum tree. See Picea mariana. 
Spurge, flowering. See Euphorbia corollata, 
Spurge, ipecac-. See Huphorbia ipecacuanhae. 
Spurge, large spotted. See Euphorbia nutans. 
Spurge, pill-bearing. See Euphorbia pilulifera. 
Spurge-laurel. See Daphne mezereum. 
Spurge-olive. See Daphne mezereuin. 
Squawberry. See Mitchella repens. 
Squawbush. See Viburnum opulus. 
Squawflower. See Trillium erectwm. 
Squawmint. See Hedeoma pulegioides. 
Squawroot. See Caulophyllum thalictroides and Cimicifuga racemosa. 
Squaw-vine. See Mitchella repens. 
Squaw-weed. See Hupatorium ageratoides. 
Squaw-weed, swamp. See Senecio aureus. 
Squirrel-corn. See Bikukulla canadensis. 
Squirrel-ear. See Peramium repens. 

Staff-tree. See Celastrus scandens. 

Stagbush. See Viburnum prunifolium. 
Staggerweed. See Bikukulla canadensis. 
Stag’s-horn. See Lycopodium clavatum. 
Stammerwort. See Ambrosia artemisiaefolia. 
Star-grass. See Aletris farinosa. 


Starwort. See Chamaelirium luteum. 


68 WILD MEDICINAL PLANTS OF THE UNITED STATES. 


Starwort, drooping. See Chamaelirium luteum. 

Statice caroliniana Walt. Same as Limonium carolinianum. 

Steeplebush. See Spiraea tomentosa. 

Stellaria media Cyr. Same as Alsine media. 

Stillingia. See Stillingia sylvatica. 

Stillingia sylvatica L. Spurge family (Euphorbiaceae). 
Stillingia; queen’s-root; queen’s-delight; silverleaf. 


Native, herbaceous perennial, 1 to 3 feet in height, occurring in dry, sandy soil, 
and pine barrens from Maryland to Florida, west to Kansas and Texas. 


Part used.—Root (official) . 
Stonecrop, ditch-. See Penthorum sedoides. 
Stonecrop, Virginia. See Penthorum sedoides. 
Stonemint. See Cunila origanoides. 
Stone-oak. See Quercus alba. 
Stoneroot. See Collinsonia canadensis. 
Stramonium. See Datura stramonium. 
Strawberry, scarlet. See Fragaria virginiana. 
Strawberry, Virginia. See Fragaria virginiana. 
Strawberry-shrub, hairy. See Butneria florida. 


Stylosanthes biflora (L.) B.S. P. Pea family (Fabaceae). 
Synonym.—Stylosanthes elatior Sw. 
Pencil-flower; afterbirth-weed. 


Wiry, perennial herb, 6 inches to 2 feet in height, native; occurring in dry soil 
from New York to Florida, west to the Indian Territory. 


Part used. —Herb (nonofficial ). 
Stylosanthes elatior Sw. Same as Stylosanthes biflora. 
Succory. See Cichorium intybus. 

Sumac, fragrant. See Rhus aromatica. 
Sumac, mountain-. See Sorbus americana. 
Sumac, scarlet. See Rhus glabra. 

Sumac, smooth. See Rhus glabra. 

Sumac, sweet-scented. See Rhus aromatica. 
Summer-savory. See Satureia hortensis. 
Sundew, round-leaved. See Drosera rotundifolia. 
Sunflower, swamp-. See Helenium culumnale. 
Swamp squaw-weed. See Senecio aureus. 
Swamp willow-herb. See Epilobium palustre. 
Swamp-cabbage. See Spathyema foetida. 
Swamp-dogwood. See Cornus amomum. 
Swamp-hellebore. See Veratrum viride. 
Swamp-laurel. See Magnolia virginiana. 
Swamp-maple. See Acer rubrum. 
Swamp-milkweed. See Asclepias incarnata. 
Swamp-sassatras. See Magnolia virginiana. 
Swamp-silkweed. See Asclepias incarnata. 
Swamp-sunflower. See Helenium autumnale. 


Swamp-willow. See Salix nigra. 


SWEATW EED—THIMBLEBERRY. 69 


Sweatweed. See Althaea officinalis. 
Sweet-cicely. See Washingtonia longistylis. 
Sweet-flag. See Acorus calamus. 
Sweet-gum. See Liquidambar styraciflua. 
Sweetroot. See Polemonium reptans. 


Symphytum officinale L. Borage family (Boraginaceae). 
Comfrey; healing-herb; blackwort; bruisewort. 


Erect, perennial herb, 2 to 3 feet high, naturalized from Europe; found in waste 
places, Newfoundland to Minnesota, south to Maryland. 


Part used.—Root (nonofficial ). 
Symplocarpus foetidus Nutt. Same as Spathyema foetida. 


Tag-alder. See Alnus rugosa. 
Tamarack. See Larix laricina. 
Tanacetum. See Tanacetum vulgare. 


Tanacetum vulgare L. Aster family (Asteraceae). 
Tanacetum; tansy; double tansy; bitter-buttons; parsley-fern. 


Strong-scented, perennial herb, 13 to 3 feet high, introduced from Europe; 
escaped from cultivation and found along roadsides from Nova Scotia to Min- 
nesota, south to North Carolina and Missouri. 


Parts used.—Leaves and flowering tops (official in U. S. P. 1890). 
Tanbark-tree. See Tsuga canadensis. 
Tansy. See Tanacetum vulgare. 
Tansy, double. See Tanacetum vulgare. 
Taraxacum. See Taraxacum officinale. 


Taraxacum officinale Weber.¢ Chicory family (Cichoriaceae). 
Synonym.— Taraxacum taraxacum (L.) Karst.¢ 
Taraxacum; dandelion; blowball; cankerwort. 


Low, perennial weed, 5 to 10 inches high, naturalized from Europe; very abun- 
dant in lawns, meadows, and waste places throughout the United States, with 
the exception of the South. 


Part used.—Root, collected in autumn (official ). 
Taraxacum taraxacum (L.) Karst. Same as Taraxacum officinale. 
Tea, continental. See Ledwm groenlandicum. 

Tea, James-. See Ledum groenlandicum. 

Tea, Jersey. See Ceanothus americanus. 

Tea, Jerusalem. See Chenopodium ambrosioides. 
Tea, Labrador. See Ledum groenlandicum. 

Tea, Mexican. See Chenopodium ambrosioides. 
Tea, mountain-. See Gaultheria procumbens. 
Tea, New Jersey. See Ceanothus americanus. 
Tea, Oswego. See Monarda didyma. 

Tea, Spanish. See Chenopodium ambrosioides. 
Teaberry. See Gaultheria procumbens. 
Tephrosia virginiana Pers. Same as Cracca virginiana. 


Tetterwort. See Chelidonium majus and Sanguinaria canadensis. 


70 WILD MEDICINAL PLANTS OF THE UNITED STATES. 


Thimbles. See Digitalis purpurea. 
Thimbleweed. See Rudbeckia laciniata. 4 
Thistle, bitter. See Cnicus benedictus. 

Thistle, blessed. See Cnicus benedictus. 
Thistle, Canada. See Carduus arvensis. 
Thistle, creeping. See Carduus arvensis. 
Thistle, cursed. See Carduus arvensis. 
Thistle, holy. See Cnicus henedictus. 

Thistle, St. Benedict’s-. See Cnicus henedictus. 
Thistle, spotted. See Cnicus benedictus. 
Thorn-apple. See Datura stramonium. 
Thoroughwort. See Hupatorium perfoliatum. 
Thousandleaf. See Achillea millefolium. 
Throwwort. See Leonurus cardiaca. 


Thuja occidentalis L. Pine family (Pinaceae). 
Arbor-vitae; white cedar; yellow cedar. 
Indigenous, evergreen tree, 20 to 50 feet in height; in wet soil and along banks 
of streams, Canada to North Carolina, Illinois, and Minnesota. Especially 
abundant in Canada and the Northern States. 


Parts used.—Branchlets and leaves (nonofficial). 
Tiarella cordifolia L. Saxifrage family (Saxifragaceae). 
Coolwort; false miterwort; foamflower; gemfruit. 


Slender, indigenous perennial, 6 to 12 inches high, found in rich, moist woods, 
Nova Scotia to Minnesota, south, especially along the mountains, to Georgia 
and Indiana. 


Part used.—Herb (nonofiicial). 
Tickweed. See Hedeoma pulegioides. 


Tilia americana L. Linden family (Tiliaceae). 
Synonym.— Tilia glabra Vent. 
Basswood; American linden; whitewood. 


Large, indigenous forest tree, 60 to 125 feet in height; in rich woods, especially 
along the mountains, from Canada to Georgia, west to Texas and Nebraska. 


Part used.—Inflorescence of this and of other species of Tilia (nonofficial). 
Tilia glabra Vent. Same as Tilia americana. 
Tinker’s-weed. See Triostewm perfoliatum. 
Tobacco, Indian. See Lobelia inflata. 
Toothache-tree. See Fagara clava-herculis and Xanthoxylum americanum. 
Touch-me-not, pale. See Impatiens aurea. 
Touch-me-not, spotted. See Impatiens biflora. 
Toywort. See Bursa bursa-pastoris. 
Tree-primrose. See Oenothera biennis. ; 
Trefoil, marsh-. See Menyanthes trifoliata, 
Trefoil, shrubby. See Ptelea trifoliata. 
Trefoil, sour. See Oxalis acetosella. 


Trifolium pratense L. Pea family (Fabaceae). 
Red clover; meadow-clover; purple clover. 


Perennial herb, 6 inches to 2 feet high; common in fields and meadows through- 
out the eastern United States; naturalized from Europe, and widely cultivated, 
Part used.—Blossoms (nonofficial ), 


TRILISA ODORATISSIMA—TWINLEAF. 71 


Trilisa odoratissima ( Walt.) Cass. Aster family (Asteraceae). 
Synonym.—Liatris odoratissima Michx. 
Vanilla-plant; deer’s-tongue; vanilla-leaf; Carolina vanilla. 


Rather stout, native, perennial herb, 2 to 3 feet high, with fragrant leaves; in 
pine barrens from Virginia south to Florida and Louisiana. 


Part used.—Leaves (nonoffcial ). 


Trillium erectum L. Lily-of-the-valley family (Convallariaceae). 
Wake-robin; ill-scented bethroot; birthroot; squawflower. 
Stout, native perennial, 8 to 16 inches high, growing in rich soil in damp, shady 
woods fronr Canada south to Tennessee and Missouri. 


Part used.—Rhizome of this and of several other species of Trillium (nonofficial ). 


Triosteum perfoliatum L. Honeysuckle family (Caprifoliaceae). 
Feverroot; horse-gentian; tinker’s-weed; white gentian; wild ipecac. 
Indigenous, perennial herb, 2 to 4 feet high; in rich soil in shady locations, Que- 

bee to Minnesota, south to Alabama and Kansas. 


Part used.—Root (nonofiicial) . 
Triticum. See Agropyron repens. 
Triticum repens Beauv. Same as Agropyron repens. 
Trumpetleaf. See Sarracenia flava. 
Trumpet-milkweed. See Lactuca canadensis. 
Trumpets. See Sarracenia flava. 


Tsuga canadensis te Carr. Pine family (Pinaceae). 
Synonym.—Abies canadensis Michx. 
Hemlock; hemlock-spruce; weeping spruce; tanbark-tree. 
Indigenous tree, about 75 feet in height, in forests from Canada south to Alabama 
and Wisconsin. 


Paris used.—Bark and prepared resinous exudate (nonofficial). 
_Tulip-poplar. See Liriodendron tulipifera. 
Tulip-tree. See Liriodendron tulipifera. 
Tupelo gum. See Nyssa aquatica. 
Tupelo, large. See Nyssa aquatica. 
Tupelo, sour. See Nyssa ogeche. 
Turkey-corn. See Bikukulla canadensis. 
Turkey-pea. See Bikukulla canadensis. 


Turnera aphrodisiaca Ward. Same as Turnera microphylla. 


Turnera microphylla Desy. Turnera family (Turneraceae). 
Synonym.— Turnera aphrodisiaca Ward. 
Damiana. 


A small, shrubby plant, native of Lower California, Texas, and northern Mexico, 
growing in dry soil. 
Part used.—Leaves (nonofiicial) . 
Turnip, Indian. See Arisaema triphyllum. 
Turnip, wild. See Arisaema triphyllum. 
Turtle-head. See Chelone glabra. 


Tussilago farfara L. Aster family (Asteraceae). 
Colt’s-foot; coughwort; horsefoot; gingerroot. 


Perennial herb, 3 to 18 inches high, naturalized from Europe; in moist places 
aoe roadsides and brooks, northeastern United States and Minnesota to 
Janada. 


Parts used.—Leaves and root (nonofticial). 
Twinleaf. See Jejfersonia diphylla. 


(2 WILD MEDICINAL PLANTS OF THE UNITED STATES. 


Typha latifolia L. Cattail family (Typhaceae). 
Broad-leaved cattail; cattail-flag; bulrush. 
Native marsh plant, perennial, + to 8 feet high; found in marshes, ditches, 
muddy pools, and other wet places throughout North America, except 
extreme northern part. 


Part used.—Root (nonoflicial ). 


Ulmus. See Ulinus fulva. 


Ulmus fulva Michx. Elm family (Ulmaceae). 

Synonym.—Ulmus pubescens Walt. 

Ulmus; elm; slippery elm; red elm; moose-elm; Indian elm. 

Indigenous tree, 50 to 60 feet high, growing on hills, along streams and in woods 
from Quebec to North Dakota, south to Florida and Texas. More common in 
the western part of its range. 

Part used.—Bark deprived of its periderm (official). 

Ulinus pubescens Walt. Same as Ulmus fulva. 

Umbrella-tree. See Magnolia tripetala. 

Unicorn-root, false. See Aletris farinosa. 

Unicorn-root, true. See Chamaelirium luteum. 

Upland-cranberry. See Arctostaphylos uva-ursi. 

Urtica dioica L. Nettle family (Urticaceae). 

Stinging nettle; great nettle. 

Herbaceous, perennial plant, 2 to 4 feet high, with stinging hairs; naturalized 


from Europe and found in waste places from Canada and Minnesota south to 
South Carolina and Missouri. 


Parts used.—F lowers, leaves, and root (nonofficial). 
Uya-ursi. See Arctostaphylos uwva-ursi. 
Uvedalia. See Polymnia uvedalia. 
Uvularia perfoliata L. Bunchflower family (Melanthiaceae) . 
Perfoliate bellwort; Mohawk-weed. 
Native, perennial herb, 6 to 20 inches high; in moist woods and thickets, Quebec 
to Florida and Mississippi. 
Part used.—Root (nonofficial ). 


Vagnera racemosa (L.) Morong. Lily-of-the-valley family 
(Convallariaceae). 


Synonyms.—Convallaria racemosa L.; Sinilacina racemosa Desf. 

False Solomon’s-seal; small Solomon’s-seal; wild spikenard; false spikenard. 

Indigenous, perennial herb, 1 to 3 feet high, found in moist woods and thickets 
from Canada south to Georgia and Arizona. 


Part used.—Root (nonoflicial ). 
Valerian. See Valeriana officinalis. 
Valerian, American. See Cypripedium hirsutum. 
Valerian, American Greek. See Polemonium reptans. 
Valerian, garden-. See Valeriana officinalis. 
Valeriana. See Valeriana officinalis. 
Valeriana officinalis L. Valerian family (Valerianaceae). 
Valeriana; valerian; garden-valerian; vandal-root. 
Perennial herb, 2 to 5 feet high, native of Europe; escaped from gardens to 
roadsides in New York and New Jersey. 
Parts used.—Rhizome and roots (official). 


Vandal-root. See Valeriana officinalis. 


VANILLA, CAROLINA—-VIBURNUM LENTAGO. ie 


Vanilla, Carolina. See Trilisa odoratissima. 
Vanilla-leaf. See Trilisa odoratissima. 
Vanilla-plant. See Trilisa odoratissima. 
Velvet-plant. See Verbascum thapsus. 
Veratrum. See Veratrum viride. 
Veratrum viride Ait. Bunchfiower family (Melanthiaceae). 


Veratrum; American hellebore; swamp-hellebore; green hellebore. 

Native, perennial herb, 2 to 7 feet high, growing in swamps, wet woods, and 
meadows, Canada and Alaska, Minnesota south to Georgia. 

Parts used.—Rhizome and roots of this or |. albuin (official). 

Verbascum thapsus L. Figwort family (Scrophulariaceae). 

Mullein; velvet dock; velvet-plant; flannel-leaf. 

Tall, erect, biennial weed, sometimes 7 feet in height; naturalized from Europe 
and growing in fields, pastures, and waste places, Nova Scotia to Minnesota, 
southward to Florida. : 

Parts used.—Leaves and flowers (nonofficial ). 


Verbena hastata L. Vervain family (Verbenaceae). 
Vervain; simpler’s-joy; wild hyssop. 
Erect, indigenous perennial, 3 to 4 feet high, found in fields, meadows, and, 
waste places, Canada to Nebraska, New Mexico, and Florida. 
Parts used.—Root and herb (nonofficial). 
Veronica officinalis L. Figwort family (Scrophulariaceae). 
Common speedwell; Paul’s-betony. 


Perennial herb, 3 to 10 inches high; in dry fields and woods, Nova Scotia to 
Michigan, south to North Carolina and Tennessee. 


Part used.—Herb (nonofiticial). 
Veronica, tall. See Veronica virginica. 
Veronica virginica L.“ Figwort family (Scrophulariaceae). 
Synonym.—Leptandra virginica (L.) Nutt.@ 
Leptandra; Culver’s-root; Culver’s-physic; blackroot; Bowman’s-root; tall speed- 
well; tall veronica. 
Indigenous, perennial p'ant, 2 to 5 feet high, in moist, rich ground in woods, 
meadows, and thickets from Canada to Alabama and Nebraska. 
Parts used.—Rhizome and roots (official). 


Vervain. See Verbena hastata. 


Viburnum dentatum L. Honeysuckle family (Caprifoliaceae). 
Arrowwood; mealy-tree. 


Smooth, indigenous shrub, about 15 feet in height, growing on low ground and in 
damp woods and thickets from New Brunswick and Ontario south along the 
mountains to Georgia, and westward to Minnesota. 


Part used. —Bark (nonofficial). 


Viburnum lentago L. Honeysuckle family (Caprifoliaceae). 
Nannybush; sheepberry; sweet viburnum. 
An indigenous shrub, sometimes asmall tree; in rich soil from Canada to Georgia 
and Missouri. 
Part used.—Bark of the root of this species or of V. prunifolium official under the 
name ‘‘ Viburnum prunifolium.”’ 


aSome authors hold that this plant belongs to the genus Leptandra and that its name should be 
Leptandra virginica (L.) Nutt. The Pharmacopceia is here followed. 


74 WILD MEDICINAL PLANTS OF THE UNITED STATES. 


Viburnum opulus. See Viburnum opulus L. 


Viburnum opulus L. Honeysuckle family (Caprifoliaceae). 
Viburnum opulus; cramp-bark; high-bush cranberry; squawbush. 


Indigenous shrub, 4 to 10 feet in height, found in low, rich woods and borders 
of fields from New Jersey, Michigan, and Oregon, northward. 


Part used.—Bark (official). 
Viburnum prunifolium. See Viburnum lentago and V. prunifolium L. 


Viburnum prunifolium L. Honeysuckle family (Caprifoliaceae). 
Black haw; sloe; stagbush. 


Indigenous shrub or small tree, growing in dry woods and thickets and on rocky 
hillsides, Connecticut to Florida, west to Michigan and Texas. Most abun- 
dant in the South. 


Part used.—Bark of the root of this species or of J”. /entago official under the 
name ‘‘ Viburnum prunifolium.”’ 


Viburnum, sweet. See Viburnum lentago. 
Vine-maple. See Menispermuim canadense. 


Viola odorata L. Violet family (Violaceae). 
English violet; sweet violet; March violet. 
Low herb, native of Europe; escaped from gardens, Nova Scotia to New York 
and New Jersey, and on the Pacific coast. 
Part used.—F lowers (nonofficial ). 
Viola pedata L. Violet family (Violaceae). 
Bird’s-foot violet; wood-violet; snake-violet. 


Native plant, perennial, 3 to 10 inches high, occurring in dry fields and on _hill- 
sides from Maine to Minnesota, south to Florida and Missouri. 


Parts used.—Herb and root (nonofificial ). 


Viola tricolor L. Violet family (Violaceae). 
Pansy; heartsease. 


Small herb, 4 to 12 inches high, introduced from Europe; found in waste places, 
sparingly escaped from gardens. 


Part used.—F lowering herb (nonofficial ). 
Violet, bird’s-foot. See Viola pedata. 
Violet, dog’s-tooth. See Hrythronium americanum. 
Violet, English. See Viola odorata. 
Violet, March. See Viola odorata. 
Violet, rattlesnake-. See Hrythronium americanum. 
Violet, snake-. See Viola pedatu. 
Violet, sweet. See Viola odorata. 
Violet, wood-. See Viola pedata. 
Violet-bloom. See Solanum dulcamara. 
Virginia creeper. See Parthenocissus quinquefolia. 
Virgin’s-bower. See Clematis virginiana. 
Viscum flavescens Pursh. Same as Phoradendron flavescens. 


Vomitwort. See Lobelia inflata. 


Wafer-ash. See Pelea trifoliata. 

Wahoo. See Luonymus atropurpureus. 

Wake-robin. See Arisaema triphyllum and Trillium erectum. 
Walnut, white. See Juglans cinerea. 


Wartwort. See Gnaphalium uliginosum. 


WASHINGTONIA LONGISTYLIS—-WOOD-FERN, EVERGREEN. 5. 


Washingtonia longistylis (Torr.) Britton. Parsley family (Apiaceae). 
Synonym.—Osmorrhiza longistylis DC. 
Sweet-cicely; anise-root; sweet chervil. 


Erect, rather stout, perennial herb, 2 to 3 feet high, native; in rich, moist woods 
and banks of streams from Canada to Alabama and Texas. 


Part used.—Root (nonofficial ). 
Water-avens. See Geum rivale. 
Water-bugle. See Lycopus virginicus. 
Watercup. See Sarracenia flava and S. purpurea. 
Watercup, yellow-flowered. See Sarracenia flava. 
Water-eryngo. See Eryngium yuccifolium. 
Water-flag. See Iris versicolor. 
Water-hemlock. See Cicuta maculata. 
Water-hoarhound. See Lycopus virginicus. 
Water-lily. See Castalia odorata. 
Water-lily, sweet-scented. See Castalia odorata. 
Water-pepper. See Polygonum hydropiper. 
Water-shamrock. See Menyanthes trifoliata. 
Water-smartweed. See Polygonum punctatum. 
Waxberry. See Myrica cerifera. 
Wax-myrtle. See Myrica cerifera. 
Waxwork. See Celastrus scandens. 
Waythorn. See Rhamnus cathartica. 
White-bark. See Populus alba. 
Whiteroot. See Asclepias tuberosa. 
Whitethorn. See Crataegus oxyacantha. 
Whitewood. See Liriodendron tulipifera and Tilia americana. 
Wickopy. See Dirca palustris. 
Wickup. See Chamaenerion angustifolium and Epilobiuin palustre. 
Willow, black. See Salix nigra. 
Willow, European. See Salix alba. 
Willow, pussy-. See Salix nigra. 
Willow, rose-. See Cornus amomum. 
Willow, swamp-. See Salix nigra. 
Willow, white. See Salix alba. 
Willow-herb, great. See Chamaenerion angustifolium. 
Willow-herb, night. See Oenothera biennis. 
Willow-herb, swamp. See Epilobium palustre. 
Wingseed. See Ptelea trifoliata. 
Winterberry, Virginia. See Ilex verticillata. 
Winterbloom. See Hamamelis virginiana. 
Wintergreen. See Gaultheria procumbens. 
Wintergreen, bitter. See Chimaphila umbellata. 
Witch-hazel. See Hamamelis virginiana. 
Woodbine, wild. See Gelsemiwm sempervirens. 
Wood-fern, evergreen. See Dryopteris marginalis. 


76 WILD MEDICINAL PLANTS OF THE UNITED STATES. 


Wood-sorrel, white. See Oxalis acetosella. 
Wood-violet.. See Viola pedata. 

Worm-grass. See Spigelia marilandica. 

Wormseed. See Chenopodium anthelminticum. 
Wormseed, American. See Chenopodium ambrosioides. 
Wormwood. See Artemisia absinthium. 


Wormwood, Roman. See Ambrosia artemisiaefolia. 


Xanthium spinosum L. Ragweed family (Ambrosiaceae). 
Spiny clotbur; spiny burseed; thorny clotweed; thorny burweed. 


An annual weed, 1 to 3 feet high, naturalized from Europe or Asia; in waste 
ground, Ontario to Florida, westward to Missouri and Texas. 


Part used.—Leaves (nonofficial ). 


Xanthorrhiza apiifolia L’ Her. Crowfoot family (Ranunculaceae ). 

Shrub yellowroot; southern yellowroot. 

Low, shrubby, indigenous perennial, 1 to 2 feet high, growing in woods and 
along river banks, southwestern New York to Florida, chiefly in the moun- 
tains. 

Parts used.—Rhizome and roots (nonofficial). 

Xanthoxylum. See Fagara clava-herculis and Xanthoxylum americanum. 


Xanthoxylum americanum Mill. Rue family (Rutaceae). 
Synonym.— Xanthoxylum fraxineum Willd. i 
Xanthoxylum; northern prickly ash; toothache-tree. 


Indigenous shrub or small tree, maximum height about 25 feet; common in 
woods and thickets and along river banks from Virginia, Missouri, and 
Nebraska northward to Canada. 


Parts used.—Bark of this or of Fagara clava-herculis official under the name 
‘‘Xanthoxylum.’’ Berries (nonofficial). ’ 


Xanthoxylum clava-héerculis L. Same as Fagara clava-herculis. 


Xanthoxylum fraxineum Willd. Same as Xanthoxylum americanum. 


Yam, wild. See Dioscorea villosa. 

Yarrow. See Achillea millefolium. 

Yellowroot. See Coptis trifolia, Hydrastis canadensis, and Jeffersonia diphylla, 
Yellowroot, shrub. See Xanthorrhiza apiifolia. 

Yellowroot, southern. See Yanthorrhiza apiifolia. 

Yellowthorn. See Fagara clava-herculis. 

Yellowwood. See Fagara clava-herculis. 

Yerba buena. See Micromeria chamissonis. 

Yerba reuma. See Frankenia grandifolia. 

Yerba santa. See HMriodictyon californicum. 


Youthwort. See Drosera rotundifolia and Heracleum lanatum. 


O 


a 
Ap 
U. S. DEPARTMENT OF AGRICULTURE. 
BUREAU OF PLANT INDUSTRY—BULLETIN NO. 90, PART I. 


B. T. GALLOWAY, Chief of Bureau. 


THE STORAGE AND GERMINATION OF 
WILD RICE SEED. 


BY 


oe Weel -DUVEL, 


ASSISTANT IN THE SEED LABORATORY. 


IssuED SEPTEMBEE 7, 1905. 


— 


MATION Of 
TSS 


WASHINGTON: 
GOVERNMENT PRINTING OFFICE. 
19:05: 


LORS HRS Sey Tay 2 Soe ea Se ae ce rr ee Sa siety ee Pea ee 
ce SS ELUEODRD GE, 2 es a a ee ere ae a ee 


aseeccamn versus spring Seeding .-/-/2.2...-... 2-22) -.-l scene cess eee. = 
De NnnEEEE SEOTIN® the secu 27. 2s See te 2 a ee ee 2 
Detailed conditions and results of storage experiments -...........---------- 
Penman shOriation! 5... Us soi) o£ eo a. Jb S2 elon eile loeb eke ane 
P_ieaodinOLmaking sermination tests... =o. .c...--4<--- -+2-----~-i bi. --- 
Peer eeiemperiture On germination ..2...+..-:...2--.--2---.-s202+- +--+ 
SLED 22 bac ESE eee oo ees een ee yo ae a 
SPEEDS Toi Settee a eke oe oa os ani ee ein ocacecae- 205% 


3 


“I o> Or or’ 


~I 


co CO 


ILLUSTRATIONS: 


Page. 
Pirate I. Wild rice growing in water after being kept wet in cold storage at a 
temperature of 32-34° F., from October 19, 1903, to November 15, 
1904 . 22 - b asec tees sc ce ne ae es fens e ee oe ee 16 
II. Stages of germination of wild rice, showing the development of the 
root system and the relative position of the seedling and the parent 


~ 


B.: P. I.—178. 


THE STORAGE AND GERMINATION OF WILD 
RICE SEED.’ 


INTRODUCTION. 


The seed of wild rice, sometimes called Indian rice or water oats 
(Zizania aquatica L.), has always been a very valuable food among 
the Indians, especially those of the upper Mississippi Valley. Of 
recent years wild rice has found a place on the menu cards of some 
of our best American hotels. The rich and highly nutritious grains, 
together with the slightly smoky flavor it has when properly pre- 
pared, make it an extremely palatable article of diet. If it were not 
for the difficulties of harvesting the seed and preparing the finished 
product for market it is probable that wild rice would find a place in 
many American homes. 

At present, however, the greatest interest in wild rice is created by 
the value of the seed as a food for wild waterfowl, particularly wild 
ducks. Asa result of this interest the propagation of wild rice from 
seed has become a question of considerable importance, especially to 
the members of the gunning clubs throughout the United States and 
Canada. 

DISTRIBUTION. 


The distribution of wild rice is now reported from New Brunswick 
and Assiniboia south to Florida, Louisiana, and Texas. There are, 
however, comparatively few localities in which it grows abundantly. 


a Wild rice is considered one of the most important foods for wild ducks and other 
waterfowl, and a large number of inquiries have been received from members of 
gunning clubs throughout the United States asking where good, germinable seed can 
be secured. It is quite generally refognized that wild rice seed loses its vitality if 
allowed to become dry, and better methods of storing the seed during the winter 
have long since been demanded. 

The results of investigations begun two years ago show that wild rice seed can. be 
handled without any deterioration in vitality if it is harvested and stored according 
to methods outlined in the present paper. 

J. W. T. Duvet, Acting Botanist in Charge of Seed Laboratory. 


Seep Lasorarory, 
Washington, D. C., July 20, 1905. 


6 STORAGE AND GERMINATION OF WILD RICE SEED. 


Good reasons exist for assuming that this area can be extended to 
include all fresh-water lakes, as well as swamps and river bogs, where 
the water does not become stagnant, throughout the whole of North 
America south of latitude 55° north. Wild rice also grows luxuri- 
antly along the lower parts of many of the rivers of the Atlantic Coast 
States, the waters of which are affected by the action of the tide to a 
considerable degree, and consequently contain an appreciable quantity 
of salt. It has been shown” that the maximum degree of concentra- 
tion of salt water in which wild rice plants can grow successfully is 
equivalent to a 0.03 normal solution of sodium chlorid. This concen- 
tration corresponds to 0.1755 per cent by weight of sodium chlorid, 
which is sufficient to give a slight salty taste to the water. 


HABITAT. 


While it is well recognized that the habitat of the wild rice plant is 
in shallow fresh water, it is now known that it will grow luxuriantly 
in water containing little less than two-tenths of 1 per cent of sodium 
chlorid. Occasional plants have been found growing in water which 
contained, for short periods at least, nearly double that amount of 
salt. These facts indicate the possibility of a much wider range of 
conditions to which this plant may be subjected without hindering its 
development. It is not beyond the range of possibility—indeed, it is 
quite probable—that by careful selection plants may be obtained which 
will thrive on soil that is comparatively dry, at least in places in which 
the water can be drawn off gradually during the latter part of the 
growing season. 

In September, 1904, Mr. G. C. Worthen, of the Bureau of Plant 
Industry, collected a cluster of wild rice plants which were growing 
on the Potomac Flats, near Washington, D. C., in soil which was suf- 
ficiently dry to permit the use of a 2-horse mowing machine for cutting 
down the rank growth of vegetation. This was newly made land, and 
in all probability the seed giving rise to this cluster of plants was 
pumped in with the dirt from the Potomac River the year previous. 

This amphibious type once established, it will undoubtedly carry 
with it a strain of seed which can withstand considerable drying with- 
out any marked injury to its vitality. Such being true, the methods 
and difficulties of propagation from seed would be greatly simplified. 

Simultaneous with establishing an amphibious type should come the 
selection of seed plants which are capable of retaining their seed until 
the larger part of it has reached maturity. These two steps once 
made, the future of wild rice as a cereal will be assured. 


«The Salt Water Limits of Wild Rice. Bulletin No. 72, Part Il, Bureau of Plant 
Industry, United States Department of Agriculture, 1905. 


FALL SEEDING VERSUS SPRING SEEDING. 7 
GERMINATION OF THE SEED. 


The greatest difficulty to be overcome in extending the area for 
growing wild rice is the poor germination of the commercial seed. 
Inasmuch as wild rice constitutes one of the most important foods of 
wild ducks and other wild waterfowl, many individuals and most of 
the gunning clubs east of the Rocky Mountains have been asking the 
question, How can we propagate wild rice from seed in order to estab- 
lish better feeding and fattening grounds for our game birds? 

The many failures in the propagation of wild rice from seed have 
been due to the use of seed that had become dry before sowing, or to 
the fact that the seed when sown fresh in the autumn had been eaten 
by ducks or other animals or was carried away by heavy floods before 
germination took place. 

It is now very generally known that the seed of wild rice, if once 
allowed to become dry, will not germinate, save possibly an occasional 
grain. In its natural habitat the seed, as soon as mature, falls into the 
water and sinks into the mud beneath, where it remains during the 
winter months, germinating the following spring if conditions are 
favorable. 

Heretofore the plan generally followed, and the one usually recom- 
mended by those who have given some attention to the propagation of 
wild rice, was practically that of natural seeding; that is, to gather the 
seed in the autumn, as soon as thoroughly mature, and, while still fresh, 
to sow it in | to 3 feet of water. 


FALL SEEDING VERSUS SPRING SEEDING. 


It must be remembered that the bulk of the seed remains dormant 
during the winter, germinating first the spring after maturing; con- 
sequently, with but few exceptions, fall seeding is unsatisfactory and 
unreliable. Fall seeding is likely to prove a failure for three reasons: 
(1) Wild ducks and other animals of various kinds eat or destroy the 
seed in considerable quantity before it has had time to germinate the 
following spring; (2) much of the seed is frequently covered so deeply 
with mud that washes in from the shore during the winter that the 
young plants die of suffocation and starvation before they reach the 
surface; (3) in some cases a large quantity of the seed is carried away 
from the place where sown by the high waters and floating ice prev- 
alent during the latter part of the winter and early spring. 

In exceptional cases these difficulties can be overcome; under which 
circumstances autumn sowing may be preferable to spring sowing. 
In the majority of cases, however, much better results will be 
obtained if the seed is properly stored and sown in the early spring, 
as soon as the danger of heavy floods is passed and the water level 
approaches normal. 


8 STORAGE AND GERMINATION OF WILD RICE SEED. 


In sowing the seed considerable care must be exercised in selecting 
a suitable place, securing the proper depth of water, etc. Good 
results can be expected if the seed is sown in from 1 to 3 feet of water 
which is not too stagnant or too swiftly moving, with a thick layer of 
soft mud underneath.” It is useless to sow wild rice seed on a gravelly 
bottom. or in water where the seed will be constantly disturbed by 
strong currents. 

Previous to this time, save in a few reported cases, the seed which 
was allowed to dry during the winter and was sown the following 
spring gave only negative results. It is now definitely known that 
wild rice, if properly handled, can be stored during the winter without 
impairing the quality of germination to any appreciable degree, and 
that it can be sown the following spring or summer with good success. 


DIRECTIONS FOR STORING THE SEED. 


The vitality of wild rice seed is preserved almost perfectly if kept 
wet in cold storage—Nature’s method of preservation. This method 
of storage implies that the seed has been properly harvested and cared 
for up to the time of storage. The seed should be gathered as soon 
as mature, put loosely into sacks (preferably burlap), and sent at once 
to the cold-storage rooms. If the wild rice fields are some distance 
from the cold-storage plant the sacks of seed should be sent by express, 
and unless prompt delivery can be guaranteed it is not advisable to 
send by freight even for comparatively short distances. It is very 
important, that the period between the time of harvesting and the 
time when the seed is put into cold’ storage be as short as possible. 
If this time is prolonged to such an extent as to admit of much fer- 
mentation or to allow the seed near the outside of the bags to become 
dry during transit, its vitality will be greatly lowered. 

It is not practicable to give any definite length of time which may 
elapse between harvesting and storing, inasmuch as the temperature, 
humidity, and general weather conditions, as well as the methods of 
handling the seed, must be taken into consideration. Let it suffice to 
say, however, that the vitality of the seed will be the stronger the 
sooner it is put into cold storage after harvesting. 

As soon as the seed is received at the cold-storage plant, while it is 
still fresh and before fermentation has taken place, it should be put 
into buckets, open barrels, or vats, covered with fresh water, and 
placed at once in cold storage. If there is present a considerable 
quantity of light immature seed or straw, broken sticks, ete., it will 
be profitable to separate this from the good seed by floating in water 


« Wild Rice: Its Uses and Propagation. Bulletin No. 50, Bureau of Plant Industry, 
United States Department of Agriculture, 1903. 


STORAGE EXPERIMENTS. 9 


preparatory to storing. The storage room should be maintained at 
a temperature just above freezing—what the storage men usually 
designate as the *‘ chill room.” 

When taken from cold storage in the spring the seed must not be 
allowed to dry out before planting, as a few days’ drying will destroy 
every embryo. 

Seed which was stored under the foregoing conditions from October 
19, 1903, to November 15, 1904, 393 days, germinated from 80 to 88 
per cent. Another lot of seed, which was stored on October 6, 1904, 
and tested for vitality on April 17, 1905, germinated 79.8 per cent. 

Plate I shows the luxuriant growth made by the seed which was kept 
wet and stored at a temperature of 32° to 34° F. for 393 days. 


DETAILED CONDITIONS AND RESULTS OF STORAGE EXPERI- 
MENTS. 


The foregoing conclusions are based on the results obtained from 
two series of experiments, as follows: 


In October, 1903, a box of wild rice seed was received from Ontario. 
Canada. This seed, as soon as gathered, was loosely packed in moist 
sphagnum and sent by express to the Seed Laboratory of the United 
States Department of Agriculture. After a few days, while it was 
yet moist and before any fermentation had taken place, the seed was 
divided into four lots for special treatment, as follows: 

(1) Seed submerged in water and placed in cold storage at a temper- 
ature of 32° to 34° F. 

(2) Seed submerged in water and placed in cold storage at a temper- 
ature of 12° F. The seed was soon embedded in a solid mass of ice 
and remained so until samples were taken for test. 

(3) Seed, without the addition of water, put into cloth bags and kept 
in cold storage at a temperature of 32° to 34° F. 

(4) Seed, without the addition of water, put into cloth bags and kept 
in cold storage at a temperature of 12° F. 


In October, 1904, a seeond consignment of seed was received from 
Minnesota, and the following additional storage experiments were 
made by Mr. C. 8. Scofield, of the Bureau of Plant Industry. 

(5) Seed submerged in water and placed in cold storage at a temper- 
ature of 32° to 34° F., as in No. 1. 

(6) Seed submerged in water and placed in cold storage at a temper- 
ature of 12° F., as in No. 2. 

(7) Seed submerged in water in a galyanized-iron bucket and stored 
on the roof of the laboratory building. The water was changed daily 
when not frozen. 


10 STORAGE AND GERMINATION OF WILD RICE SEED. 


(8) Seed submerged in water in a galvanized-iron bucket and stored 
on the roof of the laboratory building, as in No. 7. In this case the 
water was not changed save to replace the loss due to evaporation. 

(9) The conditions for No. 9 were the same as those for No. 8, 
except that air was forced into the water daily when not frozen solid. 


Samples of seed were taken from the different lots and tested for 
vitality at irregular intervals throughout the time of storage, which, 
in the former series, extended over a period approximately thirteen 
months and in the latter series over a period of little more than six 
months. 

Experiments Nos. 1 and 5.—The seed which was submerged in water 
and stored in the ‘‘chill room” showed no deterioration in vitality. 
The results of the final tests gave a germination varying from 79.8 to 
- 88 percent. This is practically Nature’s method of preserving the 
vitality of the seed during the winter. 

Experiments Nos. 2 and 6.—The seed which was submerged in 
water and stored at a temperature of 12° F. was all killed before the 
spring following the date of storage. Soon after being placed in stor- 
age the water was frozen solid and the seeds were embedded in a mass 
of ice, in which condition they remained throughout the experiment, 
a portion being cut out from time to time for germination tests. The 
complete loss of vitality in these two lots of seed is attributed not to 
the freezing directly, but to the thorough desiccation as a result of 
the continuous low temperature. 

Experiments Nos. 3 and 4.—The samples of seed which were stored 
in cloth bags at the temperatures of 32° to 34° F. and of 12° F. had, 
for all economic purposes, entirely lost their vitality. The average 
percentage of germination, as shown by the 37 tests made from each 
of the two lots, was less than five-tenths of 1 per cent. 

Experiment No. 7.—The seed which was submerged in water and 
stored on the roof of the laboratory building, the water being changed 
daily, showed a good percentage of germination when the last vitality 
tests were made. If only a small quantity of seed is desired for the 
spring planting and cold storage can not be readily secured, good 
results may be obtained ‘by this treatment; but it is much less certain 
and probably more expensive than keeping the seed in cold storage, 
and for this reason is not recommended. The success of this method 
will likewise depend largely on the temperature of the water. 

Experiments Nos. 8 and 9.—On April 22, 1905, samples taken from 
each of these two lots of seed showed a marked deterioration in vitality. 
Thoroughly mixed samples from No. 8 showed a vitality of only 58 
per cent, while No. 9 had deteriorated to 14.3 per cent. 


METHODS OF MAKING GERMINATION TESTS. 11 
PACKING FOR TRANSPORTATION. 


Too much care can not be given to the matter of packing the seed 
for transportation, for unless the packing is properly done the vitality 
of the seed will be destroyed during transit. What is here said applies 
to fresh seed which is to be sown in the autumn, as well as to seed 
which has been kept in cold storage during the winter. It must not 
be forgotten, however, that the vitality of cold-storage seed is more 
quickly destroyed on drying than that of fresh seed. 

For transportation the seed should be carefully packed, with moist 
sphagnum, cocoanut fiber, or fine excelsior, in a loosely slatted box. 
If the time of transportation does not exceed five or six days no spe- 
cial precautions need be taken as to the temperature. During the 
period of transportation it is quite probable that some of the seed will 
germinate, but if sown at once growth will not be retarded and the 
roots will soon penetrate the soil and anchor the young plants. 

If the time of transportation is necessarily long, it is recommended, 
if the best results are desired, that some provision be made for a 
reduced temperature. The nearer the temperature approaches that of 
freezing the better. It has been demonstrated, however, that a fair 
percentage of seed will remain germinable for a considerable time if 
packed as above described. 

On October 10, 1904, Mr. C. S. Scofield sent a small quantity of 
wild rice, packed in moist sphagnum moss in a well-ventilated box, to 
Doctor De Vries, of Amsterdam, Holland. On October 14 or 15 this 
box was placed in cold storage on the steamer in New York Harbor. 
The box of seed was received by Doctor De Vries in good condition on 
November 2, twenty-one days after the seed was packed for shipment. 


METHODS OF MAKING GERMINATION TESTS. 


The samples were tested (1) between folds of blotting paper—our 
regular method for testing the germination of most seeds—and (2) in 
water, Nature’s method of sowing wild rice seed. The latter method 
gave much better results and was the one finally adopted for the 
laboratory tests. The seed should be covered with water, the water 
in the dishes to be changed daily. 

Plate I shows the importance of making the germination tests in 
water, as described in the foregoing paragraph. The seed was covered 
with water and placed in a germinating chamber maintained at an 
alternating temperature of 20° C. (68° F.) for eighteen hours, and 
30° C. (84° F.) for six hours, until the majority of the seeds had 
germinated. At this stage the dish containing the seeds was trans- 
ferred to the worktable, which was exposed to the temperature of the 
laboratory—approximately that of a living-room. The water in the 


12 STORAGE AND GERMINATION OF WILD RICE SEED. 


dish was changed daily during the period of germination, and water 
was afterwards added at irregular intervals to replace the loss by 
evaporation. 

Plate II] shows somewhat in detail the different stages in the germina- 
tion of wild rice seeds. The seeds and seedlings are shown in natural 
size. In/andc the first sheath has just burst through the seed coats, 
taking a position at right angles to the seed proper. The lateral roots 
begin to emerge when the first sheath leaf has attained a length of 4 to 
13 inches. From this time growth continues rapidly, and by the time 
the seedlings are 2 or 3 inches long the root system is very well 
developed (7 and g). At this stage under favorable conditions the 
plants have a good hold in the soil and will not be washed away by an 
ordinary freshet. The relative position of the actively growing 
seedling is always at right angles to that of the old seed, as shown in 
fand g. 


EFFECT OF TEMPERATURE ON GERMINATION. 


Germination tests were made at constant and alternating tempera- 
tures, ranging from 15° to 35° C. (59° to 95° F.). While no effort 
was made to show the minimum and maximum temperatures of ger- 
mination, the percentage was somewhat reduced at a constant tempera- 
ture of 35° C., and the maximum is not much above that. All of the 
other temperatures gave good results. The lower temperatures, how- 
ever, were slightly more favorable than the higher. These facts are 
valuable to show that the wild rice plant can thrive in either warm or 
cold water, but better, perhaps, in northern than in southern latitudes. 


SUMMARY. 


(1) Under no circumstances should wild rice seed which is intended 
for planting be allowed to dry. Dried seed will germinate but rarely 
and should never be sown. 

(2) Wild rice seed can be stored without deterioration if it is gath- 
ered as soon as matured, put into barrels or tanks, covered with fresh 
water, and, before fermentation has set in, stored at a temperature of 
32-34- F. Seed treated in this way germinated as high as 88 per cent 
after being in storage 393 days. Fresh seed seldom germinates better, 
and usually not so well. 

(3) After the seed is taken from cold storage it should not be 
allowed to dry. The vitality of cold-storage seed is destroyed on 
drying even more quickly than that of fresh seed. 

(4) For transportation the seed should be packed in moist sphagnum, 
cocoanut fiber, or fine excelsior. If not more than five or six days 
are required for transit, no special precautions need be taken for con- 
trolling the temperature; but if the time for transportation exceeds 


SUMMARY, 13 


six days, provision should be made for a temperature sufliciently low 
to prevent marked fermentation. A temperature approximately 
freezing will give the most satisfactory results. 

(5) Wild rice can be sown either in the autumn or in the spring. 
Spring sowing is preferable, thus avoiding the danger of having the 
seed eaten or destroyed by wild ducks or other animals during the fall 
or winter, or of its being buried or washed away by the heavy floods 
of late winter or early spring. 

(6) Wild rice should be sown in the spring in from 1 to 3 feet of 
water which is neither too stagnant nor too swiftly moving, as soon as 
the danger of heavy floods is passed. 

(7) Wild rice is of the greatest importance as a food for wild water- 
fowl, likewise a delicious breakfast food for man, and the area in which 
it is extensively grown should be extended. It will grow luxuriantly 
in either warm or cold water; furthermore, it can be grown success- 
fully in water which is slightly salty to the taste. 

(8) In determining the vitality of any sample of wild rice seed the 
germination tests should be made in water—the condition under which 
the self-sown seed germinates. 

(9) The seed will germinate well at temperatures ranging from 
15° to 30° C. The maximum temperature of germination is above 
35° C. (95° F.), but better results are obtained at lower temperatures. - 


te u 7 cs i 
haps So pam 7? 


vy Ween, OS 


say 


~ 


Pees BO 


DESCRIPTION OF PLATES. 


Piate I. Wild rice growingin water. This seed was submerged at a temperature of 
32-34° F. for approximately thirteen months. In making the germination test 
the seed was covered with water and placed in a germinating chamber main- 
tained at a temperature of 20° C. (68° F.) for eighteen hours, and at 30° C. 
(86° F.) for six hours. After the majority of the seeds had germinated the dish 
was transferred to the worktable of the Seed Laboratory. 


Pate II. Progressive stages in the development of wild rice seedlings; f and g, seed- 
lings showing the relative position of the growing seedlings and the parent seed, 
which take a position at right angles to each other when grown normally in 
water. (Natural size. ) 

16 


Bul 90, Pt. |, Bureau of Plant Industry, U. S. Dept. of Agriculture. PLATE |. 


> 


piste 2 


WILD RICE GROWING IN WATER AFTER BEING KEPT WET IN COLD STORAGE AT A TEM- 
PERATURE OF 32-34° F., FROM OCTOBER 19, 1903, TO NovemBeER 15, 1904. 


Bul. 90, Pt. |, Bureau of Plant Industry, U. S. Dept. of Agriculture. PLATE II. - 
a 


\ 
| 


5) =e] 


| | 
| | a 
, 7 
4 


H a C 


* 


ia 


L 


STAGES OF GERMINATION OF WILD RICE, SHOWING THE DEVELOPMENT OF THE 
Root SYSTEM AND THE RELATIVE POSITION OF THE SEEDLING AND THE 
PARENT SEED. NATURAL SIZE. 


Ap 


eo VEPAKRTMENT OF AGRICULTURE. 
BUREAU OF PLANT INDUSTRY—BULLETIN NO. 90, PART II. 


B. T. GALLOWAY, Chief of Bureau. 


THE CROWN-GALL AND HAIRY-ROOT DISEASES 
OF THE APPLE TREE. 


BY 


GEORGE G. HEDGCOCK. 


ASSISTANT IN PatHoLoay, MIssissrppr VALLEY LABORATORY. 


- e 
VEGETABLE PATHOLOGICAL AND PHYSIOLOGICAL 
INVESTIGATIONS. 


IssUED NOVEMBER 17, 1905. 


WASHINGTON: 
GOVERNMENT PRINTING OFFICE. 
1905. 


4 
CONTENTS. 

Te a sah erie» a ae ae a 0 ee ae eee 
Two distinct diseases, crown-gail and hairy-root....................-------- 
EE NREINE UH EE NER Se Se 5 rs Peo ics pn ne See Som a ce ee 
Effect upon the length of life of the apple tree..........-...-.-.--.-.------- 
SPEennn ee MEIRAIVINOGH 920. fc 22-2 ee ee eee eee ae een be 
a AS a EE eS ee 

3 


on transplanted seedling. Fig. 3.—Hairy-root 

apple tree. Fig. 4.—Hairy-root disease on gr 1 

II. Fig. 1.—Apple seedlings diseased with hairy-root. 
seedlings diseased with soft crown-gall .....-- 

III. Fig. 1.—Healthy fibrous-rooted apple tree, pot 
Apple seedlings diseased with hairy-root...----. 


4 


2 
oN 
é ° <i 
we 
i ie, 
a 
a¢ i - 
- i 
+e } 
: oust Av ant, 
- r av 
3 ; > a 


Bul. 90, Pt. Il, Bureau of Plant Industry, U. S. Dept. of Agriculture. PLATE I. 


Fic. 1.—APPLE CROWN-GALL ON Fic. 2.—APPLE CROWN-GALL ON 
GRAFTED TREE. TRANSPLANTED SEEDLING. 


Fic. 3.—HairRy-RooT DISEASE ON Fia. 4.—HairyY-RootT DISEASE ON 
GRAFTED APPLE TREE. GRAFTED APPLE TREE. 


PLATE II. 


Bu). 90, Pt. Il, Bureau of Plant Industry, U. S. Dept. of Agriculture. 


Fic. 1.—APPLE SEEDLINGS DISEASED WITH HAIRY-ROoT. 


Fia. 2.—APPLE SEEDLINGS DISEASED WITH SOFT CROWN-GALL. 


*"NMOUF) LOd 'sayu) AlddV GSLOOY-SNOUSIA AHLIVAH—' | ‘DdI4 


“LOOY-AYIVH HLIM G3aSV3SIg SONIIGSAS AlddyW—'? ‘Ol 


Bul. 90, Pt. ll, Bureau of Plant Industry, U. S. Dept. of Agriculture 


PLATE III. 


B. P. 1.—186. V. P. P. I.—145. 


THE CROWN-GALL AND HAIRY-ROOT DISEASES 
OP HE APPLE. TREE. 


INTRODUCTION. . 


- The diseases of the apple which have been classed under the name 
crown-gall have, during the last few years, attracted much attention, 
due partly to an increase of these diseases and partly to the enacting 
of more stringent State laws governing the shipment and inspection of 
. trees. 

A series of investigations into the nature of crown-gall upon the 
apple, pear, raspberry, peach, almond, grape, rose, and other plants 
has been in progress for some time in the Mississippi Valley Labora- 
tory of the Bureau of Plant Industry at St. Louis, Mo., and also at 
other points in the Mississippi Valley. It is not to be assumed, how- 
ever, that such diseases are more common in this locality than in some 
other portions of the United States. Apple crown-gall and hairy-root 
have been found in all nurseries that have been examined in various 
portions of the country. 

This preliminary report is sent out, not with the intention of giving 
the results of all our investigations, but for the purpose of calling the 
attention of apple-tree growers to the different diseases hitherto known 
as apple crown-gall, and to endeavor to interest them in the collection 
of data regarding the predisposition of varieties to these diseases. 


TWO DISTINCT DISEASES, CROWN-GALL AND HAIRY-ROOT. 


Our investigations have resulted first in separating apple crown-gall 
into two diseases, which are considered distinct. The disease now 
designated as crown-gall isa callous-like gall growth of hypertrophied 
tissue following wounds on some portion of the root system of the 

tree, which rarely occurs above the ground on parts of the trunk or 
limbs. (See PI. I, figs. 1 and 2.) 

The malady now called the hairy-root disease is evidently the same 
as the one first given this name by Stewart, Rolfs, and Hall in Bulletin 
191 of the New York State Experiment Station. It is characterized 
both in seedlings (PI. II, fig. 1, and PI. III, fig. 2) and in grafted or 


5 


6 CROWN-GALL AND HAIRY-ROOT DISEASES OF THE APPLE TREE. 


budded trees (PI. I, figs. 3 and 4) by a stunted root system, accom- 
panied with an excessive production of small fibrous roots, often origi- 
nating in clusters from the main root, or taproot. Galls often occur 
in connection with hairy-root, but these are a result of wounds rather 
than a form of this disease. Seedlings of the hairy-root type, unless 
wounded, remain free from galls. 


TYPES OF APPLE CROWN-GALL. 


Apple crown-gall is of two types. <A hard callous form is common 
on grafted trees at the union of the root and scion, and at any other 
point of the root system where wounds occur in either the cultivation 
or transplanting of trees (PI. I, fig. 1). The results of extensive inoc- 
ulations with this type have failed to prove that this disease is of a 
contagious nature. ; 

A second type is a soft form more common on seedlings (PI. I, 
fig. 2), occurring more rarely on grafted trees (PI. I, fig. 2). These 
softer galls resemble those of the raspberry and peach, in that they 
are soft and often rot off. It is not certain, however, that they, like 
the latter, are replaced the following year by a new gall growth from 
the adjacent live tissues of the host, nor is there proof yet that they 
are of a contagious nature. 


EFFECT UPON THE LENGTH OF LIFE OF THE APPLE TREE. 


Careful data are being collected from orchards and nurseries as to 
the effect of these diseases upon the life and fruitfulness of trees. 
Any information as to the locality of orchards in which diseased trees 
have been planted will be highly appreciated. In our crown-gall 
orchard there are more than 200 trees diseased with the hard type of 
crown-gall, and 200 healthy trees of the same grade planted under 
similar conditions. After two years’ growth six of the crown-gall 
trees and nine of the healthy ones have died. No difference in the 
growth of the trees is noticeable. However, it can not be assumed 
from the results so far that, on the one hand, the disease may not yet 
shorten the life of the trees, or, on the other, that the trees may not 
entirely overcome its effects. A tree having crown-gall on its roots, 
however, can never be correctly graded with a smooth-rooted tree. 
The root system of a healthy fibrous-rooted apple tree is shown in 
Plate II], tigure 1. 


SUGGESTIONS TO NURSERYMEN. 


Nurserymen are advised to be careful in the selection of seedlings 
for grafting and budding. All rough, warty, or galled seedlings 
should be thrown out, for most of them will form rough-rooted trees. 
Seedlings with tufted or hairy roots should also be rejected, for these, 


DATA DESIRED. 7 


as shown by our experiments, develop into hairy-rooted trees with a 
very deficient root system. The hairy-root disease, as it appears from 
the results of two years’ experiments, is not contagious. It is hoped 
in the near future to be able to offer some practical means of reducing 
the percentage of trees affected with these diseases in the nursery. 


DATA DESIRED. 


The hearty cooperation of nurserymen and orchardists in securing 
data is desired. It is hoped to secure the help of the leading nursery- 
men of this country in getting an accurate count from each nursery 
of the number of diseased trees in at least one row of every variety 
in all fields where the trees are all dug in one season. Such data are 
desired from every locality where apple trees are grown. Printed 
blanks with directions for tabulating such data have been provided 
and these will be sent to all who request them. Address the Missis- 
sippi Valley Laboratory, St. Louis, Mo. 


O 


feo enrAklT MENT OF AGRICULTURE. 
BUREAU OF PLANT INDUSTRY—BULLETIN NO. 90, PART Il. 


B. T. GALLOWAY, Chief of Bureau. 


ee PR Mei NE, 


BY 


ALICE HENKEL, 


ASSISTANT, DRUG-PLANT INVESTIGATIONS. 


IssurED DECEMBER 28, 1905. 


WASHINGTON: 
GOVERNMENT PRINTING OFFICE. 
1905. 


CONTENTS. 


we AU ALOT 2 oe SLE ea 8 A da Ba 
aE NREL OTOW Io See ge SN oe ee ee 
Peppermint cultivation in the United States: 29555, 2 Mise See tee ee eke eptete 


CRUE 2 aE) 22 3 0 ee ee eae SEE 2h pepe > ey eed Werk eee ee 


Conditions injurious to crop____._- WET SAT Is bop A eM Pe tO oe Sah 2 
unum ishiiition ! °- |. 28 is. esse oo oe Ae losses eka 
ere aMLOTINC EMU oorrhre A ye Re Sie Se a 
PPEmminOtang menthol 92. 422 ses Sk et eh eee 
BENG Ct ES Ee Ss 
SEE eSEenibrarreibar (07 fobs 465. 82 OF oe 


Fig. 1 
9 


~ 
2 
2 


ILLUSTRATIONS. 


. Peppermint ‘‘runners,’’ showing method of propagation 
. Leaves and flowering top of peppermint... -_...---- .----------2-=- 


. Peppermint still (after Dewey, in Bailey’s Cyclopedia of American 
Horticulture) 


4 


B. P. I—189. 


Ee EGE VENT 


DESCRIPTION. 


One of the most important essential oils produced in the United 
States is distilled from the peppermint plant and its varieties. The 
three kinds of mint grown in this country for the distillation of pep- 
permint oil are the so-called American mint (J/entha piperita L.), 
the black mint (Jentha piperita vulgaris Sole), and the white mint 
(Mentha piperita officinalis Sole), the two last named being varieties 
of the American mint. 

The American mint, although introduced from England many 
years ago, is so called from the fact that it has long been cultivated 
in this country, and the name “ State mint ” has been applied to it in 
the State of New York for the same reason. 

The peppermint, or American mint, is now naturalized in many 
parts of the eastern United States, occurring in wet soil from the New 
England States to Minnesota, south to Florida and Tennessee. It is 
an aromatic perennial belonging to the mint family (Menthacez), 
and propagates by means of its long, running roots (fig. 1). The 
smooth, square stems are erect and branching, from 1 to 3 feet in 
height, bearing dark-green, lance-shaped leaves, which are from 1 to 
2 inches long, and from one-half to 1 inch wide. The leaves are 
pointed at the apex, rounded or narrowed at the base, sharply 
toothed, smooth on both sides, or with hairy veins on the lower sur- 
face. The flowers are borne in whorls in dense, terminal spikes; they 
are purplish, with a tubular, five-toothed calyx, and a four-lobed 
corolla. (Fig. 2.) 


aIn response to a steady demand for information relating to the peppermint 
industry, Miss Alice Henkel, Assistant in Drug-Plant Investigations. has been 
requested to bring together the most important facts regarding the history, 
culture, and utilization of the peppermint plant. The information here pre- 
sented has been obtained in large part from scattered articles on the subject, 
and in part from experience with the plant in the Testing Gardens of the Depart- 
ment of Agriculture. 

Ropney H. True, Physiologist in Charge. 
OFFICE OF DRUG-PLANT INVESTIGATIONS, 
Washington, D. C., October 14, 1905. 


6 PEPPERMINT. 


The two varieties mentioned are closely related botanically, al- 
though in general appear- 
ance they are quite differ- 
ent. The variety known 
as black mint (J/entha 
piperita vulgaris) has pur- 
ple stems and slightly 
toothed, dark-green leaves, 
while the white mint: 
(Mentha piperita  offici- 
nalis) has green stems, 
with brighter green leaves, 


Fig. 1.—Peppermint ‘‘runners,” showing method of which are more lance- 

propagation. shaped and more deeply 
toothed. Black mint is much more hardy and productive than either 
the American mint or the white mint, 
and is grown on nearly all pepper- 
mint farms in this country. The white 
mint, which produces a fine grade 
of oil, is rarely cultivated on a com- 
mercial scale in this country on ac- 
count of its inability to withstand the 
climate and its smaller yield of essen- 
tial oil. 

The oils spoken of as Japanese and 
Chinese * peppermint ” oils are not ob- 
tained from the true peppermint plant, 
but are distilled from entirely different 
species, namely, Mentha arvensis piper- 
ascens Malinvaud and Mentha arvensis 
glabrata Holmes, respectively. 


COUNTRIES WHERE GROWN. 


The most important peppermint- 
producing countries are the United 
States, England, and Japan. Pepper- 
mint is grown on a smaller scale in 
Germany, France, Italy, Russia, China, 
and southern India. 

In Japan, peppermint cultivation 
is said to have been undertaken 
before the Christian era. The plant 


grown there is not. as already Fic. 2.—Leaves and flowering top of 
E peppermint. 


stated, the peppermint cultivated in 
our country, but Mentha arvensis piperascens, which is entirely dis- 


CULTIVATION IN THE UNITED STATES. 7 
tinct from the true peppermint, not only botanically but also in taste 
and odor. 

Peppermint is cultivated on many drug farms in England, espe- 
cially at Mitcham, the middle of the eighteenth century marking the 
beginning of peppermint cultivation in that country. Up to 1805, 
however, there were no stills at Mitcham, and the crops obtained 
there were sent to London for distillation. About 1850, at which 
time the peppermint industry in England was at its height, the effect 
of American competition began to be felt, and caused a decided 
check in the production. 


PEPPERMINT CULTIVATION IN THE UNITED STATES. 


Wayne County, N. Y., in 1816, was the first locality in this country 
to distill peppermint on a commercial scale. The supply of root- 
stocks was obtained from the wild plants found growing along the 
banks of streams and brooks. Adjacent counties soon undertook the 
cultivation of peppermint, but Wayne County was then, and is now, 
the principal peppermint district in New York. 

The cultivation of peppermint was extended to Ashtabula, Geauga, 
and Cuyahoga counties in Ohio, and also to northern Indiana. Roots 
were taken from Ohio into St. Joseph County, Mich., the first plan- 
tation being made on Pigeon prairie in 1835. Other plantations in 
St. Joseph County were established the following years, and adjoin- 
ing counties soon took up the cultivation of peppermint, and south- 
western Michigan has been for thirty-five years or more the greatest 
peppermint-producing section in the United States. 

About 1844 an interesting peppermint-oil monopoly ¢ was under- 
taken by a New York firm, which seems to have put an end to pepper- 
mint cultivation in Ohio, for none of the counties just. mentioned 
has since been heard from as a peppermint-producing section. 

The first step taken by this New York firm in its efforts to con- 
trol the peppermint-oil market was to send a representative to 
Liverpool, England, to ascertain the amount annually demanded by 
that market, which was found to be about 12,000 pounds. This done, 
another agent was sent West to determine the amount produced annu- 
ally, with the result that it was found that the farms in New York 
_did not produce enough oil for their purposes, the plantations in 
Ohio too much, while those in Michigan seemed to produce just about 
the right amount to satisfy the Liverpool demand. <A contract was 
then entered into by this agent with the producers in New York and 
Ohio “ whereby he bound them under heavy penalties to plow up 
their mint fields and destroy the roots, and not plant any more mint, 
or sell or give away any roots, or produce or sell any mint oil for the 


aProc. Amer. Pharm. Assoc., 7: 449-459 (1858). 


8 PEPPERMINT. 


period of five years.” For this wholesale destruction of their mint 
fields the producers received a bonus of $1.50 per acre. Next a con- 
tract was made by the agent with the producers of St. Joseph County, 
Mich., agreeing to pay them $2.50 a pound for their mint oil, every 
ounce of the mint oil to be delivered for a period of five years to the 
agents named in the contract. They also were prohibited during 
this period from extending their plantations and from selling roots 
to anyone. The producers held to these contracts for about three 
years, after which period the New York firm was not so anxious to 
enforce them, having, in the meantime, acquired a large fortune 
through its peppermint-oil monopoly. 

Since that period the area devoted to peppermint cultivation in 
Michigan has steadily increased, and northern Indiana, with its prin- 
cipal centers of production in St. Joseph, Steuben, and La Grange 
counties, continues to place on the market a considerable quantity of 
oil. Ohio seems to have abandoned peppermint cultivation, at least 
on a commercial scale, and New York, for a number of years and until 
very recently, had greatly reduced the area under peppermint, thou- 
sands of acres formerly devoted to this crop having been given over 
to sugar beets, onions, and celery. In 1889 Wayne County, N. Y., 
had 3,325 acres of peppermint, whereas in 1899 there were only 300 
acres. In 1905, about 933 acres were under cultivation. 

Special canvassers appointed by the State of Michigan“ made a 
canvass of 299 growers in the peppermint district in that State, cover- 
ing 39 townships in nine counties (Allegan, Berrien, Branch, Cass, 
Kalamazoo, Oakland, St. Joseph, St. Clair, and Van Buren), and 
the total number of acres under peppermint cultivation, the number 
of pounds of oil distilled, and the average number of pounds per aere, 
as ascertained by this canvass, for the years 1900, 1901, and 1902, are 
as follows: 


Items. | 1900. | 1901. 1902. 
Potal number of Acres STOW: Ll: oe ieee oes soe eee ee eee 2,112 | 2, 7824 6, 4008 
Total number of pounds distilled --..-.--.-..---.-.-.- wacencecseeeeen|- 47,6281] GBTIBS oe ees 
Average number of pounds per acre. --2-- ---2.- bie 32. seen anne 22.5 23.9 | 12.8 
CULTIVATION. 


Peppermint cultivation is most profitable on muck lands, such as 
are now used in Michigan for this crop and for celery and cranberry 
culture. These muck lands were formerly marshes and swamps, 
which have been reclaimed by draining, plowing, and cultivating, 
the swamp vegetation having been thus subdued, and the decayed 


«Twentieth Annual Report of the Bureau of Labor of the State of Michigan, 
1903, pp. 458-447. 


CONDITIONS INJURIOUS TO CROP. 9 


vegetable matter resulting in a very black soil which is most admir- 
ably adapted to mint cultivation. Formerly peppermint was grown 
exclusively on upland soil in Michigan, but it is a very exhausting 
crop on such land. Only two crops can be obtained from upland 
plantations, and after the second year’s harvest the land is plowed and 
a rotation of clover, corn, etc., is practiced for five years before pep- 
permint is again planted. But on the rich muck land peppermint 
can be grown year after year for six or seven years, the land being 
plowed up after each crop is harvested, and the runners turned under 
to form a new growth the succeeding year. The ground is harrowed 
in autumn and again in spring, and carefully weeded. Peppermint 
will grow, however, on any land that will produce good crops of corn, 
the ground being prepared by deep plowing and harrowing. 

In Michigan’ the land is plowed in the autumn, and early in spring 
it is harrowed and marked with furrows about 3 feet apart. The 
roots selected for planting are from one-eighth to one-quarter of an 
inch thick, and from 1 to 3 feet long; and the workmen engaged in 
“setting mint,” as the process is called, carry these roots in sacks 
across their shoulders and place them in the furrows by hand, cover- 
ing the roots with one foot and stepping on them with the other. 
The roots are planted so close together in the furrow as to form a 
continuous line. An expert workman can plant about an acre in 
a day. 

In about two weeks the young plants will make their appearance, 
and are carefully hoed and cultivated until July and August, when 
the plants have usually sent out so many runners as to make further 
cultivation difficult. The crop is cultivated with horse cultivators, 
but if the land was very weedy in the first place, the weeds will 
have to be pulled by hand. It is very necessary that the land be free 
from weeds, as any collected with the peppermint crop will seriously 
injure the quality of the oil. 

It may be interesting to note here that on muck lands, when 
necessary, the horses are usually provided with mud shoes to prevent 
their sinking into the soft, wet ground, these mud shoes consisting 
of wide pieces of iron or wood about 9 by 10 inches, fastened to the 
hoofs and ordinary shoes by means of bolts and straps. 


CONDITIONS INJURIOUS TO CROP. 


Cold and wet weather or extremely dry periods have a very unfa- 
vorable effect on the mint crop. Insect enemies also tend to cut down 
the mint harvest—grasshoppers, crickets, and cutworms sometimes 
doing considerable damage. <A rust, causing the foliage to drop off 


«Twentieth Annual Report of the Bureau of Labor of the State of Michigan, 
1903, pp. 4838-447. 


10 PEPPERMINT. 


and leaving the stems almost bare, is apt to follow if very moist 
weather occurs toward the latter part of the season. Weeds are 
especially to be avoided in a mint field, since, as stated, the quality 
of the oil will be seriously impaired if these are harvested with the 
peppermint. The weeds generally found in a peppermint field are 
Canada fleabane (Leptilon canadense), fireweed (Evrechtites hieraci- 
folia), giant ragweed (Ambrosia trifida), pennyroyal (Hedeoma 
pulegioides), Katon’s grass (Hatonia pennsylvanica), June grass 
(Poa pratensis), and other low grasses. 


HARVESTING AND DISTILLATION. 


The first crop of mint is harvested in the latter part of August, 
when the plants are in full flower, and the gathering continues until 
about the middle of September, the stills running night and day 
until all the mint is disposed of. The first crop is usually eut with a 
scythe, as mowing machines do not work well on soft cultivated land. 
The succeeding crops are cut with a mowing machine or sweep-rake 
reaper. The highest yield per acre and the best quality of oil are 
obtained from the first year’s crop. Sometimes, if the weather con- 
ditions have been very favorable, a second cutting is made. The 
yield of oil from peppermint obtained from the same field sometimes 
varies very much, the condition of the atmosphere seeming to exert 
xn influence upon it, as it 1s said that mint cut after a warm and 
humid night will yield more oil than that cut after a cool and dry 
night. It requires about 330 pounds of dried peppermint to produce 
1 pound of oil, and the yield of oil from an acre ranges from 12 to 50 
pounds. 

If the mint crop has been grown on muck land, all that is necessary 
after the crop has been harvested is to plow up the land and turn the 
runners under for a new crop. If grown on upland, after the second 
year’s crop is in, or, at the most, after the third year’s harvest, the land 
is plowed and then given up to other crops. Peppermint exhausts 
the land, and it is necessary to practice rotation of crops for about five 
years in order to put the land in condition if it is desired to use it 
again for peppermint cultivation. 

After the plants are cut they are usually placed in windrows until 
they are dried, but are not allowed to become so dry as to permit the 
leaves to shatter off, and are then taken to the distillery. Some grow- 
ers believe that if the plants are allowed to dry there will be a smaller 
oil content owing to the escape of some of the oil into the atmosphere, 
and so have the plants brought to the distillery in the green state; 
but Mr. A. M. Todd @ is of the opinion that no loss of oil will result 


aAmer. Jour. Pharm., 60: 328-332 (1888). 


DESCRIPTION OF STILL. 11 


from drying, his experiments along this line showing that the dry 
plants can be distilled three times as rapidly as the green plants, and 
that a larger quantity of oil may be obtained. He states that— 

To obtain the best results, both as to quality of essential oil and economy of 
transportation and distillation, the plants should be dried as thoroughly as pos- 
sible without endangering the loss of the leaves in handling. Distillation 
should then take place as soon as convenient to prevent the oxidation of the oil 
in the leaf by atmospheric action. 


The smaller producers, who have no stills of their own, have their 
mint crop hauled to the nearest peppermint distillery, where it is 
distilled for them at a cost of 25 cents per pound of oil. 


DESCRIPTION OF STILL. 


The apparatus used in peppermint distillation in the early years 
of the industry in this country consisted of a copper kettle, from 
the top of which a pipe connected with a condensing “ worm.” 
Water was placed in the kettle and the plants were immersed in it, 
and direct heat was applied to the bottom from a furnace. With 
such a still only about 15 pounds of oil could be obtained from a 
charge. In 1846, large wooden vats were substituted for the copper 
kettles, and the plants were distilled by steam passing through them. 
The kettle formerly used as the still was now employed to generate 
steam, a long pipe conveying the steam to the bottom of the vats. 
With this method of distillation from 75 to 100 pounds of oil could 
be obtained from a charge without much additional expense. 

A modern peppermint still (fig. 3) may be briefly described as fol- 
lows: The apparatus required consists of a boiler, a pair of large 
circular wooden vats, a condenser, and a receiver. The boiler, of 
course, is used for the generation of steam. 

Two wooden vats are used in order that they may be filled and 
emptied alternately. These vats are about 6 feet high and about 
5 feet in diameter, with tight-fitting removable covers and perfo- 
rated false bottoms. Steam pipes are led from the boiler into the 
bottom of the vats. 

The condenser consists of a series of pipes of block tin, either 
immersed in tanks of cold water or over which cold water is kept 
running, the condenser being connected with the top of the dis- 
tilling vats. The condensed steam, together with the oil, flows into 
a metallic receiver, in which the oil, being lighter than the water, 
rises to the top and can be drawn off. 

The perforated false bottoms with which the vats are supplied 
permit the passage of steam. A strong iron hoop is placed about 
this false bottom, and two pairs of stout chains, which meet at the top 


12 PEPPERMINT. 


of the vat in a pair of rings, are attached to it. After the charge 
has been distilled it is drawn from the vats by means of this arrange- 
ment. 

The plants are thrown into the vats and are closely packed by two 
or three men tramping upon them, and as the vat becomes about 
one-third full the packing is still further assisted by turning in a 
small supply of steam, which softens the plants. When the vat is 
filled the tight cover is replaced and a full head of steam turned on. 
In the largest distilleries the vats have a capacity of from 2.000 to 
3,000 pounds of dried plants each. 


J 


HR 


Hert 


ju 


Fic. 3.—Veppermint still. (After Dewey, in Bailey's Cyclopedia of American 
Horticulture. ) 


A, boiler; B, steam pipes leading to vats; C, valves for shutting off steam; D, mint 
packed in vat ready for distilling: #, mint being lowered into vat; F, tight-fitting cover 
used alternately for both vats; G, pipe from top of vat, joined at H so as to swing to 
other vat; J, perforated pipe, from which cold water drops over condensing tubes; K, 
supply pipe for cold water; WM, condensing pipes; \, outlet for condensed oil and water ; 
O and P, water and oil in separating can; R, outlet for water; S, floor of distilling room. 


Large tanks are used for storing the oil, and cans holding 20 
pounds each are employed for shipping, three of these cans being 
placed in a wooden case. 

The peppermint hay which remains after distillation is used as a 
fertilizer or is fed to stock. 


PEPPERMINT OIL AND MENTHOL. 


Peppermint leaves and flowering tops are official in the Eighth 
Decennial Revision of the United States Pharmacopoeia. as are like- 
wise the following products and preparations derived from these 
parts: Oil of peppermint, menthol, spirit of peppermint, and pepper- 
mint water. 


EXPORT OF PEPPERMINT OIL. 13 


The United States Pharmacopceia describes oil of peppermint as 
“a colorless liquid, having the characteristic strong odor of pepper- 
mint and a strongly aromatic pungent taste, followed by a sensation 
of cold when air is drawn into the mouth.” It is largely used in medi- 
cine, internally as a stimulant and carminative, and externally to 
relieve neuralgic and rheumatic conditions. It is also used for flavor- 
ing and scenting confectionery, cordials, and cosmetics. There is a 
slight difference in the odor of white and black peppermint oil, the 
black being more pungent and less agreeable in fragrance than the 
white, which has a much finer odor, but, as already indicated, the 
white mint is less hardy than the black and yields a smaller quantity 
of oil. 

The Japanese oil of peppermint, which, as pointed out elsewhere in 
these pages, is obtained from a different species of mint than that 
which produces the true oil of peppermint, is very inferior to the last 
named. It has a very unpleasant odor and a bitter, disagreeable 
taste, but it is a heavy oil and contains a higher percentage of menthol 
and, being a very much cheaper oil, it is hable to be used as an adul- 
terant of true peppermint oil. 

Menthol, formerly known as peppermint camphor, is the solid con- 
stituent of oil of peppermint, obtained by subjecting the distilled oil 
to an exceedingly low temperature by means of a freezing mixture. 
Its properties are about the same as those of oil of peppermint, only 
somewhat intensified. It is very largely made up into cones or pencils, 
which furnish a popular remedy, to be applied externally or inhaled, 
for the relief of headache, neuralgia, catarrh, asthma, and kindred 
affections. It is also largely employed in other forms of medication. 
The name “ pipmenthol” has been applied to the menthol obtained 
from the American oil, to distinguish it from the Japanese menthol. 
Pipmenthol is said to have a distinct odor of peppermint, while the 
Japanese menthol has but a slight peppermint odor. 


EXPORT OF PEPPERMINT OIL. 


The exports of peppermint oil during the fiscal year ended June 30, 
1904, amounted to 42,939 pounds, valued at $124,728. Germany and 
_the United Kingdom were the largest consumers, the former receiving 
22,372 pounds, valued at $65,505, and the latter 11,558 pounds, worth 
$31,798. 

The following tables show the export of peppermint oil, by coun- 
tries, for the fiscal year ended June 30, 1904, and the quantities and 
values of peppermint oil exported for a period of ten years, from 
July 1, 1894, to June 30, 1904, inclusive: 


i4 PEPPERMINT. 


Haeports of peppermint oil, by countries, for the fiscal year ended June 30, 1904.4 


Quan- 


Country. tity. Value. 
: Pounds, 
DG 6 92 2 Ssh ee eee ee See an ee See ee 473 $1,585 
BYRNGR oe 8 2 3 ee ee es ee eek wee Be ee ie 3, 054 10,059 
Germany 225 'us £5 Oe ees ee ee eee bt Ass ae Si eee 22,372 65, 505 
Netherlands =. ...-<05 toe ee. 22 ee eee 590 1,934 
United Kingdom: 4.289220.) 5.25 2b ee Se 11,558 | 31,798 
Dominion of Canada: - 
Nova’ Scotia, New Brunswick, ete.2- 220532222 2 See &5 234 
Quebec, Ontario; Manitoba, étce-22 222 2-2 ae eee 1,165 3, 306 
Noewtoundland andiiabrador: --.5.5-2. 2 ee cede.) eee. ee 94 204 
West Indies: 

British. 2 ccs! 2. ce ete Sasa got = eben 183 7 
Ours 3. eee a Le Se ee ce ee eee 87 
Danigh- o. be. -25 eee. eo eee eed ee 17 55 
When 2A ee foes 2 ch eee ee a ee Ce 20 61 
JAVPONTINA: & Acc Seeo wo ech ce tombe ee se eee ee a ee 1, 237 3, 504 
British G@uiane-22332. ci. ee te eee ae eee 10 | 81 
Porn) he eRe eee eee Lcdalaldotcad Minne SOUeE en 50 | 175 
British Anstralasia 2. 0222.22-0: 206s. cdo asd dd 1,176| 3,019 

Total... Gsshes cose ee eS eee 42,939 | 124, 7: 


“The Foreign Commerce and Navigation of the United States for the year ending 
June 30, 1904, vol. 1, p. 531, Bureau of Statistics, Department of Commerce and Labor. 


Quantities and’ values of peppermint oil exported during the fiseal years 1895 
to 1904, inclusive.@ 


Fiscal year. Sey Value. | Fiscal year. ph | Value. 
| 
Pounds. | Pounds. | 
i OE a ls 87,633 | $194,616 || 1900 - --------2------| 89,558 | $90, 298 
1806 ok Sith Seton gerey 2 acais $5, 200'|  174,810,,|| 1901 22.5525 3 ae ee 60,166 | 63,672 
ts 0 Gene DO ad RET 162,492:| 957,484 || 1902... 28 i ce.0.ce eee | 36,301 | 54,898 
(PEI BRSE A 6) Ok gee OIE 145, 975:| 180/810 || 1908- 28.) 2) eee ee 13.033 | 34,943 
rT Ce OSS at ar, ge ae 137,462 | 118,227 || 1908 _-o oe: 2. es ee | 42.930 124,728 


«From The Foreign Commerce and Navigation of the United States for the year ending 
June 30, 1902, vol. 2, p. 309, Bureau of Statistics, Treasury Department; and The 
Foreign Commerce and Navigation of the United States for the year ending June 30, 
1904, vol. 1, p. 192, Bureau of Statistics, Department of Commerce and Labor. 


PRICES OF PEPPERMINT OIL. 


The price of peppermint oil was very low for a few years prior to 
1900, the enormous production of 1897 resulting in a great drop in 
price. The lowest price paid for it was in 1899, when it brought 
only 75 cents per pound. As a result of the low price a great many 
mint farmers restricted the area of their mint plantations or alto- 
gether abandoned peppermint cultivation. The smaller output of . 
the following seasons again sent prices up, and in 1902 the oil sold 
as high as $4.75 a pound, which price was maintained until early in 
1903, when it gradually declined, until toward the end of that year it 
reached $2.20 per pound. 


PRICES OF PEPPERMINT OIL. 15 


The following table* gives the highest and lowest prices of pepper- 
mint oil in bulk from 1873 to September 16, 1905: 


| | | 
Year. Highest.| Lowest. Year. Highest.| Lowest.) Year. Highest. Lowest. 
| 
$3.15 3 Plata |abcy eee $3. 00 $2. $2. 00 $1.70 
5.25 oy fl | lel koalas ere Be 4, 37 2. 1.85 | 1.20 
5.50 3. ; 3. 60 2. 1.25 | 90 
3.75 2. 2.75 at: 90 80 
3.00 i 2.40 le 90 | 75 
2. 00 ale 2.30 i 1.10 | 80 
2.65 ap 2.40 al6 1.80 | 1.10 
2.87 2. 2.50 | 2. 4.75 | 1.7 
2.85 2.¢ 2.50 | 2.15 4.75 2.20 
2.50 i: 2.45 | 2. 3.75 | 2.65 
2.60 2 2.45 UF 3.45 2. 25 


* To September 16. 


The good prices of the past few years have caused many farmers 
to look again to peppermint as a profitable crop, as noted in increased 
areas under cultivation in many localities. This is the case not only 
in Michigan and Indiana, but also in New York, where for many 
years the peppermint industry has been declining. Thus, if favor- 
able conditions of growth prevail, an increased production may be 
looked for within the next few years, which will have the effect of 
again depressing prices. 

As is the case with other products the prices of which are subject 
to great fluctuations, the condition of the market for peppermint oil 
needs to be closely observed. The cost of cultivation per acre has 
been stated at from $12 to $14, and, with a charge of 25 cents per 
pound of oil for distillation, the market price may easily fall below 
the cost of production. 


@¥rom Oil, Paint, and Drug Reporter, September 18, 1905, p. 7. 


O 


U. S. DEPARTMENT OF AGRICULTURE, 
BUREAU OF PLANT INDUSTRY—BULLETIN NO. go, PART IV. 


B. T. GALLOWAY, Ohief of Bureau. 


ALBERT C. CRAWFORD, 


PHARMACOLOGIST, POISONOUS PLANT INVESTIGATIONS. 


ISSUED JANUARY 17, 1906. 


‘es ae we 3 ao 
Ve = 


a SES 
bb > e 


WASHINGTON: 
GOVERNMENT PRINTING OFFICE. 
1906. 


B. P. 1.—194. 


THE POISONOUS ACTION OF JOHNSON GRASS: 


' Johnson grass, which was introduced from Turkey into this country 
about 1830,? has spread so that in many places it is considered as a 
weed and pest.* Some farmers, however, have utilized the dried grass 
as hay with advantage, either alone or combined with other food ma- 
terial,t and chemical analyses have proved its value as feed. Recently 
reports have come to this office from California of the death of cattle 
under such circumstances as to point to Johnson grass as the causative 
agent—the cattle dying in thirty minutes after eating the grass. John- 
son grass belongs to the same genus of the Gramineae as sorghum. 
This group has been partially investigated chemically, and it has been 
found that the fresh green plants of various members yield hydrocyanic 


1 This office has from time to time received communications from stockmen, 
especially in the lower part of California, Arizona, and adjacent territory, ex- 
pressing a suspicion that the eating of Johnson grass had caused the death of 
stock with rather sudden and violent symptoms. There has seemed to be little 
ground in poisonous-plant literature to support such an explanation. Last sum- 
mer, however, convincing observations were reported from California by a stock- 
man who had lost heavily, and a supply of the grass in question was obtained. 
The result of the study of this material was so positive, and the possibility of 
damage due to this unsuspected forage plant so clear, that this preliminary notice 
is put out in the hope of getting observations and material for study from many 
sources, in order, if possible, to determine the conditions under which the 
poisonous properties are developed and over how wide an area they are likely 


to appear. 
RODNEY H. TRUE, Physiologist. 


OFFICE OF POISONOUS PLANT INVESTIGATIONS, 
Washington, D. C., December 11, 1905. 


2 Ball, C. R. Johnson Grass. Bul. No. 11, Bureau of Plant Industry, U.S. 
Dept. of Agriculture, 1902. 
‘Spillman, W. J. Extermination of Johnson Grass. Bul. No. 72, Part III, 
Bureau of Plant Industry, U. 8. Dept. of Agriculture, 1905. 
4North Carolina Agr. Expt. Sta. Bul. 97, p. 92. 
Vasey, G. Grasses of the South. Bul. No.3, Division of Botany, U.S. Dept. 
of Agriculture, 1887. 
Report of the Commissioner of Agriculture for 1881, pp. 231, 232, 239, 241; 


Report of the Secretary of Agriculture, 1890, p. 381. 
3 


4 


acid as a result of the action of enzymes on more highly complex 
bodies.? 

Ball? in 1902 stated that at that time there had been no official re- 
ports to his office of cases of poisoning by Johnson grass, but that there 
were some newspaper statements to that effect. He thought these 
accounts were probably not authentic, but stated that “since Johnson 
grass is closely related to sorghum, which is known to be poisonous 
under some circumstances, it would not be surprising if Johnson grass 
should also be poisonous under like conditions. * * * In eompari- 
son with the great number of cattle fed or pastured in Johnson grass, 
the reported cases of poisoning are extremely rare.”’ 

The first report of the poisonous action of Johnson grass which 
reached the Department came from Miles City, Mont. Mr. William 
Story reported that he and a neighbor had lost several head of cattle 
after they had eaten small quantities of the grass, and that they had 
died very suddenly. Mr. Story suggested that there was “something 
peculiarly poisonous about the grass.’> The Commissioner of Agricul- 
ture in publishing this report stated that “although the grass has been 
cultivated in the South for forty or fifty years, no similar charges have 
been made against it.’’? 

In India this plant is widely used as a fodder for cattle,t and the 
natives make use of the seeds for food. It has been noted there that 
deaths in cattle frequently occur when, on account of the failure of rain, 
the plants which have reached a certain size become stunted and 
withered. The toxic principle appears simultaneously over a wide area, 
but soon disappears if a rainfall occurs.® The deaths of cattle have 
been attributed by some to an insect living upon the plant, and in 
Australia it is the belief that Sorghum vulgare, which also yields hydro- 
cyanic acid, becomes more poisonous when attacked by an insect dur- 
ing a drought. A similar observation has been made with Sorghum 
vulgare in the Sudan. Balfour® found that one specimen of the plant 


Dunstan, W. R., and Henry, T. A. The Nature and Origin of the Poison of 
Lotus Arabicus. Phil. Trans. Roy. Soc. London, 1901, vol. 194, B., p. 515. 
Dunstan, W. R., and Henry, T. A. Cyanogenesis in Plants. Phil. Trans. 
Roy. Soc. London, 1902, vol. 199, A., p. 399. 
Slade, Henry B. Prussie Acid in Sorghum. Jour. Amer. Chem. Soc., 1903 
vol. 25, pp. 55-59. 
Slade, Henry B. Study of the Enzymes of Green Sorghum. Fifteenth Ann. 
Report, Agr. Expt. Sta. of Nebraska, 1902, pp. 55-62. 
Briinnich, J. C. Hydrocyanie Acid in Fodder-plants. Jour. Chem. Soe., 
1903, vol. 83, part 2, pp. 788-796. 
2Loc. cit., p. 23. 
* Report of the Commissioner of Agriculture, 1885, p. 74. 
‘Duthie, J. F. Fodder Grasses of Northern India, 1888, p. 41. 
*Pease, H.T. Poisoning of Cattle by Andropogon Sorghum. Jour, Compar. 
Med. and Vet. Arch., vol. 18, 1897, p. 679. See also Agr. Ledger, 1896, No. 24. 
* Balfour, Andrew, Cyanogenesis in Sorghum Vulgare. First Report, Well- 
come Research Laboratory, at Gordon Mem, College, Khartum, 1904, p. 47. 


On 


which harbored aphids yielded more hydrocyanic acid than a second one 
without parasites. Pease has lately claimed that the deaths from John- 
son grass in India were really cases of nitrate poisoning, as he found 25 
per cent of nitrate of potassium in the stem of the plant and was able to 
produce somewhat similar symptoms in animals by feeding them this 
salt. Johnson grass is being introduced into Australia as a fodder plant, 
but as yet no reports of its poisonous action there have been noted by 
the writer.! 

There has been some chemical study of Johnson grass, but not with 
reference to any poisonous principle.” 

A fresh, green, mature, nonflowering specimen of Johnson grass, moist- 
ened with a little water and preserved with chloroform, was sent from 
Santa Rosa, Cal., in sealed glass vessels, to this laboratory. This was 
botanically identified here as Johnson grass. This specimen was not 
immediately worked up, but remained in the jars for about a month. 
At that time on opening the jars a marked odor of hydrocyanic acid, 
together with that of chloroform, was detected. The ground-up plant, 
with the water in which it came, was distilled, and the distillate was 
caught in sodium hydrate solution. This distillate, on mixing with 
ferrous sulphate and acidulation with hydrochloric acid, gave a heavy 
blue precipitate with ferric chlorid. Yellow ammonium sulphid was 
added to the same filtrate, and the mixture was evaporated to dryness 
on the bath. The dried residue was then taken in hydrochloric acid 
water, and on the addition of ferric chlorid the fluid gave the charac- 
teristic red reaction for hydrocyanic acid. The nitro-prussid, picric 
acid, and silver nitrate reactions were all positive for hydrocyanic acid. 
The aqueous fluid in which the plant was shipped was filtered off from 
the plant and gave on distillation all the above reactions for hydro- 
eyanic acid. 

According to our California correspondent, this plant is poisonous 
when grown on irrigated as well as on nonirrigated lands, but especially 
so when grown on irrigated soil and the growth has become rank. 

Recently Dunstan* has shown that Lima beans (Phaseolus lunatus) , 
which when grown wild in Mauritius yield sufficient hydrocyanic acid to 
produce poisoning, when cultivated in Burma lose this toxicity almost 
entirely, although it may return most unexpectedly.* He was unable, 
however, to determine the condition which increased its poisonous 
properties. 


' Maiden, J. H. Useful Australian Plants. Dept. Agr. New South Wales, 
Misc. Pub. No. 22, 1896. 
* Annual Report of the Commissioner of Agriculture, 1878, p. 168. 
* Dunstan, W. R. Phaseolus Lunatus. Agr. Ledger, 1905, No. 2. 
*Chureh, A. H. Food-Grains of India. 1886, p. 155. 
Watt, George. Dictionary of the Economic Products of India, vol. 6, part 1, 
1892, p. 187. 


6 


It is interesting to note, besides this production of hydrocyanie acid 
from complex glucosids, that proteids, when subjected to oxidation 
under certain conditions, also yield it.1 In fact, hydrocyanic acid may 
exist in plants in two forms, either as the acid or as one of its salts, or 
in the form of complex glucosids.2 Under the circumstances, the 
conclusions of Briinnich® should be held in mind, viz, that “‘ all fodder 
plants related to sorghum must be used with discretion in either the 
green or the dried state, and should not be given in large amounts to 
animals which have fasted for some time.”’ 

In reference to other forage plants, Avery? says that “ Kafir-corn 
leaves also contain this poison, but other forage plants—clover, alfalfa, 
grasses, and corn—give no test for prussic acid,’ and Briinnich also 
found it in Guinea grass or Panicum maximum and P.muticum. Many 
facts have been collected relative to the distribution of hydroeyanic acid 
in plants, yet its exact significance in their metabolism is unknown.” 
The question as to the relationship of parasites® to the production of 
hydrocyanic acid remains to be solved. 

Later investigations will be carried on to determine the nature of this 
cyanogenetic compound, to determine whether hydrocyanic acid is 
present in all stages of its growth, but disappears on drying the plant, 
whether the hydrocyanic acid production occurs under all conditions or 
only when grown on certain soils, and the amount produced. Hydro- 
cy anie acid will also be looked for in other member of this genus. 


eitieriten Re ae YY The Formation of Prasat Abia by the ‘Oxaucn of 
Albumins. Jour. Physiol., vol. 31, 1904, p. 65; vol. 32, 1904, p. 50. 

2 Les Nouveaux Remédes, vol. 14, 1898, p. 272. 

$ Loc. cit., p. 792. 

4 Avery, S. Laboratory Notes on Poison in Sorghum, Jour. Compar. Med. 
and Vet. Arch., vol. 23, 1902, p. 705. 

5Czapek, F. Biochemie d. Pflanzen, 1905, vol. 2, p. 259. 

‘ Literature on some parasites of the sorghum pate can be found in Bot, Gaz., 
vol. 28, 1899, p. 65. Also in Busse, W., Untersuch, u. d. Krank. der Sorghum 
Hirse, Arb. a. d. biol. Abtheil. f. Land u. Forstw. am kaiserl. Gesundheitsamt, 
1904, vol. 4. 


O 


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