IN THE EVENING.
MISSOURI
BOTANICAL GARDEN.
SEVENTEENTH ANNUAL REPORT.
ST. LOUIS, MO.:
PUBLISHED BY THE BOARD OF TRUSTEES.
1906.
BOARD OF TRUSTEES OF THE MISSOURI
BOTANICAL GARDEN.
EpwarpD OC, ELIOT.
JOHN GREEN, M. D.
GEORGE C. HITCHCOCK.
LEONARD MATTHEWS.
WILLIAM H. H. PETTUS.
JOHN F. SHEPLEY.
Davin S. H. SMITH.
EDWARDS WHITAKER.
«
President,
RUFUS J. LACKLAND.
Vice-President,
DAVID F. KAIME.
ADOLF ALT, M.D.,
President of The Academy of Science
of St. Louis.*
WINFIELD S. CHAPLIN,
Chancellor of Washington Univer-
sity.*
GEORGE R. LocKwoop,! 3
President of the Board of Public
Schools of St. Louis.*
DANIEL S. TUTTLE,
Bishop of the Diocese of Missouri.*
ROLLA WELLS,
Mayor of the City of St. Louis.*
A. D. CUNNINGHAM, Secretary.
*Ex- Officio.
1 Elected President of the Board of Public Schools of St. Louis, October 9, 1906,
to succeed Robert Moore, who had held that office for one year. :
(2)
PREFACE.
Under direction of the Board of Trustees, the seven-
teenth annual report of the Missouri Botanical Garden is
presented to the public.
The sixteenth volume was issued on May 31st, 1905,
which is to be regarded as the date of publication of the
scientific papers it contains, except that by Professor
Hitchcock, separates of which were distributed on March
29th, 1905.
These reports are sent to scientific institutions and
journals in exchange for publications or specimens desir-
able for the Garden, and, when possible, reprints of the
botanical articles they contain are presented to botanists
occupied with a study of the subjects they refer to. Any
of the Garden publications not out of print may be pur-
chased at approximately the cost of publication from Mess.
R. Friedlinder & Sohn, Berlin, Germany; W. Wesley &
Son, London, England, or the undersigned.
Witiram TRELEASE.
St. Louis, Mo., October 10, 1906.
(3)
CONTENTS.
PaGr.
1. REPORTS FOR THE YEAR 1905:—
a. Report of the Officersofthe Board . ....... #7
b. Seventeenth Annual Report of the Director . . . . . 28
2. ScrentTIFIc Papers :—
a. Studies on the lignin and cellulose of wood.—
ee eae ee Oa 41
b. Studies upon some chromogenic fungi which Mickhen
wood.—
By George Grant Hedgcock . .. . a 59
-¢. Zonation in artificial cultures of dics had otheottien and
other fungi.—
By George Grant fo Oe ee an 115
d. Some new Texas plants.—
By B.D ae 119
e. Ascidia in Gasteria and Agave.—
By J. Arthur Harri = Sekt : 126
J. Prolification of the fruit in Capsicum ap Passiflora.—
By J. drthus Fes Ce Ort. ee oe ee
_g. Fasciation in Oxalis crenata and experimental produc-
tion of fasciations.—
: By Henri Hus Se ek es oe A Pee eb boste Bene 2 4
h. Constriction of twigs by the bag worm and ichiitans evi-
dence of growth pressure.—
By Hermann ven Schrenk . . 2... 1: , 153
AS
(5)
LIST OF ILLUSTRATIONS..
Frontispiece: In the evening.
Miepg the DrOOK =~. - 6s et the
Without the walls .
Improvement plans . +--+ +s * * * * * ° ;
A tangle of Ee Or a ee oe ed
Se a ee eae ae Sean hn | A ea
In the North American garden . ++ +
Plates 1-2 Si a RE Tree ee 5 |g a aS
Wee Sale... oy 5. eae et
Piates 10-16"... 0 eH
Plates 17-19 7 . . . . . . . . . . . ® . .
Plates 20-26, Diagrams 1, 2 ee ee eT a
(6)
. Facing p.
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x.
BROOI
ALONG THE
REPORTS FOR THE YEAR 1905.
REPORT OF THE OFFICERS OF THE BOARD.
SUBMITTED TO THE TRUSTEES JANUARY 10, 1906.
To the Board of Trustees of the Missouri Botanical Garden:
This report marks the close of the sixteenth calendar
year of the Board’s administration of the Missouri Bo-
tanical Garden and of the revenue property devised for
its support.
PURPOSES.
The General Assembly, by an act approved March 14,
1859, authorized and empowered Mr. Henry Shaw to con-
vey certain real estate and personal property to Trustees,
for the establishment of a perpetual fund for the support
of the Missouri Botanical Garden, to be maintained for
all time for the use and enjoyment of the public, and as a
means of education and research in botany, horticulture
and allied sciences.* Fuller details of Mr. Shaw’s pur-
poses are contained in his will.f
TRUSTEES.
Mr. Shaw died on the 25th of August, 1889, and his
will was admitted to probate on the second of September
of thesame year. As Trustees, he designated «* M. Dwight
Collier, Henry Hitchcock, Wm. H. H. Pettus, Dr. John B.
Johnson, Adolphus Meier, Wm. G. Eliot who is now Chan-
cellor of the Washington University, and his successor in
office, Charles F. Robertson, who is now Bishop of the
* Rept. Mo. Bot. Gard. 1: 26.
+ Rept. Mo. Bot. Gard. 1329.
(7)
ee eee ee aN Se
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8 MISSOURI BOTANICAL GARDEN.
Episcopal Church of the diocese of Missouri and his suc-
cessors, David F. Kaime, James Yeatman, Judge Saml.
Treat, Joseph W. Branch, Gerard B. Allen, Rufus Lack-
land, Judge G, A. Madill, the President for the time being
of the Public Schools, and -his successors in office, the
President for the time being of the Academy of Science of
St. Louis, and his successors, the Mayor of the City of St.
Louis and his successors in office, Dr. Asa Gray of Cam-
bridge, Massts., and Profr. Spencer F. Baird, Secretary of
the Smithsonian Institution, Washington City. The two
last named as honorary Trustees are added to the trust in
recognition of their scientific eminence and ability.”’
Both of the honorary members, as well as the Chancel-
lor of the University and the Bishop of Missouri, had died
before the composition of the Board was made known by
the publication of Mr. Shaw’s will, as had Mr. Allen and
Mr. Meier of the personally designated members. By
direction of the courts the membership of the Board was
therefore fixed at fifteen, comprising the ten then living
designated members and five ex officio. Organization was
promptly effected by a majority of the members thus con-
stituting the Board, Mr. Lackland being chosen President —
to which office he has been re-elected each year, and Mr.
Pettus acting as Secretary for a short time until provision
was made for the regular employment of a Secretary,
when Mr. A. D. Cunningham was appointed to that office —
which he has since held.
The Board were promptly put in control of the Garden
and revenue property, and assumed their responsibility as
Trustees from the nominal date of September 1, 1889.
One of their first acts was to secure care for the Garden
by the appointment of a managing Director for it, Mr.
Shaw’s selection of Professor William Trelease of Wash-
ington University being confirmed and the duties of the
Director being defined as ‘the duties prescribed for that
office in the last will of Henry Shaw, deceased, and such
REPORT OF THE OFFICERS OF THE BOARD. 9
other duties as may from time to time be prescribed by
this Board in pursuance of the trusts declared in said
will.”’
Of the original members of the Board appointed as
individuals, only three remain in office, Mr. Lackland,
Mr. Kaime and Mr. Pettus. Mr. Collier resigned in 1889
because of removal from the city. Judge Treat, because
of his advanced age, retired in 1890. Mr. Yeatman and
Judge Madill died in 1901. Mr. Hitchcock, who from
the first had been Vice-President of the Board and Chair-
man of its Garden Committee, died in 1902, and Dr. John-
son and Mr. Branch died in 1903. Vacancies were filled
for atime by Dr. George J. Engelmann, — elected in
1889 to succeed Mr. Collier, and resigned because of
removal from the city in 1895, and Mr. George 8. Drake, —
elected in 1890 to succeed Judge Treat, and resigned in
1895. Membership in the Board is now held further by
Mr. Leonard Matthews, — elected in 1895 to succeed Mr.
Drake, Dr. John Green, — elected in 1896 to succeed Dr.
Engelmann, Mr. John F. Shepley, — elected in 1901 to
succeed Mr. Yeatman, Mr. Edwards Whitaker, — elected
in 1902 to succeed Judge Madill, Dr. D. S. H. Smith, —
elected in 1902 to succeed Mr. Hitchcock, Mr. George C.
Hitchcock, — elected in 1903 to succeed Mr. Branch, and
Mr. Edward C. Eliot, — elected in 1903 to succeed Dr.
Johnson.
Bishop Tuttle, who had succeeded the late Bishop
Robertson before the organization of the Board, is the
only ex officio member who has met with it continuously
from the first. Mr. Chaplin, who became Chancellor of
Washington University in 1891, has served since that time
as a member of the Board. A number of well known
gentlemen have further found place on it, from time to
time, while serving as Mayor of the City, President of
the Academy of Science or President of the School Board.
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10 MISSOURI BOTANICAL GARDEN.
INCOME, TAXES.
When the Trustees were charged with their duties the
revenue property was appraised at $1,241,274.00,—
much of it being unimproved real estate, yielding little if
any revenue and burdened by taxes which year by year
have become more oppressive. By gradual increments the
gross rents have grown 324%, from $91,258.32 in 1890
(when the general taxes were $22,470.53) to $120,742.94
in 1905, the general taxes meantime having increased to
$36,397.57, or 62%,—nearly double the percentage in-
crease in rents.
Special street and sewer taxes average $10,075.75 per
year for the entire sixteen years, in addition to unusual
large expenses for sewer and street improvements, averag-
ing $47,360.05 for each of the last two years. The average
annual cost of insuring the revenue property and keeping
it in suitable repair has amounted to $12,940.94. Com-
missions and necessary legal and other occasional expenses
have averaged $2,336.75, and the office expenses of the
Board average $5,000.00, per year, — leaving an average
net income, nominally available for the purposes of the
Garden, of $52,261.85. Out of this, however, no less
than $137,384.22, an average of $8,586.10 per year, has of
necessity been reserved and has been ultimately entirely
spent for street and other purposes apart from the Garden
itself, to which the remainder, except for special bequests,
has been devoted for improvement or maintenance.
TESTAMENTARY CHARGES.
The testamentary provisions of the founder of the
Garden constituting fixed charges on the revenue are: (1)
Maintaining a net income of $3,500.00 from property
deeded during his lifetime to Washington University for
the support of the School of Botany; this has resulted in
WITHOUT THE WALLS.
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*
REPORT OF THE OFFICERS OF THE BOARD. 11
an average payment of $1,272.60 per year. (2) The sum
of $1,000.00 per year for a banquet to the Trustees of the
Garden and invited guests, the average expenditure for the
past sixteen years being $948.56. (3) For a banquet to
the gardeners of the institution and invited florists, nur-
serymen and market gardeners, $400.00 per year, of which
an average of $368.15 has been used. (4) For premiums
or prizes to be offered at a flower show held in St. Louis,
$500.00 per year, the expenditure averaging $360.75. (95)
For a sermon to be preached each year on the wisdom and
goodness of God as shown in the growth of flowers, fruits
and other products of the vegetable kingdom, $200.00
per year, which has been paid in full.
REAL ESTATE IMPROVEMENTS.
Among the real estate improvements made by the Board,
in addition to sharing in general street and sewer develop-
ment, is to be noted particularly the improvement of Flora
Avenue, by which the main gate of the Garden is ap-
proached from Grand Avenue. This street, nearly a mile
long, with the co-operation of other persons owning prop-
erty along it, has been converted from a narrow dirt road
into a wide macadamized boulevard with granitoid walks,
shade trees, and a well planted park strip down the cen-
ter, the expenditure of the Board on it amounting to
$44,840.78.
UNPRODUCTIVE PROPERTY.
It has been found impracticable to carry out Mr. Shaw’s
plan of long term residence leases for the unimproved real
estate in the vicinity of the Garden, and the courts have
granted to the Board power to sell this property, on con-
dition that the proceeds shall be reinvested so as to yield an
income for the maintenance of the Garden. Under this
permission some sales of building sites have been made and
as 7 aed ae 2 ae Be a’ i ee ae ke - thie Sid | +
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12 MISSOURI BOTANICAL GARDEN.
the Board hope gradually to sell all of the better class of
this property at a fair valuation; but much of the land is so
situated with reference to railroads and the factories, etc.
that have clustered along them as to have little prospective
value for residences, while, as has been sdid, the taxes
on it are becoming heavier each year. The Board are
therefore being forced to the conclusion that this part of
the endowment real estate must sooner or later be parted
with as a whole, and the proceeds reinvested in property |
of a better description, capable of yielding revenue. Upon
this conversion of the entire endowment into revenue-pro-
ducing property obviously rests the sole hope of the Board
, to meet the Director’s wishes— which they share—for a
much needed addition to the buildings used for the Garden
office, library, herbarium and laboratories, for additional
and larger plant houses to relieve the overcrowding of the
present collections so that they may be better displayed
ie and suitably enlarged, for further extension of the grounds
according to plans already prepared, and for the development
of the institution into the great and productive research —
pet center planned by the Director in accordance with the wish
of Mr. Shaw, — the realization of all of which, however,
is believed to be only a question of time.
THE PARK STRIP,
When Mr. Shaw presented land to the City for the
4 establishment of Tower Grove Park, he deeded with it a
strip 200 feet in width entirely surrounding this park, ex-
cept for the necessary entrance ways, with the provision,
agreed to by the City, ‘*that the Board of Commissioners
of Tower Grove Park shall from time to time cause to be
leased the said strip of land of 200 feet in width surround-
ing said park, in convenient lots not to exceed 200 feet in
front, nor less than 100 feet in front, to any one person
for periods of thirty years before renewal, for the purpose
REPORT OF THE OFFICERS OF THE BOARD. 13
of erecting villa residences thereon only; and all the gross
rents received from said leases, without deduction, shall be
forever paid over to said Henry Shaw and to his heirs,
executors, administrators and assigns, so that he and they
shall forever enjoy said rents; and said city shall execute
the proper leases therefor, which shall contain a clause
that there shall be only one residence on each tract so
leased.’’ The purpose was distinctly stated to be to make
of the 200 foot strip about the park not only a source of
ornament to the park but a source of revenue for the
maintenance of the Garden. A small portion of this strip,
adjoining the north gate of the park, was subsequently
released by Mr. Shaw to the City asa building site for a
residence erected for the superintendent of the park. When
the Board assumed charge of the Garden property it was
found that a few houses for park employees had been
erected, at Mr. Shaw’s expense, on the strip, andthe rental
from these was turned over to the Board; but these un-
sightly buildings have all been removed by direction of the
Park Commissioners and, with the exception of the small
temporary rents referred to, no revenue has yet been se-
cured from this portion of the revenue endowment of the
Garden. The Trustees should asearly as possible take steps
either to derive such income as may be secured by having
the park strip used in the manner described by Mr. Shaw
in his deed of gift, or to induce the City of St. Louis to
lease or purchase by condemnation the entire strip sur-
rounding Tower Grove Park, and thus enable the Board
to realize from it an income for the maintenance of the
Garden.
GARDEN IMPROVEMENTS.
At the Garden itself, the Board are pleased to note con-
stant and great though necessarily expensive change for
the better, and its valuation, with permanent improve-
ments, has nearly doubled, having increased from the
14 MISSOURI BOTANICAL GARDEN.
original appraisal of $125,160.00 to $227,185.94. When
the property came under the charge of the Trustees it was
a country home on which for some years inadequate
expenditure had been made for maintenance. The streets
about it were without sidewalks, and the walks within were
of such construction as to be impassable in wet frosty
weather. The surrounding walls were crumbling, the
plant houses were limited in capacity, of antiquated and
inadequate construction, and very badly out of repair, while
the residence assigned to the Director was found by a
committee of physicians to be in a most unsanitary condi-
_tion and a little museum that had been maintained for
many years had so deteriorated under the care of house-
hold servants that the Director was advised to close it.
The provision of room for administration, library, her-
barium and research purposes also confronted the Board as
an immediate necessity, as did provision for a supply of
city water, improvement of existing temporary sewers, and
some arrangement other than open grates for heating the
residence and museum building.
On these needed improvements the Board have heen
compelled to spend large sums of money. The full length
of the Garden front on Tower Grove Avenue has been ter-
raced, provided with a granitoid sidewalk and planted to
an avenue of maples,—already becoming beautiful and
yielding in summer a pleasing shade,—while cinder paths
have been maintained along the Shaw Avenue and Magnolia
Avenue sides; the larger of these improvements costing
$3,087.56. The principal walks of the Garden have been
suitably remade; the expenditure for this and the replace-
ment of much of the brick edging amounting to about
$3,000.00. The gate-house, walls, and fences were put in
repair at a cost of $5,056.03. The area covered by plant
houses has been more than doubled, at a cost of $29,608.45 —
the new houses being of modern construction, except for
two small temporary structures, and permitting the growth
REPORT OF THE OFFICERS OF THE BOARD. 15
of plants that could not otherwise have been cultivated.
Granitoid ponds, steps, etc., needed for the replacement of
decaying old wooden structures, or tofurnish opportunities
for growing such plants as the Amazon lily, have cost
$1,931.26.
At the Director’s residence the temporarily built, un-
sanitary and discordant east wing was replaced in 1890
by a conveniently planned wing, at a cost of $19,000.00.
A -water supply, including necessary piping along Tower
Grove Avenue, supplemented by a provisional sewerage
system, necessitated the expenditure of $4,589.10.
In compliance with an express direction of Mr. Shaw’s
will for the provision of a cottage to be occupied by @
person charged with caring for the mausoleum and its sur-
roundings, a neat building has been erected which also
serves as a gate-house for the private gate at Cleveland
Avenue by which the Director’s residence is reached from
the street. The cost of this improvement was $4,538.05.
Under another provision of Mr. Shaw’s will, requiring
its ultimate reconstruction on Tower Grove Avenue in some
convenient situation in contiguity to the Garden, his
former city residence was removed in 1892 from the cor-
ner of Seventh and Locust Streets and rebuilt, mainly in
fire-proof construction, in the Garden, on Tower Grove
Avenue between Shenandoah and Botanical Avenues, the
large expenditure of $33,478.89 for this purpose being
made by direction of the court. In connection with the
renovation of the residence and the removal of this build-
ing, which for the time afforded adequate room for the
office, library and herbarium, a boiler pit was provided at
a safe distance, from which steam heat is supplied to the
office, residence, and museum buildings ; $1,204.07 having
been spent for this purpose in addition to subsequent
improvement charges.
Among the provisions in Mr. Shaw’s will, explained in
some detail in certain manuscript suggestions not made a
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16 MISSOURI BOTANICAL GARDEN.
part of that document, was one looking to the education
of a limited number of garden pupils, to be lodged conve-
niently for their work. For this parpose a small building
at the southeastern corner of the Garden, found to be in
very bad repair, was renovated as far as possible and suit-
ably furnished; the cost of this being $1,933.25.
STORM AND FIRE LOSSES.
The foregoing represent the larger items of construction
and improvement that it has been found imperatively
necessary to make. Several hail and fire losses have been
met, from time to time, amounting to $3,988.97. In 1896
the Garden suffered severely from the tornado which did
such great damage generally in the City, and the Board were
compelled to spend $4,479.36 in making such repairs as
were possible, though a complete renovation of the Garden
would have necessitated almost entirely clearing and
replanting its wooded parts.
EXTENSION. :
In connection with the tornado damage attention was
directed to the need of general plans along which the
grounds might be harmoniously and adequately improved
and enlarged. The preparation of such plans was en-
trusted to Messrs. Olmsted, Olmsted & Eliot, whose final
report has been adopted for the future development of the
property at such times and in such degree as the Board
may find expedient. The total expenditure in connection
with these plans has amounted to $5,037.75. Acting on
the advice of the landscape architects, the Board, in 1897,
purchased, at a cost of $10,000.00, a narrow triangular
strip of land containing 2} acres, needed to square the
property out to Alfred Avenue.
In accordance with the plans which have been prepared,
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IMPROVEMENT PLANS.
‘ ‘
REPORT OF THE OFFICERS OF THE BOARD. 17
the southern extension of the grounds as so enlarged,
limited by Tower Grove, Magnolia and Alfred Avenues,
has been graded, drained and supplied with water pipes, and
planted with a collection of about 1,400 species of plants,
forming a synopsis of the North American flora, which is
soon to be opened to the public. Thecost of this addition,
aside from the land purchased, has been $21,286.34, — of
which $9,045.99 was for grading, $3,116.66 for the water
supply, $2,900.00 for drains, and the remainder for plant-
ing and minor improvements.
‘GARDENING.
On the maintenance of the Garden, including ordinary
improvements, an average of $43,675.33 a year has been
spent. Of this, an average of $23,271.39 is for wages
of gardeners and laborers, repairs and supplies, fuel,
water, additions to the living plants, and other sim-
ilar gardening purposes. The annual expenditure on
the Garden office, including salaries of the Director and
Superintendent, averages $5,217.67. The Board have
acquiesced in the Director’s belief that it is impossible for
the Garden to stand still, and provision for maintenance
has therefore been on a basis permitting rather small but
continuous development from year to year.
By the addition of the North American Synopsis, the
area of the grounds has been enlarged from 44.7 to about
65 acres, or nearly one-half. In 1889 the plant houses cov-
ered 14,840 square feet of ground; they now occupy 30,740
square feet, an increase of 107%. Propagating frames have
been increased from 2,200 to 5,750 square feet, or 161%.
It has been estimated that not over 2,000 or 3,000 kinds of
plants were cultivated at the time when the Board took
charge of the Garden. In 1895, after an accurate system of
recording plant accessions had been introduced, the number
of species and varieties was inventoried at 3,921, exclusive
2
18 MISSOURI BOTANICAL GARDEN.
of, perhaps, 1,000 annuals. In 1898 a new inventory
showed the presence of 8,009 kinds; the inventory of 1903
showed that there were then 11,357, and the number now
reported is 15,976, — an increase of 431% over the larger
of the estimates for 1889. Several of the special collec-
tions of plants are among the largest of'their kind in the
world. This is especially true of the cacti, comprising 678
named species representing 17 genera, and of the agaves and
yuccas. The collections of palms and sago plants are also
unusually large. From almost nothing, the collection of
bromeliads, or plants of the pineapple family, has grown to
204 species. Orchids, of which originally there were few,
and though two-thirds of the collection were destroyed by
fire a few years since, have been recollected to the number
of 942 species representing 116 genera, and an expert dealer
in plants of this group has recently spoken of the Garden
collection of orchids as being the largest in the United
States. With this increase in the variety of living plants,
their scientific value has been constantly raised by the in-
corporation of specimens collected in their native homes
by the Director, who has visited Mexico and Central
America for this purpose, and by others, thus permitting
a definite record to be kept of their origin, — which is not
usually true of purchased plants.
One of the early provisions made by the Board was for
the adequate labeling of the plants,—a need specially
mentioned in Mr. Shaw’s will; and the entire large col-
lection is now provided with names as far as the state of
maturity of the specimens makes this possible. With the
greatly increased variety and the accurate naming of the
plants they are better cared for than formerly and
their decorative use has received greater attention. The
former promiscuous beds of flowers have given place to
well set instructive groups of attractive species. As an
example of this may be mentioned the recent solid planting
of the sunken garden in front of the gate with beds of tulips
EOE SE TPES fe ee
REPORT OF THE OFFICERS OF THE BOARD. 19
for the spring and of foliage plants in summer, producing
an unrivaled display of the best varieties of these plants.
A collection of decorative species that are hardy in this
vicinity occupies a separate section of the central part of
the grounds, and the names of these plants are frequently
noted by visitors for use in planting their own gardens.
For the convenience of teachers and classes, several
hundred instructive plants are arranged in _ botanical
sequence, and forage plants, savory herbs and medicinal
plants are similarly grown in separate groups. One of the
features of the Garden for several years past has been the
growth through the summer of chrysanthemum plants for
display in the fall. Notwithstanding the limited space
available for this purpose, the chrysanthemum show of the
Garden.has won recognition as a competitor in attractive-
ness with the florists’ show held in the city, and visitors
who have traveled largely characterized that of this year
as the finest they had ever seen — not even excepting those
of Japan.
VISITORS.
The number of visitors to the Garden varies so greatly
from year to year, especially as affected by fair or unpleas-
ant weather on the two Sunday afternoons on which the Gar-
den is opened by direction of Mr. Shaw’s will, that it is hard
to furnish comparative figures. Until the middle of 1898
only estimates were made: since then a regular count
has been kept. In 1904, the World’s Fair year, the vis-
itors numbered 316,747. The average from 1899 to 1903,
was 83,503. In 1905 there were 100,830. The visitors
on the open Sundays average a little over a fourth of the
yearly total. .
Though the number of visitors is evidently increasing,
it is a matter of regret that more of our citizens do not
avail themselves of the privileges given by Mr. Shaw’s
bequest, and that so few seem to know that, excepting
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20 MISSOURI BOTANICAL GARDEN.
Sundays and holidays, the Garden is always open to every-
body, and free of all charge.
LIBRARY.
The provision of a library and herbarium for purposes
of scientific research, in addition to the maintenance of a
beautiful and instructive garden, are among the designated
purposes of the founder of the Garden.
A small but good nucleus of a botanical and gardening ‘
library came into possession of the Trustees when they as-
sumed charge of the property. To this the special libraries
of the late Dr. George Engelmann and of the Director, and,
later, the collection of early books of the late Dr. E. L.
Sturtevant, have been added by gift. Further smaller
gifts have been received every year, and purchases have
been liberally made, a yearly average of $4,418.82 having
been spent on the likrary, including salaries, fuel, pur-
chases and binding. From somewhat less than 5,000
numbers, the library has thus been brought to a present
total of about 51,000 books and pamphlets, —a tenfold
increase. It is now justly ranked as the foremost botan-
ical library in the country, and compares favorably with
the great libraries of the Old World. Its present valuation
is $84,248.35.
HERBARIUM.
In the same manner, the Bernhardi and Riehl herbaria,
bought by Mr. Shaw and numbering about 60,000 speci-
mens, were almost immediately added to by the gift of the
invaluable Engelmann herbarium, of nearly 100,009 speci-
mens; and gifts, exchange and purchase have further
brought the number of. specimens of dried plants at the
Garden up to a present total of about 524,000, — eighty-
seven times the original number. On the herbarium, an
average of $2,531.91 has been spent yearly, for salaries,
fuel and purchases; its present valuation is $79,216.75.
REPORT OF THE OFFICERS OF THE BOARD. 21
INSTRUCTION IN GARDENING,
The requirement that instruction in gardening and hor-
ticulture should receive attention at the Garden, in addition
to the provision of a lodging house for pupils, led to the
entire renovation of the fruit orchard some years ago, at a
cost of $444.44, and two small vegetable houses have been
built for further experimental and educational use. The
annual expenditure on the gardening course averages
$930.34, |
The Director reports that of the 39 pupils thus far
enrolled, of whom 15 completed the course, ten are now
successful florists or gardeners, two have become land-
scape architects, three hold responsible park positions, two
are college horticulturists with teaching as well as practi-
cal duties, one is a surveyor, one is a government plant
experimenter, and one is a forester in the Philippine
Service.
‘
INSTRUCTION IN BOTANY.
Mr. Shaw’s provision for a close connection between the
School of Botany, which he had endowed in Washington
University, and the Garden has been of great assistance to
the undergraduate department of the University, and
through the Garden opportunities for work have been
offered to graduate students of whom five have received
the Master’s degree and six the degree of Doctor of Phi-
losophy with botany as a major study. The Board expect
to see a large increase in this utilization of the Garden
facilities commensurate with the very gratifying growth of
Washington University. Indirectly the Garden has been
of much use to the young men who have served as assist-
ants in its office, library or herbarium, or as teachers in
the school of botany, for with very few exceptions they
have gone to college, government or other positions of high
BGs ses od
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22 MISSOURI BOTANICAL GARDEN,
responsibility in botany or horticulture, for which their
service here gave excellent training.
RESEARCH AND PUBLICATION.
Repeated mention is made in Mr. Shaw’s will of his pur-
pose to have scientific research, no less than decorative
gardening and _ botanical and horticultural instruction,
included among the activities of the Garden. Details of
administration have been left necessarily to the Director in
consultation with a special Garden Committee of the Board,
but his purpose to develop the institution symmetrically
under the broad plan of its founder has always met with
the support of the Trustees, who have approved his policy
of not only allowing but expecting a part of the time of
capable employees to be spent ininvestigation. Though he
looks forward hopefully to the time when the revenue of the
Board shall permit the employment at the Garden of a
corps of highly trained and expert investigators giving a
large part of their time to such study, he has been able as
yet to devote to this work only a small part of the time
of otherwise indispensable employees, the salary apportion-
ment and incidental expenses for this purpose averaging
$1,000.83 per year, for the last 12 years. He therefore
points with pride —and the Board share his gratification —
to the fact that under existing conditions not a year passes
without the accomplishment of valuable research work at the
Garden, and that its publications win commendation for
their useful contents as well as for the attractive and con-
venient form in which they are brought out and the
liberal conditions on which they are furnished to botanical
libraries and investigators everywhere. The yearly cost of
publication, including reissues of some of the volumes from
electrotyped plates which are preserved, has averaged
$2,000.37, but the Garden’s Reports have given it a stand-
ing in the scientific world that it could have attained in no
other way, and by far the largest part of the gifts to the
A TANGLE OF MOONSEED.
ix ee Ae
REPORT OF THE OFFICERS OF THE BOARD. 23
library and herbarium — averaging $1,775.36 per year for
the past 13 years— are received in exchange from cor-
respondents to whom the Reports are sent.
EXCHANGES.
Though already planted and partly equipped, the Garden
was without scientific affiliations when its development was
entrusted to the Board. To bring it into exchange relations
with other establishments, the publication of these annual
volumes, devoted to administrative reports, scientific mono-
graphs, etc., was early decided on. Through the medium
of this publication the Garden now stands in exchange re-
lations with 859 institutions interested wholly or in part in
botany, gardening, horticulture or forestry. The exchange
of plants and seeds with other gardens has recently been
- greatly increased by the issuance of an annual exchange
seed list which is sent to the officers of such establish-
ments, who may thus readily ascertain and request such of
their desiderata as can be furnished. The first seed list
included 813 species, and the list just issued contains the
names of nearly 1,400 kinds of plants of which seeds were
saved for exchange purposes in 1905.
POLICY OF ADMINISTRATION.
Through its entire history under the management of the
Board the Garden has been treated as a public institution
inthe fullest and most liberal meaning of the term. None
of its Trustees receive any form of remuneration for their
services. It is not managed as a source of revenue. Such
of its publications as are sold are charged for at cost. Its
employees, who are always required and ready to render
any proper service in their power, are forbidden to accept
fees. The office staff devote a large part of their time to
examining into and reporting on questions put in connec-
tion with a large correspondence or asked by visitors.
opt ee. Pe cen hee
Wa cece
Pa ed Oe Pay here ars
ED et Aaah ptt oe PR PO ES
ae
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ee che z oh Tie gig ei Rca Lg) Sect Ons : Pe
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7
24 MISSOURI BOTANICAL GARDEN.
The library and herbarium are opened to any one capable
of using them, and books and specimens are freely loaned
to responsible students thus enabling an increasing number
of persons to make scientific use of the Garden facilities
each year without incurring the expense of a visit to St.
Louis. Above all, restrictions on visitors are limited to
the few simple rules necessary to preserve the property
and insure its pleasant and profitable use by all.
FINANCIAL REPORT FOR 1905.
We submit now for your consideration the financial re-
sults for the year ending December 31st, 1905.
Our rental receipts show an increase of only $521.34
over the previous year, a few vacancies in the early part of
the year interfering with an anticipated increase; but as
all properties. are now occupied we may hope for an increase
of at least $5,000.00 over last year.
We have disposed of 220 feet on Flora Boulevard for a
total sum of $17,510.00.
The Board deemed it wise to dispose of a piece of prop-
erty at the northeast corner of Seventh and Chestnut
Streets fronting 21 feet on Chestnut, for the sum of
$100,000.00, it having been under lease for 89 years at
$3,700.00 per year; we hope to invest the proceeds in
real estate yielding a greater income. , '
We have purchased a piece of property 70X70 feet on
the north side of Poplar Street between 12th and 13th
Streets, adjoining property already belonging to the Board
and improved with five small houses renting for $1,200.00
per year, for the sum of $10,000.00, for the purpose of
investment.
We have again been called upon by the City for heavy
special taxes for streets and sewers, amounting in all to
$47,151.47, distributed as follows:
REPORT OF THE OFFICERS OF THE BOARD. 25
_
Rock Spring District Sewer No.14 ....... +. § 7,309 24
Manchester Road Joint DistrictSewer . . . ». «. «© « 33,899 25
Rock Spring District Sewer No. 13 . . . »« « « » « 1,979 55
Alley’ In Clty Block Wo. 4047) 0. we oe ee 1,164 00
$ 44,352 04
Sundry streets reconstructed . . . . » « » we e 2,799 43
$ 47,151 47
These improvements were anticipated in our last report.
Other improvements of a similar character will necessitate
the following expenditures during the year 1906, and,
if paid in cash, a portion of the cost must be met by sales
of residence property: — |
Old Manchester Road Sit whe wiles fauce- 0. 0° 2 cp eee
Shaw Avenue a dae eo nec Re ae ee eae ee hne ral tie 16,000 00
Molen AYORMG 6 wet he me mae: le oy 8 4,000 00
RE BORG s+ 8) 0 ge ae ee ee 14,000 00
Vandeventer Avenue . «© 6 + + + 6 ee ee es 4,000 00
$ 58,000 00
The annual bequests provided for in Mr. Shaw’s will
have been carried out at a cost of $1,884.11, with the
exception of the Trustees’ annual banquet which was not
held.
No permanent improvements have been made at the
Garden, but a large number of additions have been made
to the herbarium, and much work has been done on the
library catalogue.
_ The following amounts have been credited to the Stock
Account: —
Library . . . . . e ° °. . . e . e . . . . . . $4,332 64
Herbarium . - - - + 6 6 + + we @ ee ew ee oe 5,805 25
RECEIPTS.
Bontsin ... ec + 0, + 0 0 85m eee en
Interest and dividends . . . . + + « « « 1,828 77
Garden pasturage above expenses, etc. . . - 403 04
Garden hand-book sales . . + + + « « « 198 25
Publication sales . . «. « + © » « #*¢ « 4 08
Total income collections .. . .j $123,177 08
ON MeL Ea ee Se NaN RR Mh, ee ee Pee MS SS ema
4 26 MISSOURI BOTANICAL GARDEN.
Ce
“iy Brought forward . . . . « « . $123,177 08
. Sales of realestate . . . . . . . « « «© $100,000 00
. sig under decree of Court . . 17,510 00
Be! Insurance for loss to buildings . . ... . 1,759 87
- Le “t BE ec ee 20 00
a Shaw School of Botany, rent ...... 2,850 00
: Bonds, stocks and certificates . . . . . . 40,000 00 162,189 87
* Total receipts .. . tee pte $285,316 95
Bs Cash on hand December 31, 1904 eee 1,638 28
% $286,955 23
DISBURSEMENTS.
Garden Account,
a= Tabor pay-rolt.. . ss.» . w+ » « QI 49880
: Students’ pay-roll ey ee ae 1,474 80
ee I eg ee a 1,683 00
ee ae a 1,839 53
: ob ee ee eo Ee a ioe bs 234 50
. mepAlIp and BUppNeS. 3. ow 6 ee sl 2,638 25
: Plants and seeds , 1,046 71 $28,205 68
Herbarium Account,
PRIBMIGR ae a a es 1,688 51
wuel. . os eer ee eee 128 54
Current expenditure oe ee ee ee eee 2,599 12 4,416 17
Library Account,
oe ee ee ee ee ee 2,893 92
BMG]. ¢ 6 0 pees gts? er besa e cots 146 68
cc; Current expeniditinne Cae ee 1,966 89 —-5,007 49
Office Account,
| ee ee eee 4,681 06
Fuel tee te Gh one eel a os 116 47
Current expenditure . . . . »« « « » « 943 52 5,741 05
Research Account,
Saigrich «.7 <4 Coma Seer Cs 774 96
Current experdttnre 463 35 1,238 31
Scholarship Account,
-
°
.
.
.
REECTCHION, ns ey ee 723 60
Gare-OF Podge 2s Gee 240 00
Poel: fo fe os) heey ee) 105 82
Current pS eae eee ee ae ee ee 101 15 1,170 57
‘Total maintenhnce . 4. 2. 6 sm 8 6 is $45,779 27
Garden Improvement,
if North American synopsis, plans. . .. . 1,479 60
Total amount expended on Garden , $47,258 87
REPORT OF THE OFFICERS OF THE BOARD. 27
Brought forward « «+ + + © + + $ 47,258 87
Publication Account,
Sixteenth annual volume ...-+ ++ + $ 2,000 00 ;
First “c he ewostns’. 3 oa 254 85 2,254 85
Property Expenses,
State, school, city and sprinkling tax . . . 86,397 57
Streets, sidewalks andsewers. . . + + + 47,151 47
Insurance Be i Gg te he ya e 7,501 56
OME oe en se ee a) 2, ES 8,625 O4
Sroyerientl ee + tt eS 1,456 06 101,131 70
Office Expenses, :
meg. ie ¢ ee ee ee 4,200 00
Ofsice rent <5 sia een 8 ee 900 00
Printing, advertising, telephone, etc. . . + 777 25 5,877 25
Bequests,
Annual Flower Sermon * .. + + + © * 200 00
Annual Flower Show . +--+ ++ «+ « « 423 00
Gardeners’ Annual Banquet ... + + + 405 30
Washington University, School of Botany . 855 81 1,884 11
Sundries,
Realestate ..- - . Wa ee ee ee ee
Bonds, stocks and certificates Cg Stee ee oe
Legalexpenses. . + + + San a 1,816 90
Repairs to buildings damaged py fire ce 1,759 87 }
Shaw School of Botany, rent, etc. . . + + 2,200 00
Commissions «© «oor 8 eee te eee 477 00 126,253 77
Total disbursements . . ATT $284,660 55
Cash balance December alist, 1905 mien 2,294 68
$286,955 23
, —<———
Respectfully submitted,
R. J. LACKLAND, President.
Attest:
A. D. CuNNINGHAM, Secretary.
SEVENTEENTH ANNUAL REPORT OF THE DIRECTOR.
SUBMITTED TO THE TRUSTEES JAN. 10, 1906.
To the Board of Trustees of the Missouri Botanical Garden:
The following report for the year 1905, on the Missouri
Botanical Garden and the School of Botany connected
therewith, is respectfully submitted in compliance with the
rules of the Board.
¢
GARDENING,
Decorative gardening, from the nature and limits of
the formal grounds devoted to this purpose, presents much
the same problems year after year, and they are neces-
sarily similarly met. Last year about 37,500 plants were
used in ornamental bedding, an increase of some 2,000 as
compared with the preceding year. The early planting of
the parterre with bulbs, begun in 1904, has been repeated,
and with equal success. Through the summer the same
tract was devoted to a collection of coleus in masses rep- |
resentative of choice varieties. These new features, and
the bulb and foliage bedding on either side of the main
entrance-way, met with general commendation.
Among the special greenhouse collections, the succu-
lents, palms, cycads, and orchids have attracted particu-
lar attention, and each of these has received material en-
largement. Out of doors, massed economic plants from
the tropics, asters, pinks and dahlias, were satisfactorily
grown in more than usual number and variety. For the
fortnight beginning with November 13, a notable feature
of the Garden was a collection of about 2,000 chrysan-
themum plants, representing 211 varieties, well grown as
standards, for massed bloom, or to single heads. These
were displayed under canvas, and many flattering com-
(28)
i
4
i
4
i
4
D WALK.
CLUDE
A SE
ar
Ped
He aaa eee
me dre Me
+ y ae ‘
a Le edy
SEVENTEENTH ANNUAL REPORT OF THE DIRECTOR. 29
ments have been heard on the exhibition, which was
viewed by about 25,000 people, — one-fourth of the visi-
tors for the entire year.
Accessions number 314, comprising 17,620 plants
or packets of seeds. Of these, 680 plants, valued at
$288.45, and 360 seed packets, valued at $22.65, were col-
lected, and 26,845 plants, valued at $1,879.15, were prop-
agated, by Garden employees; 7,745, representing 199 of
the 314 entries and valued at $645.55, were presented or
received in exchange for material or for Garden publi-
cations; and 9,875, representing 67 entries, were pur-
chased, the Secretary’s books showing an expenditure of
$1,046.71 for such purchases, including transportation,
duty or other charges. ;
The records show that 2,345 species or varieties were
added to the collection of living plants, while 576 were
lost or discarded, leaving a net gain for the year of 1,769,
and bringing the total of cultivated species up to 15,976, in
contrast with the 14,207 noted for the preceding year.*
In January the Gardenissued its first ‘exchange seed list,”’
which included 813 species or varieties, of which about 18 %
were collected from wild plants of this vicinity, —the
others being from Garden plants. The convenience of this
list for correspondents was indicated by the receipt of
greatly increased requests for seeds, so that the distribu- -
tion of exchange material has been far greater than usual ;
the total during the year amounting to 4,173 packets of
seeds, valued at $417.30, and 1,320 plants, valued at
$77.80. As in earlier years, surplus bedding plants,
together with many of those removed from the grounds at
the end of the season, including duplicates of ferns,
begonias, etc., not needed for the houses, have been given
to charities and schools. The latter distribution, of some
800 plants, was effected largely through the effort of Miss
* Rept. Mo. Bot. Gard. 163 14.
?
OS £ a ite SRT ie Eth awe Ins eee eT a —S ee a i er ae, ee ee es ee
cp hey ‘ < Fi | a
30 MISSOURI BOTANICAL GARDEN.
Mary C. McCulloch, Supervisor of Kindergartens in the
public schools of the city. About 300 surplus plants have
also been given to High Schools and Normal Schools, to be
used for purposes of instruction.
As in the preceding year, the summer of 1905 was
cool. The average mean daily temperature for the en-
tire year was 1.2° F. below the normal. The months
of January and February were much colder than usual,
necessitating a considerable increase in the expense of
maintaining the plant houses. The entire month of
March, on the other hand, was unusually warm, so that
DIAGRAM A.
proo° 8 saa
P75: . satin ~
o
. 1905 w >
z & & & = Zz = 4 & 5 fo) a
me ee eS eS ee ee
at ! |
MEAN TEMPERATURES.
operations usually performed in April were found to be
necessary in March, thus increasing the length of the
open season by about a month. The precipitation for the
year has amounted to 38.54 inches, in contrast with the
average of 37.14 inches given by the sheets issued by the
local Weather Bureau office, — from which the accompany-
ing diagrams A and B have been compiled. In the early
spring, in June and in all the cold months the precipitation
was considerably below the average; but in May, July and
August the average was a little exceeded and the Septem-
ber and October rainfall was very high. Through most of
the growing season it was rather uniformly distributed,
SEVENTEENTH ANNUAL REPORT OF THE DIRECTOR. 31
DIAGRAM B.
r—5-INs /
ee ASS . / \
pa bse \ Zee
UL
JUNE
AUG
r-SEPT
r-—-OCT=
P-NOV
—DEC
-—MAR;=
APR
—MAY
PRECIPITATION.
favoring the growth of most bedding plants but also
greatly increasing the labor necessary to keep the grounds
free from weeds. For several reasons, therefore, the cost
of gardening this year has been unusually high.
VISITORS.
The total number of visitors for the year reached
100,830, of whom 12,355 were recorded for the open
Sunday afternoon in June, and 15,208 for the open
Sunday afternoon in September, the remaining 73,267
being week-day visitors. Diagram C shows the distribution
of the latter by months. The contrasted dotted line, —
representing the average for the period covered by accu-
rate records (beginning with midsummer in 1898) with
exception of the aberrant years 1902* and 1904,f—
seems to correspond closely with the normal out-door
* Rept. Mo. Bot. Gard. 14317.
+ Rept. Mo. Bot. Gard. 16317.
|
{
|
32 MISSOURI BOTANICAL GARDEN.
DIAGRAM C,
}-25000
20000 / |
r-15000 / \
10000 2
Fe Rimes MO ca Da oe ; ‘ \
WAN. FEB. MAR. APR. MAY. JUNE. JULY. AUG. SEPT. OCT. NOV. D&G
VISITORS ON WEEK-DAYS.
activity of our citizens. The great increase in the
November visitors this season, due primarily to the chry-
santhemum exhibit already mentioned, is directly attribut-
able to the fact that the officers of the street-car company
placarded the cars in such a way as to attract public atten-
tion to the Garden. Special recognition is due to the
company for this effort, which resulted in mutual ad-
vantage. The relation of Sunday to week-day visitors,
who this year reached 27% of the total, is graphically
shown on diagram D.
@ DIAGRAM D.
60000
ee Se oe ee ae a as
3—3 S—B—B-—B-60-Bo0o 3 —— 8 —$ —8—S Ff S-$
c - - al am oan a - vd = = =- - -
me ae 14 Sy ]
bate 0000 oe %
A & 3 WY 9 i
, STUDIES ON THE LIGNIN AND CELLULOSE OF WOOD. 55
any doubt that the thickness of the healthy wall is greater
by a decided difference than is that of the cellulose fiber
walls which are left in the badly rotted tissues. Such
measurements have shown a difference of about a third of
the total thickness of the original healthy wall between the
two series. It has been stated distinctly by practically all
workers with this group of fungi that delignification is a
very common phenomenon as a result of the action of the
mycelium of these fungi, which grows in the wood and dis-
integrates it to a greater or less extent as the case may be.
That cellulose is left by many of the fungi in the last
stages of decay of the wood in which they are growing is
but a very natural phenomenon to one who is familiar with
the structure of woody cell walls. Chemical analyses have
shown that cellulose is the basis of the structure of the wall
and makes up a large percentage of it by dry weight. The
methods of analysis are, to be sure, not satisfactory for
exact quantitative results since each method gives results
varying from the others by a small margin, yet the figures
obtained may be used provided we keep in mind the fact
that they are but approximate. Czapek * has given a sum-
mary of what has been done in this connection by differ-
ent chemists. The following table shows the results ob-
tained by two of the best methods of analysis.
LANGE. SCHULZE.
Beech ...- =. . + 54,0-53.0-53.0 51.0-50.5-50.0
MPS ee ees cee ce 51.0-50.0-50.6 48,0-48.2-49.0
tie Sa 55.0-56.0-56.0 52.0-52.0-52.5
The figures indicate the percentage of cellulose found
in the woods named by the two methods of analysis and
also show the amount of variation in the results obtained.
Analyses of pine wood showed that the percentage of cel-
lulose varied from 47.5 to 53.5 per cent, and it was found
that the sap wood was richer in cellulose than was the
* Czapek. Biochemie der Pflanzen. 1: 563-564. ( 1905.)
A
i Sy ip alpen ie an Se ea
ee
oo) ier
[Eee
56 MISSOURI BOTANICAL GARDEN.
heart wood. In general it may be said that from 45 to 60
per cent of wood is cellulose. Besides all this, pure cul-
tures of several of the more common of the wood rotting
fungi have been grown upon blocks of wood, which so far
as could be determined, had no unlignified cellulose in the
cell walls, and delignification has resulted while similar
control blocks have shown no such delignification.
EFFECT OF STERILIZATION UPON WOOD BLOCKS,
When it is considered for how long a time the thin micro-
scopic sections of the various woods were submitted to boiling
at extreme temperatures before any decided effect could be
detected one can hardly say that the effect of boiling at 100
degrees Centigrade for the time necessary for sterilization can
be appreciable in cell walls in the interior of relatively large
blocks of wood such as are commonly utilized for cultures
of the wood rotting fungi. It seems that the effect upon
the lignin of a block of wood which the investigator is sure
has no unlignified cellulose originally, is so slight that it
may be neglected. This is still more true when we remem-
ber that simple soaking has a more or less solvent effect
upon the lignin in any method of culture work. There
can be no doubt that when finely divided wood is soaked in
water or is boiled, lignin is extracted in sufficient quantity
for lignin reactions to be obtained in the filtrate from the
extracted material. But this is very different from the
detection of that extraction by micro-chemical examina-
tion of the treated walls themselves; for it must be con-
fessed that micro-chemistry furnishes methods which are
far from accurate for quantitative analysis. .
In testing the action of fungi upon wood we must keep
in mind the solvent action of boiling and of continual
soaking in water and not be too hasty in drawing conclusions
from meager results ‘which might be confused with such
action of boiling or soaking. Control tubes of wet wood
STUDIES ON THE LIGNIN AND CELLULOSE OF woop. 57
and careful examination of the material used should obvi-
ate much of this difficulty however.
ENZYMES OF FUNGI.
Potter also states that, ‘‘ the delignification cannot be
entirely attributed to an enzyme secreted by fungi.’’ As
above stated it has been shown that the original thickness
of wood fiber walls is very materially reduced by some
agent at the very time when the fungus is attacking the
tissues in question. It is also shown that the cellulose which
may or may not be present in the walls defore they are at-
tacked certainly is not the same unlignified cellulose which
is present after the destruction of the wood by the fungus.
It seems certain then that fungi do delignify woody cell
walls.
There can be no doubt that enzymes, or some substance
exhibiting the characteristics of enzymes, have been proved
to exist in some of the fungi and indeed in some of the
wood rotting ones. Such investigations as have already
been published in this connection would seem to prove be-
yond any reasonable doubt that enzymes are secreted by
fungi and that they are of the most varied character.
Diastase seems to he proved to be present in many of the
different fungi and in the wood rotting ones. The disap-
pearance of starch in the early stages of attack of some of
the wood rotting fungi upon wood seems to be generally
attributed to the secretion of diastase by the fungi. Why
should the disappearance of lignin from the fiber walls,
leaving cellulose in the last stages of decay, be attributed
to any other cause than the secretion of a delignifying
enzyme until we have proof of some other agent which is
capable of such action? ‘To be sure, the separation of an
enzyme or a juice showing enzymic action from the fungus
being studied is the exact proof of the existence of an
enzyme; yet when we consider that enzymes are accepted
58 MISSOURI BOTANICAL GARDEN.
by practically all botanists as being present in fungi we can
hardly come to but one conclusion regarding the existence
of a delignifying enzyme.
EXPLANATION OF PLATES.
Plate 1.— 1, Portion of transverse section of wood of Populus tremu-
loides showing several wood fibers with a third, inner, cellulose layer.
This layer is colorless, as the section is stained with phloroglucin which
is a test for lignin. This supernumerary layer is characteristic of the
species of the genus Populus so far as they have yet been examined, five
of the endemic North American species from the various parts of the
country having been tested. 2, Portionof a transverse section of wood of
Pinus palustris showing one annual ring of growth. The thicker walled
cells form the summer wood of the ring, while the thin walled ones make
up the spring wood. This section has been boiled until the lignin was
extracted from the spring wood, and was then stained with phloroglucin,
atest forlignin. The red walls have reacted with the chemical show-
ing that lignin is still present in the summer fibers. The yellow cells
have retained their normal color to a greater or less extent and gave no
reaction with the phloroglucin, thus showing that they have been de-
lignified quite completely. The section shows the relative solubility of
the lignin in the two parts of the annual ring.
Plate 2.—1, Portion of a transverse section of wood of Picea rubens
boiled for 27 hours. This has been stained with chlor-iodide of zinc,
a test for cellulose. The cellulose has been colored blue while the other
parts of the walls are of their normal color. This shows the delignify-
ing action of boiling water upon the lignin of the fibers. 2, Transverse
section of wood of Populus balsamifera cut from a freshly felled tree and
stained with chlor-iodide of zinc. The thick supernumerary layer of
cellulose is colored blue. 38, Transverse section of wood of Sassafras
Sassafras boiled 20 hours and stained with chlor-iodide of zinc. This
is drawn with a smaller magnification than are the other figures. It
shows the frequency of the cellulose layer, in the fibers of the wood.
’ Note that there is no cellulose present in the last cells formed in the ring,
which arein a vertical line near the left of the figure. 4, Transverse
section of wood of Sassafras Sassafras drawn on the same scale as the
_ other figures. This was boiled and treated with chlor-iodide of zinc
‘and shows the character of the delignified layer in the fibers. Most
of the fibers adjacent to the vessels and tracheids do not have the
cellulose layer.
re
2
o
a
“d
4
hier
SAME CMe Meee Te ee eta pet at eM mag: oe ae | é * Seb. tre Si 1. eee ea ee ee ee ee ee ee ee ee
REPT, MO. BOT. GARD., VOL. 17. PLATE 1.
LIGNIN anp CELLULOSE.
REPT. MoO. Bot. GARD., VOL. 17. PLATE 2.
LIGNIN AND CELLULOSE
STUDIES UPON SOME CHROMOGENIC FUNGI WHICH
DISCOLOR WOOD.*
BY GEORGE GRANT HEDGCOCK,
GENERAL REMARKS.
The discoloration of unpainted woodwork exposed to
moisture and to oxidation by the air is a familiar phenom-
enon. Of course much of this discoloration is due to the
dirt and soot which accumulate on the surface of boards
in addition to the chemical changes which take place
through weathering. There is also a class of stains of an
entirely different nature and more striking to the eye.
These occur on freshly sawn lumber in piles. It was in
the investigation of the most important of these lumber
stains in the study of the western yellow pine, Pinus
ponderosa,t under Dr. Hermann von Schrenk of the Mis-
sissippi Valley Laboratory, of the United States Depart-
ment of Agriculture, that the studies upon which this
paper is based were begun. In addition to the blue stain
in pine lumber, brown, black, pink, purple and yellow
stains and blotches were noted, not only on pine boards,
but also on gum, poplar and other kinds. These were
found to be caused by a number of fungi, some of which
were quite different from that which causes the blue stain
of the western yellow pine, viz: Ceratostomella pilifera
(Fr.) Winter. Again the bluing of wood was found to be
the result of the action of more than one species of
Ceratostomella. |
* A thesis presented to the Faculty of Washington University, in
edndidacy for the degree of Doctor of Philosophy, April, 1906. Pub-
lished by. permission of the Secretary of Agriculture. — Type cultures
of all of the species described as new in this paper have been divided into
parts deposited respectively in the United States National Herbarium, the
Physiological and Pathological Herbarium of the United States Depart-
ment of Agriculture, and the Missouri Botanical Garden Herbarium.
+ Von Schrenk, H. Bull. Pl. Ind. 36: 1-27. (1903). (5
9)
60 MISSOURI BOTANICAL GARDEN.
The work of investigation has been undertaken primarily
on account of the value of the knowledge which might
accrue through the study of the chromogenic fungi or
bacteria which are concerned in the color reactions which
take place in stained lumber; yet, on the other hand, the
economic value of the work has not been lost sight of,
owing to the loss of hundreds of thousands of dollars to
the lumber industry each year due to the lowering of the
grade of lumber in piles through the rapid action of many
tiny wood staining fungi.
In the study of the fungus flora of the lumber pile a
large number of forms have been observed not all of which
discolor the substratum upon which they grow. Attention
has been given not only to those which penetrate wood
deeply and stain it, but also to those which discolor it only
superficially.
In the isolation and culture of the fungi herein de-
scribed, recourse was had to the most careful bacterio-
logical methods, such as are described by Dr. Erwin F.
Smith * in his excellent monograph on ‘ Bacteria in rela-
tion to plant diseases.’’ In a number of instances new
conidial stages of fungi have been discovered, and in every
instance the relation of the new form to the older known
forms has been repeatedly established by starting with the
newly discovered stage and following the fungus through
all its known stages and back again to the beginning. The
saprophytic nature of the forms studied made it possible to
start with single colonies on a poured plate of agar medium
and make transfer cultures to test tubes of sterile wood or
other media. Each fungus was grown upon a number of
kinds of wood, as well as upon potato, rice, bean, sweet,
potato, and other similar media in tubes, in addition to
cultures upon agar media made from wood and other veg-
etable decoctions.
In order to better establish varietal and specific charac-
* Smith, E. F. Bacteria in relation to plant diseases. 1. (1905).
eae ag? te,
ED Se Pa Le Re A Re ee Eee ee ee ae oe OUR?
Fal ots Oe AP. By SP ng ee ; ae
CHROMOGENIC FUNGI WHICH DISCOLOR WwoopD. 61
ters, parallel cultures of related fungi have been grown on
sets of the same media in conditions as nearly identical as
can obtain in an ordinarily well-equipped bacteriological
laboratory. In order to make drawings of the delicate coni-
dial stages of some species it was often found necessary to
grow the colonies in Petri dishes of thin glass, and study
them in the open in the dish, without placing a cover glass
over the very fragile hyphae.
It is thought best to arrange the results under the fol-
lowing groups or heads : —
WOOD-STAINING FUNGI.
I. Wood-bluing fungi.
1. Ceratostomella.
II. Wood-blackening and»wood-browning fungi. .
1. Graphium.
2. Hormodendron.
3. Hormiscium.
4. Other wood-blackening fungi.
III. Wood-reddening fungi.
1. Penicillium.
2. Fusarium.
In conclusion of these introductory remarks, grateful
mention should be made of the valuable assistance to the
work of investigation accruing through the use of the
library and other facilities of the Missouri Botanical Gar-
den, kindly tendered by Dr. William Trelease, the Direc-
tor. Acknowledgment also should be made of the helpful
co-operation of Dr. Hermann von Schrenk, Mr. Perley
Spaulding and Miss Laura L. Eames, of the Mississippi
Valley Laboratory, and to Dr. A. D. Hopkins, of the Bu-
reau of Entomology of the United States Department of
Acriculture, for securing material for study and granting
other similar favors.
I. WOOD-BLUING FUNGI.
The blue stain in pine wood has been known in Europe for
62 MISSOURI BOTANICAL GARDEN.
many years. Hartig * and Frank f both refer to it in their
publications on plant diseases. Hartig ascribes the cause
of bluing to Ceratostoma piliferum (Fr.) Fuckel, which is
now placed under the genus Ceratostomella according to
Winter. This species was first described by Fries,
who placed it under the genus Sphaeria, where it remained
with other species until Fuckel placed it under the genus
Ceratostoma. Winter,t in the revision of the genus Cera-
tostoma, placed all species with colorless ascospores under
the new genus Ceratostomella.
Von Schrenk § in a bulletin of the Bureau of Plant In-
dustry, U. S. Department of Agriculture, fully describes
both the fungus Ceratostomella pilifera and its effect and
mode of entrance into the wood of Pinus ponderosa.
A Jarge number of species of Ceratostomella are de-
scribed by Saccardo,{ many of which occur on wood, but
no reference is made to the staining effect of any of these
species. There is little doubt that a number of these
are wood-staining fungi, but from the results of our inves-
tigations it appears that the common bluing fungus is C.
pilifera.
1. CERATOSTOMELLA.
Winter in Michelia characterizes the genus Ceratosto-
mella as follows: ‘* Perithecia superficial or partly im-
mersed, usually tough, leathery or carbonaceous, glabrous
or invested with filaments, with prominent well developed
beak; asci without paraphyses, 8-spored; ascospores con-
tinuous, globose, ovoid or oblong, hyaline; spermogonia
and conidia present in some instances.”’
Although Winter mentions that conidia are present ina
few instances, a careful search through the literature on
the subject reveals that the number of species which have
* Hartig, R. Lehrbuch der Pflanzenkrankheiten. 75, 106. (1900).
; Frank, A.B. Krankheiten der Pflanzen. 1: 107. (1895).
¢ Saccarado, P. A. Michelia. 1: 370. § lc.
{| Saccardo,P. A. Sylloge Fungorum. 1-17.
CHROMOGENIC FUNGI WHICH DISCOLOR Woop. 63
s
conidia in their descriptions is very limited indeed, only
two being noted, and where such descriptions have been
given, no mention is made of the connection between the
conidial and the perfect or perithecial stages as being
established by cultural methods. For instance, Ceratos-
tomellaalbocoronata (Ellis) Sacc. has a conidial stage where
the conidia are from 2- to 3-septate, and of abnormally
large size. So far as our investigation has proceeded, all
species of Ceratostomella have colorless conidia, that are
one-celled and of about the same measurements as the
‘ascospores. The presence of asci in the perithecia has
been the most difficult point to prove. Cultures must be
taken at the stage just preceding full maturity of the ascus,
else it apparently dissolves and frees the ascospores at ma-
turity, allowing their free ejection through the long, nar-
rew beak of the perithecium, but destroying the evidence
of the presence of a sac. In an examination of the ejected
ascospores they are frequently found adhering to each
other in fours, side by side, in the same position they
occupied in the ascus.
The conidial stage of Ceratostomeiia is very important,
owing to the immense number of conidia borne on the
mycelium in its earlier growth. These are readily dissem-
inated by the wind and are probably carried by insects
which penetrate the wood and bark of trees, like most of
the ambrosia and bark beetles. At the stage in which the
conidia form a mucilaginous mass, they adhere readily to
any insect that may pass over them. In the laboratory a
number of species of mites which feed on fungi carried
spores on their bodies from a colony in an agar plate to a
sterile portion of the surface of the medium and started
new colonies of the fungus. Bark beetles were placed in
a dish with the conidial stage of Ceratostomella, and after
allowing them to remain a short time were transferred to
sterile agar plates which were inoculated with spores
from the insects. It is probable that some species of in-
ce
cai alia ection Sai
™= os "¢ ype e
APOE LER NERS 5 RIVE NT God Rae yee eg ME ey RR eae eee ae
PEE Sa hghv OSES WE AE eID ena 7 . anata ae
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64 MISSOURI BOTANICAL GARDEN.
sects feed on the conidial stage of Ceratostomedla, especially
one or more species of ambrosia beetles and a number of
mites infesting their channels in the wood; but proof is
yet lacking on this point. The constant occurrence of this
fungus in the channels of a number of wood boring beetles
indieates that the conidia or the ascospores must be carried
in some manner by these insects. Hopkins* describes
some of these beetles in a bulletin of the U. S. Department
of Agriculture, and von Schrenkf describes how C. pilifera
follows the channels of wood-boring beetles in Pinus
ponderosa. The writer has made similar observations on
Ceratosmella in its penetration of Pinus ponderosa, P.
Arizonica, P. echinata and P. Virginiana.
Although the genus Ceratostoma is very closely related
to Ceratostomella none of the species of Ceratostoma have
been found in connection with the blue stain of wood. In
the course of the present investigation stained wood has
been collected from a large number of localities, including
wood from trees and shrubs of the following genera: Abies,
Acer, Fagus, Fraxinus, Liquidambar, Liriodendron,
Pinus, Rubus, Ulmus, Vitis and Wistaria. A number of
species and varieties of Ceratostomella have been identified,
cultivated and proved to be wood-bluing fungi. Ceratos-
tomella pilifera has been found far more prevalent than
other species. As the conidial stage of this species was
first discovered by the writer and the description has never
been published, it will now be given. It is taken fromthe
results obtained from a very large number of pure cultures
grown and studied over a period of nearly four years.
Ceratostomella pilifera (Fr.) Wint.
This fungus has been isolated in cultures taken from
stained wood from a number of species of Pinus taken
from nearly every region of the United States. It pene-
* Hopkins, A. D. Bull. Div. Ent. U. S; Dept. Agr. 32: 9, 10.
+ Von Schrenk, H. J. ¢. :
CHROMOGENIC FUNGI WHICH DISCOLOR WOOD.) 65
trates rapidly the sap wood of dead and dying trees, giving
it a peculiar blue tint whose color is hard to explain, since
the mycelium of the fungus in the cells of the wood is a
dark brown. It has also been found in the sapwood of
species of Abies, Quercus, Fraxinus, and other genera.
Ceratostomella pilifera grows readily upon a large num-
ber of culture media. Pine decoction agar made from the
bark and sapwood was found an excellent medium for its
cultivation. Two distinct fruiting forms were observed in
all the varieties, the conidial and the perfect or perithecial.
MYCELIUM.
When either the conidia or ascospores are sown In pine
agar plates, germination takes place in a few hours, and in
two days colonies with a white hyaline mycelium develop.
The filaments areseptate, and, asa rule, branch alternately.
In a day or so upright hyphae, either simple or branched,
are sent out, upon which are borne branching whorls of
conidia. Inthe course of a week or so, portions of the
mycelium in the older region of the colonies develop with
thicker walls and assume a brown color. From these dark
colored filaments the perithecia originate.
The first growth of the mycelium is usually sparse,
but under favorable conditions a secondary profuse
growth with a fluffy white appearance develops, which
bears conidia, and the formation of perithecia is retarded.
Such a growth is formed on some of the richer media
made from pine and other plant decoctions. Under these
conditions the development of conidia is enormous, and
the number of perithecia is decreased somewhat. Cultures
on the sapwood of trees in test tubes bear both stages of
the fungus abundantly. On the heartwood, however, but
few conidia are formed, the mycelium is almost invisible to
the eye, and perithecia are either sparse or entirely absent.
The filaments of the fungus never penetrate deeply into
5
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66 MISSOURI BOTANICAL GARDEN.
the tissues of heartwood of pine, oak, ash, or similar
woods in artificial cultures, which fact coincides with the
observation that in nature, for some reason, the same
thing happens, the older discolored heartwood being
usually free from attacks by the mycelium.
ConIDIA.
The conidia are borne on erect or-slanting hyphae which
branch alternately from the mycelium. They develop
terminally in branched or simple moniliform short chains
which are often in whorls (pl. 4,f.5). They are de-
tached by the least movement, and in moist air fall’
together in agglutinated masses about the terminus of the
hyphae, in a form which superficially resembles Cephalo-
sporium (pl. 4, f. 6). These masses, however, are not so
regular as the masses of conidia in the heads formed by
Cephalosporium. The conidia bear some resemblance
to those of Ovularia. The Cephalosporium-like clus-
ters are always present in agar cultures after a few
days’ growth. The conidia vary from obovate to elliptical
or cylindrical. They measure from 8 to 12 in length,
and from 2 to 4m in width. They are hyaline when
young, but often become vacuolate or guttulate when old.
They are unicellular and borne on septate hyphae which
are from 3p to 5y in diameter. Only hyaline filaments
bear conidia. In germination the conidia send out terminal
germ tubes from either end (pl. 4, f. 7).
PERITHECIA.
The first indication of the formation of perithecia on the
mycelium is the union of two or more filaments and the
formation of a knotted mass. The adjacent mycelium
changes first to a light brown color, and later to a very
dark brown. In the center of the mass the young peri-
thecium develops first as a globular black body, without a
beak or neck.
The walls of the perithecium are usually formed by one
CHROMOGENIC FUNGI WHICH DISCOLOR WOOD. 67
layer of cells united in. a rudimentary tissue. From the
upper side of the young perithecium, after it has grown to
about its normal size, there is projected a long tube or beak
composed of parallel hyphae which terminate with colorless
ends in a rounded tip. When the perithecium is mature it
contains a large number of irregularly obovate or elliptical
asci, each containing eight spores, usually arranged in
fours. The beaked ostiolum, when the ascospores are
ejected, splits at the end into a number of bristles which
spread out and form a supporting crown for the ejected
ascospores (pl. 3, f. 8). The perithecium without the
neck measures 160u in height and 180, in diameter, aver-
age, and the neck about 1 mm. in height and 20y in diam-
eter, average. The ascospores average 5.5u by 2.5p.
It is quite evident that the sac containing the ascospores
is dissolved at maturity, since it is rarely found with the
mature ejected spores, but must be looked for in perithecia
just approaching this stage. If the spores are ejected in
open air they usually collect in a globular drop at the
terminus of the neck, and when in this position the drop
resembles very much the heads of some species of
Graphium when covered with the mucus drop characteris-
tic of the latter genus. If the spores are discharged in
water they form worm-like sticky masses not readily mis-
cible with water.
Ceratostomella Schrenkiana n. sp.
The sapwood of pine lumber made from Pinus echinata
in Arkansas and other southern localities is stained by a
number of fungi, among which are species of Ceratosto-
mella, Hormodendron, Graphium, Hormiscium and Peni-
cillium. A species of Ceratostomella which greatly re-
sembles C’. pilifera has been frequently collected on the
wood of the southern pines by Dr. von Schrenk and others,
from various localities in Missouri and Arkansas.
This species, when grown side by side with cultures of
C. piliferafrom the Black Hills pine, attains the same
eee, ere ae
fn *
See
68 MISSOURI BOTANICAL GARDEN.
size in so far as the colonies and perithecia are concerned ;
but on the other hand, both the conidia and ascospores are
constantly much smaller. In nature, the perithecia that
mature are often surrounded by several closely adhering
dark bodies, which are either sclerotia or abortive peri-
thecia, apparently the latter. This feature has never been
noted with other species of Ceratostomella. In view of
the differences, it is thought best to separate this form
from C. pilifera and it is named C. Schrenkiana, in
honor of Doctor von Schrenk, who first collected the
fungus and noted its peculiarities.
The following cultural characters are taken from a large
number of natural and artificial cultures : —
MYCELIUM.
Conidia or asco-
v " . p spores, if sown on agar
VV RAY plate cultures, germi-
- Te fz nate in a few hours,
and after a day have
formed conidia, if the
temperature is favora-
\ ( ble. The most favora-
A! V ble temperature for
W tA growth is 80° to 90°
NU F. The filaments and
hyphae remain white
ae for several days, and
often become massed
in a furry outgrowth in
ments unite into upright
As clusters or false heads.
Pc LNA «(See figure). After a
~~ few days’ growth, por-
which strands of fila-—
C. SCHRENKIANA, X 50.
Dendroid fruiting tuft.
tions of the mycelium
lying next the agar
become pigmented with a brown color, and rapidly develop
Ss Ra ar)
CHROMOGENIC FUNGI WHICH DISCOLOR WOOD. 6Y
perithecia. The filaments of the mycelium measure 5p to
7m in diameter.
CONIDIA.
The conidia are borne in clusters similar to those of
C. pilifera (pl. 4,f. 1,2). They are unicellular, obovate
to elliptical or cylindrical, and measure 3p to 7p by lp to
Qu, averaging 5u by 1.84. They are hyaline and are
usually neither guttulate nor granular, but when old may
have one or two guttules.
PERITHECIA.
Perithecia are formed on the mycelium (pl. 4, f. 4), as
in C. pilifera. The shape is spherical. They are coal
black in color, and measure 120u to 200u in diameter, with
a smooth, beaked ostiolum, measuring .8mm. to 1.2mm.
in length by 10% to 254 ia diameter, surmounted at ma-
turity by a row of short, hyaline, spreading bristles, which
support the. ejected ascospores ina globular mass. ‘The
bristles measure about 10u to 15y by 2u (pl.3,f.6). The
ascospores are hyaline, pointed, elliptical, often slighily
curved, and measure 2.54 to 4p by 1p to 1.5p, averaging
3.54 by lw (pl. 4, f. 3).
This species resembles, in its gross measurements, the
description of C. echinellaE. & E. It differs, however,
_ in that the perithecia are borne superficially, and are not
glandular-pubescent or thickened at the tips. The mature
perithecia are not gregarious, although they bear numerous
small bodies at their base, under certain conditions.
Ceratostomella echinella E. & E.
The last species of Ceratostomella identified and studied
before publication was collected by Dr. von Schrenk at Kir-
byville, Texas, on the wood of the red beech, Fagus atropu-
nicea (Marsh.) Sudworth. This was growing on freshly
cut heart and sap-wood, staining the wood either a bluish or
Aly Re pies eens ek th
70 MISSOURI BOTANICAL GARDEN.
a brownish color. The fungus was isolated and grown in
pure cultures on the wood of the red gum and on pine de-
coction agar, as well as on tubes of rice, potato, etc., the
following cultural characters being from both natural and
artificial cultures. The description differs but slightly
from that given by Ellis and Everhart* and has been
emended by the addition of the conidial stage.
MyYcELium.
The colonies resulting from either ascospores or conidia
are white at first with a hyaline mycelium which later’
becomes pigmented in certain of the larger filaments, assum-
ing a dark brown color. The filaments measure from 4
to 7 in the mature forms. Conidia appear after 36
hours, and perithecia in four or five days.
CONIDIA.
The conidia are borne in short branching moniliform
chains in small clusters, falling together in irregular masses
as they mature (pl. 6,f.1). They are unicellular, hya-
line, becoming guttulate when old, obovate to elliptical,
and measure from 4p to 6.54 by 2u to 3.5p, averaging 6
by 3z.
PERITHECIA.
The perithecia are brown at first, but when they are ma-
ture they become black and carbonaceous and are covered
sparsely with short, glandular hairs. They are globose,
or sometimes slightly flattened, measuring 50p to 100p in
diameter. The hairs measure 104 to 324 in length by
1.5mto 2, Each terminatesina gland or spherical enlarge-
ment measuring 24 to 34 in diameter. The perithecium is
tipped by a long slender beak, black at the base, brown
* Ellis, J. B. and Everhart, B. M. North American Pyrenomycetes.
p- 195. (1892).
¢
CHROMOGENIC FUNGI WHICH DISCOLOR WOOD. 71
or gray at the apex, but smooth throughout measuring
1 mm. to 1.7 mm. in length by 15» to 25 in diameter
and surmounted when mature by a terminal fringe of
bristles 15 to 254 in length and tapering from 4p at the
base to lp at the apex, being hyaline throughout (pl. 3, f.
3). The asci are clavate(?), bearing biseriate spores
which are hyaline, curved or straight, elliptic to cylindri-
cal, and which are ejected in the open air ina globular mass,
at the terminus of the beak.
Since writing the above, co-type material of C’. echinella
inthe herbarium of the Missouri Botanical Garden has been
examined. ‘The perithecia are probably as described by
Ellis and Everhart,* but they have persistent asci which
contain brown two-celled ascospores, placing the fungus in
the specimen under another genus. This indicates that at
least a mistake was made in a portion of the material sent
out as type specimens if not in the description.
Ceratostomella capillifera n. sp.
The wood of Liquidambar Styraciflua L. is an excellent
pabulum for a large number of fungi, and lumber made
from this tree, when in piles, soon becomes thoroughly
stained either blue or black by the growth of a number of
species of wood-discoloring fungi. Among these was
found quite commonly a species of Ceratostomella which
is closely related to C. pilifera, but differs in the length
of the beaked ostielum, the length of the terminal bristles
and the shape and size of the conidia and ascospores.
The following characters obtain with pure cultures : —
MYCELIUM.
Cultures of either conidia or ascospores sown on pine-
decoation agar germinate quickly, forming a diffuse, white,
hyaline, septate mycelium, which in less than two days
* Ellis, J. B. and Everhart, B. M. J. c.
:
;
5.
72 MISSOURI BOTANICAL GARDEN.
bears hyaline conidia abundantly. In about a week, black,
long-beaked pycnidia appear on certain portions of the
mycelium, which assumes a brown color, the development
being similar to that described for C. pilifera. The fila-
ments of the hyaline portion of the mycelium measure
from 2u to 8 in diameter, and the older brown-colored
filaments, in wood, from 3p to 6y,
CONIDIA.
The conidia are borne like those of C. pluriannulata,
and are similar in shape (pl. 6, f. 2,3). They are unicel-
lular, hyaline, elliptical to cylindrical, and measure 4 to
8» in length, and 1.5y to 2u in width averaging 64 by 1.8y.
PERITHECIA,
The perithecia are spherical, of a black color, with
black, long-beaked ostiola, terminating in long, wavy
hyaline filaments. The body of the perithecium is usually
nearly covered with dark brown hyphae or filaments. It
averages 200m in diameter, and the neck 1.5 mm. in length
and 254 in width. The terminal filaments are long and
slender and measure 804 by 1p, average (pl. 3, f. 1).
The ascospores are elliptical to reniform in shape and
average 4.54 by 1.5uin size. It will be noted that al-
though the perithecia of this species are larger than
those of C. pilifera the conidia are somewhat smaller.
Both ascospores and conidia usually germinate terminally,
some few exceptions having been noted.
The name C. capillifera is given to this species on
account of the long minute bristles which project from the
ostiolum.
Ceratostomella pluriannulata n. sp.
A block of blued sapwood from Quercus rubra was col-
lected in 1905 by Perley Spaulding and turned over to the
CHROMOGENIC FUNGI WHICH DISCOLOR WOOD. 73
author for investigation. A species of Ceratostomella was
found among the cells of the wood and fruiting quite
abundantly on the surface. The characters of the fungus
agree in part with C’. pilifera, to which it is closely related.
‘It has ascospores which are constantly of a slightly differ-
ent shape, and is often characterized by more than one
whorl of bristles on the beaked ostiolum. The conidia are
considerably smaller, and the hyphae on which they are
borne are more frequently branched, thus giving rise to
larger clusters. Owing to the frequent presence of more
than one ring of bristles on the ostiolum, the name C.
pluriannulata is here given to the species, with the fol-
lowing cultural characters : —
MYCELIUM.
The growth of the mycelium from either conidia or
ascospores resembles that of C. pilifera. The spores
germinate in pine-decoction agar in 2 to 3 hours, and in 2 to
3 days a white, floccose mycelium bearing conidia appears.
In from 5 to 10 days perithecia appear, the development
of which is like that of C’. pilifera.
ConrIDIA.
The hyaline unicellular conidia are borne on branching -
septate, hyaline, upright hyphae in simple or compound
whorls (pl. 5, f.1). In the older portions of the colony,
the conidia fall off and adhere in masses forming clusters
resembling Cephalosporium. These clusters vary in size
from 10u to 100m in diameter. The larger clusters are
formed by the adherence of several spore clusters into one
mass. The conidia measure 54 to 8u in length and 2p to
3u in width, averaging 6p by 1.5p.
PERITHECIA.
The perithecia are borne superficially on the wood, or on
the mycelium, there being no definite stroma in this or in
74 MISSOURI BOTANICAL GARDEN.
the other species of Ceratostomella studied. The peri-
thecia are globose, varying from 904 to 200m in diameter,
and averaging 1204. The lengthened beak measures from
.9 to 2 mm. in length, and 104 to 30m in diameter, aver-
aging 1.5 mm. by 254. Some of the necks, in addition to
the ring of bristles ordinarily found at the terminus of the
beaked ostiolum, have one or two whorls at some distance
below the end (pl. 8, f.7). This character has never
been noted in cultures of C. pilifera. The terminal
bristles are tapering and average 20 in length by 2y in
width. The asci have not been measured. The ascospores
are reniform in shape, hyaline, and measure 4p to 5y in
length, and 1.54 to 1.7 in width, averaging 4.54 by 1.54
(pl. &, £..2).
This species penetrates pine wood when inoculations are
made, nearly as readily as C’. pilifera. The mycelium in
the wood cells follows the medullary rays in pine wood,
but in red gum wood it penetrates also numerous tracheary
vessels, and sometimes the wood fibres. It is most abun-
dant, however, in the medullary rays of the latter wood.
Ceratostomella minor, n. sp.
During the year 1905 some specimens of pine wood
stained with a dark blue color, from Pinus Arizonica Eng.,
were received from Dr. A. D. Hopkins. From these there
were isolated two species of wood-staining fungi; one
a species of Ceratostomella, the other a Graphium. The
Ceratostomella proved to be smaller than the species from
Pinus Virginiana, and differed in other points. The fun-
gus gained entrance through the galleries of wood beetles,
the stain radiating from cavities made by these insects.
When grown in pure culture the fungus possessed conidia
and ascospores. As the specific characters do not agree
with any previous description of Ceratostomella, the name
CHROMOGENIC FUNGI WHICH DISCOLOR woop. 175
C’. minor is assigned, with the following cultural charac-
ters: — :
MYCELIUM.
Cultures of eitner conidia or ascospores on pine agar
media germinated in a few hours, and in two days the col-
ovies began to form conidia. The mycelium is white and
sparse. The hyaline filaments of the conidial stage measure
in width 1.5y to 2.6, averaging 2.34. The brown or black
filaments in agar cultures and in wood measure in diameter
from 2u to 4p, averaging 3.54. These are often rugose,
the roughenings being coarser than those in C’. exigua.
In some portions of the mycelium on wood there are
filaments that contain cells that are unequal in diameter,
being enlarged at one end. Like C. pilifera this species
follows chiefly the medullary rays of pine wood.
CONIDIA.
The formation of the conidia and the form of fruiting
do not differ essentially from those of C. pilifera and
C. exigua. The unicellular, hyaline conidia measure from
4 to 5.54 in length, and from 1.84 to 2m in width, aver-
aging 4.54 by 2u. They areoval to elliptical in shape, and
fall off at the slightest touch, collecting in rounded masses
on the hyphae when in moist air.
PERITHECIA.
The perithecia mature in about three weeks in artificial
cultures. They are spherical, black rugose bodies, vary-
ing from 40 to 70» in diameter, averaging 524, witha
beaked ostiolum 120y to 160uin length, averaging 135p,
and from 6u to 124 in diameter (pl. 5, f.6). The beak
ends in a whorl of short, thick bristles (pl. 8, f. 4). The
asci are round to oval, hyaline, with eight spores in each,
usually arranged in fours (pl. 5, f. 7). The spores meas-
ure 3.1uto 4.2uin length and .9 to 1.94 in width, averag-
76 MISSOURI BOTANICAL GARDEN.
ing 3.54 by 1.54. They are hyaline and germinate termi-
nally.
The perithecia are smaller than those of C’. exigua, the
description of which follows; the ascospores differ in shape,
and are smaller, and the terminal bristles of the beak have
a different shape, being short and thick as compared with
the slender ones of C. exigua.
Ceratostomella exiqgua n. sp.
The wood of dead and dying trees of Pinus Virginiana
Mill., and one or two other species, is often colored a blue-
black color, even more intense than that produced by Cer-
atostomella pilifera in wood of the western yellow pine.
Specimens were first obtained for study from Dr. A. D.
Hopkins of the United States Department of Agriculture.
It was at first thought that the stain was caused by C. pili-
fera, but pure cultures of the fungus proved that it was
different ina number of particulars. The scanty mycelium,
the numerous diminutive perithecia and tbe smaller coni-
dia, as well as the more intense color of the stain, were
prominent points of difference which were considered of
specific importance. These characteristics remained quite
constant through a series of cultures on various kinds of
wood.
The life history of the fungus obtained from a series of
pure cultures is as follows: —
MYCELIUM.
Cultures grown on pine decoction agar from both conidia
and ascospores give in a short time colonies with a white
mycelium much more limited in growth than that of C. pili-
Sera. The conidial stage appears after 24 hours, and is
of the same type as that of C. pilifera, but with the con-
idia in lesser numbers. In a few days the perithecia ap-
pear on a mycelium which assumes a very dark brown,
CHROMOGENIC FUNGI WHICH DISCOLOR WOOD. 77
*
almost black, color. The development of the fungus on
wood is slower than that of C. pilifera, staining pine, oak,
elm and ash with a blue-black discoloration contrasting
sharply with the unstained portions. The mycelium pen-
etrates pine wood through the medullary rays, and in some
cases enters the wood fibres. The hyphae of the hyaline
mycelium bearing conidia average 2.5y in diameter, while
the dark brown filaments in wood measure from 2p to
6u, averaging 4 in diameter. On the surface of wood
in the vicinity of perithecia some filaments have swollen
cells. Many of the filaments in wood cells have fine trans-
parent roughenings all over the surface.
ConrpIA.
The conidia, like those of C. pilifera, are borne in
whorls of single spores and short moniliform chains inter-
mingled (pl. 6, f. 4,5). In older cultures they drop
together in clusters resembling somewhat those of Cepha-
losporium. They are oval to elliptical in shape, one-celled,
hyaline, and vary in size from 3.54 to 4.5, in length, and
1.64 to 2.2uin width, averaging 4u by 2y (pl. 6, f. 6).
PERITHECIA.
In artificial as well as in natural growths, the perithecia
are very abundant. They are formed in large numbers
under favorable conditions after 10 to 14 days. The pro-
cess of formation is similar to that of C. pilifera. The
perithecia are globose. They are black, brittle, with a
roughened surface. They measure 604 to 80, averag-
ing 73 in diameter, with a beaked ostiolum 150p to 2004
in length, and 8u to 18m in diameter, averaging 1804 by
14u (pl. 3, f. 2). The asci are hyaline, irregularly
oval to elliptical, and contain 8 spores, often arranged in
fours. The ascospores are hyaline, elliptical, sometimes
slightly curved, and measure 2.1luto 2.84 in length, and
8 to 1.1uin width, averaging 2.54 by lp (pl. 6, f. 7).
The sacs usually disappear before the spores are ejected.
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F REP a ed
78 . MISSOURI BOTANICAL GARDEN.
This species resembles the description of C. microspora
E. and E. but differs in the shape of the ascospores, the
size of the perithecia, and the length of the beaks. It has,
in addition, a conidial stage, which serves to further set it
apart as a new species. On account of the minuteness of
its perithecia and spores it is named C’. ewigua.
Ceratostomella moniliformis n. sp.
Another species of Ceratostomella was found by Dr.
von Schrenk growing on gum wood in Texas, near Kirby-
ville, occurring on Liquidambar Styraciflua. This species
stains the wood of gum a brown color, and is the most
rapid growing form studied. It is distinct from the other
species of Ceratostomella in the form of the conidial
clusters, and in the early color of the mycelium. The peri-
thecia are covered with short spines, otherwise resembling
very much those of C. pilifera. The name Ceratostomella
moniliformis is given to the fungus, with the following
cultural characters :—
MYCELIUM.
Cultures of either conidia or ascospores sown in agar
plates germinate quickly, andin 24 hours produce the con-.
idial stage. In two or three days the perithecia are pro-
duced. The colony is hyaline at first, but in a few hours
begins to turn gray, and finally becomes black. The my-
celium is coarsely granular, the filaments measuring 24 to
8 in diameter.
CONIDIA.
The conidia are found on simple or branching, upright
hyphae, both in simple moniliform chains and in terminal
clusters, the latter resembling the form occurring with
some species of Graphium (pl. 5, f. 3, 4,5). They
are formed by the abstriction of the ends of the hyphae,
and often fall together as fast as they mature, in rounded :
.
e
Pea ae a ni bee Rake Si Ried a hearer) 2" ed oO Re eg Se See
ay 4 ng a et BE LOGE mE SRP OS ta aa
iy Saka: Sa
oo ee
Seats
aes,
CHROMOGENIC FUNGI WHICH DISCOLOR WooD. 79
masses, surrounded by either water or mucilage. They
are unicellular, cylindrical in shape, hyaline, and measure
64 to 8u by 1.8% to 2.2u, being of different shape
and dimensions from the ascospores, and are thus easily
distinguished from them under the microscope. The con-
idia in mass as they become old have a gray color.
PERITHECIA.
The perithecia are formed by the union of two or more
filaments of the mycelium. First, a gray mass of irregular
cells appears at the pointof union. This rapidly develops
into a globular perithecium, from the top of which there is
then rapidly thrown out a beaked ostiolum, consisting of
parallel filaments. The perithecia are brown to black in
color, 90% to 180 in diameter, covered with sparse, con-
ical spines, 12% to 16 long, with a diameter of 6y at
the base. The beaked ostiolum is brown to black through-
out its length, striate, and is surmounted by short, rather
thick, hyaline bristles, measuring 124 to 184 by 2u
(pl. 8, f. 5). When the perithecia are mature, the asco-
spores are ejected in water in long, slimy, gray masses.
In nature they are ejected in an irregular gray mass sup-
ported by the terminal filaments of the beak. The asciare
fugacious, hyaline, oval, and average 20uby 10u. The
ascospores are hyaline, not guttulate, at least at first, oval,
often flattened on one side, measuring 4¢ to 54 by 3u to
4p.
This species of Ceratostomella is so different in color
from the other species studied as to suggest that it may be
a Ceratostoma. The gray color of the mycelium on media
in which the mycelium of other species remains hyaline,
the constant tint of gray in mature conidia and ascospores,
and the peculiar moniliform chains of conidia very distinct-
ly set this form apart at least as avery distinct species.
But since the color of the spores shows only in mass, it is
Bf
5
; 3
3
3
s
80 - MISSOURI BOTANICAL GARDEN.
thought best to place the species at present under Ceratos-
tomella.
II. WOOD-BLACKENING AND WOOD-BROWNING FUNGI.
1. GRAPHIUM.
Graphium has been associated with the decay of wood and
other organic matter ever since the original species were
named by Tode* and Corda,f and placed under Stilbum
by Corda. It has been found also to be associated with
the staining of lumber, especially of pine and gum boards.
It has been isolated from black or brown stained portions
of wood taken from Pinus, Populus, Liriodendron,
Liquidambar, Quercus, Acer and Wistaria. Species of
this genus, like those of Ceratostomella, enter most kinds of
wood through the medullary rays, being usually confined
to the sapwood, with the exception that they may grow
upon decayed heartwood that other fungi have previously
attacked. Inthe study of anumber of species of Graphium
not all species stained wood alike: some scarcely stained
the wood at all, others, like the one from Liguidambar,
are quite effective as wood stainers.
The stain given by Graphium varies from a dirty gray
to a dark brown, or, rarely, a black color.
The genus Graphium was first described by Corda. It
is described as follows, using a free translation: — ‘* Stroma
cylindrical, clavate, or capitate, brownish, rather rigid,
the upper hyphae paler, lax, and bearing the conidia,
which are elliptical or oblong, hyaline, often involved in
mucus at first.”” No mention is made either by Saccardo,t
or Engler and Prantl { of any conidial stage other than
that found on the stroma or head.
* Tode. Fungi Meckl. 1:10. + Corda. Icon. Fung. 1:8.
t Saccardo, P. A. Michelia. 2:32; Syll. Fung. 43 564, 609.
¢{ Engler, A. and Prantl, K. Die Natiiriichen Pflanzenfamilien. 1 : 493.
4
Sieh oa oe
Se Tonge
oe ;
CHROMOGENIC FUNGI WHICH DISCOLOR woop. 81
Boulanger * has proven by the use of careful cultural
methods that Sporotrichum chlorinum Link var. grisea
Boul. is astage of Graphium eumorphum Sacc. This dis-
covery is now verified by the present investigation, and
Sporotrichum-like stages of other species of Graphium are
now published for the first time. Boulanger also discov-
ered a Sporotrichum stage of Chaetomium cuniculorum,
and obtained fruits of the latter fungus in connection with
Graphium, indicating that Chaetomium cuniculorum is
the perfect stage of the other two fungi. We have not
been able to verify the latter relationship.
Graphium giganteum (Pk.) Sace. is given by Durand t
as a conidial stage of Holwaya gigantea (Pk.) Durand.
Zopf t observed a moniliform conidial stage of Chae-
tomium which failed to germinate. He makes no mention
of finding any related forms of Sporotrichum.
Some of the species we have studied contain both Sporo-
trichum forms and moniliform fruits, the latter resembling
very much the branching clusters of the conidial stage of
Ceratostomella.
From the study of several species of Graphium the fol-
lowing cultural characters are found to be common :—
MYCELIUM.
The mycelium in a culture from either of the two spore-
forms found in the species studied is from the first hyaline
and septate. On rich vegetable-decoction agar, like that
made from green pine sapwood, bean pods, potato, etc., a
fluffy white mycelium is usually produced, varying in
density with the medium and the species. The first form
of conidia is borne on erect hyphae, which may be either
simple or branched. These hyphae or conidiophores
* Boulanger. Rev. Gén. de Bot. 1895 97-102.
+ Durand, E J. Bull. Torr. Bot. Club, 28% 351. (1901).
t Zopf, W. Ueber Pilzfarbstoffe. III. Bot. Zeit. 47; 86-89. (1889).
6
82 MISSOURI BOTANICAL GARDEN.
branch alternately from the mycelium. These first conidia
are borne in clusters of three types, two of which are
simple clusters of the type of those of Sporotrichum but
not borne on prostrate hyphae, the other being a form
which is like most of the conidia described under Cera-
tostomella. In this latter form the conidia are borne in
the short branching chains. These conidia are all ephem-
eral and do not hold their vitality as long as those which are
produced later upon the heads or stromata. In order to
distinguish them from the latter, they are now for the first
time called secondary conidia; and those of the heads or
stromata are now called primary conidia. As the colony
grows older, some portions of the mycelium become more
or less pigmented, and the darker filaments grow together
in masses from which erect columns of parallel hyphae
spring forth. These bear heads composed of branched
hyphae, which produce the primary conidia terminally by
abstriction. As fast as the conidia are formed they drop —
away into a mucous matrix which forms over the surface of
the head. .
SECONDARY CONIDIA.
The secondary conidia of the Sporotrichum-type are of
two forms and are borne on simple erect or branched
hyphae. The first form of this type are attached to the
hyphae in spreading, open clusters, which do not fall to-
gether in masses until some days after they mature (pl. 7,
f.3). The second form are produced in close clusters
which fall together as rapidly as they mature (pl. 9, f. 9).
The third form of fruiting, as mentioned in the previous
topic, more nearly resembles the branching chains of the
conidia of many species of Ceratostomella (pl. 9, f. 3).
The spores in all three forms are oval, elliptical, or cylin-
drical, and correspond closely in form and measurement to
the primary conidia of the same species. They are unicel-
lular and formed by abstriction either from the terminus of
the hyphae or from short, terminal or lateral branches.
stalks is composed of a num-
CHROMOGENIC FUNGI WHICH DISCOLOR woop. 83 tae
PRIMARY CONIDIA. ; é
After the primary conidia have been detached and sep-
arated from the mucus there is in many species nothing to | a
distinguish them from the secondary conidia. In fact, the ..
real difference usually in the two forms is whether they ; .
are borne on a single hypha, or on hyphae united in a
stroma. The agglutinated cluster on a single stalk of one
hypha éorteapands morphologically to the mucous-covered
head (pl. 7, f. 2,5). The latter are more persistent, and
retain the conidia for a much longer time. A secondary :
growth of the stromata has been “noted in some species | :
after the first spores have been formed, and have dropped :
away, in which there are formed branches from the end of
an old stalk, growing out |
from the apex. (See figure ).
Each of these subsidiary
ber of parallel filaments, or
hyphae, which in turn
branch at the outer ends and
form masses of conidia like
the original head, only small-
er. Such growths take
place on rich, starchy media
like cooked potatoes. On
cultures on rice and potato
an abortive type of stromata
is often formed in the shape
of somewhat flat, branching
stalks, which grow much tall-
er than the fruiting form,
but bear no conidia. This
abortive form resembles very
much the original description
of the sterile fungus Anthina Fr., which is probably a
form of Graphium.
G. EUMORPHUM, X 50. .
Proliferation of stroma. a
84 MISSOURI BOTANICAL GARDEN.
In the gross appearance of artificial cultures of Graph-
ium and of Ceratostomella there is often a striking resem-
blance or mimicry. The secondary conidial stage of the
former resembles superficially the conidial stage of the lat-
ter, and the mucous head of the former that of the perithe-
cial stage of the latter at the time the ascospores are ejected
ina viscid drop. The mucus surrounding the conidia of
Graphium is soluble in water; on the other hand, the jelly-
like substance exuded from the perithecia of Ceratostomella
is very difficultly soluble in water. Both are well adapted
for adhering to the bodies of insects with which they come
in contact.
PEeERFECT OR CHAETOMIUM STAGE.
The Chaetomium stage of Graphium observed by Bou-
langer has not been found in connection with the species
studied; but asthe cultural work on G'raphium has not yet
been concluded, it is hoped that by the use of the proper
culture media some species may finally be stimulated to pro-
duce perfect forms, either of Chaetomium or other genera.
In the consideration of the species of Graphium studied,
it is thought best to separate them into the following groups
based on the form of fruiting in the secondary conidial
stage : —
A. Species of Graphium with a secondary conidial stage with conidia
resembling those of Sporotrichum.
B. Species of Graphium with a secondary conidial stage unlike Sporo-
trichum,
1. Species with secondary conidia borne continuously and terminally,
falling at once in clusters.
2. Species with secondary conidia borne either in simple clusters or
in clusters of short branched chains.
CHROMOGENIC FUNGI WHICH DISCOLOR WOOD. 85
A. Sprecies oF GRAPHIUM WITH A SECONDARY CONIDIAL
STAGE WITH CONIDIA RESEMBLING THOSE OF SPORO-
TRICHUM.
Graphium ambrosiigerum n. sp.
A species of Graphium was found growing in the gal-
leries of wood-boring beetles in the wood of Pinus Arizon-
ica Eng. This material was furnished by Dr. A. D. Hop-
kins in connection with the study of the relation between
ambrosia fungi and ambrosia beetles, a subject which is still
under consideration in connection with the study of such
beetles by Dr. Hopkins. Pieces were taken from the
stained wood in the interior of the block near a gallery,
and placed in test tubes of pine sapwood, which had been
sterilized after being moistened. From the mycelium in
the stained wood there grew out a white growth, bearing at
first the secondary then the primary conidia of Graphium.
Secondary conidia were found in some of the galleries
when the material was first examined, but were not recog-
nized as such at the time. The following are the cultural
characters of the fungus :—
MYCELIUM AND SECONDARY CONIDIA.
When cultures are made with either the primary or sec-
ondary conidia on pine decoction agar, the spores generally
germinate ina few hours, and in two or more days colonies
may be noted with a white, sparse mycelium. At this
stage of growth, secondary conidia appear of the Sporotri-
chum-type, borne on erect, simple hyphae (pl. 8, f. 5}.
These are hyaline, and oval in shape. After a few days,
portions of the mycelium assume a brownish color, and
from certain matted masses of filaments or stromata there
grow out tufts of erect, parallel hyphae (pl. 8, f. 6).
Re ae gee,
ah f ‘
a as pie I See
PND Ee geglet
TA aay inc hae ban ge ain Valk paeiaci tall Pete N SR RY Meee ke ete Boy ee
Ree Rae = J Fo EOE ys Sela Rag alone = PCIE Caste GET OLAS NS phe ice , ‘i fi :
Oxy 3 = Sale mess ' \ f ;
86 MISSOURI BOTANICAL GARDEN.
These tufts of hyphae elongate into a long, round stalk,
terminating in a bunch of either verticillate or alternate
branches.
PRIMARY CONIDIA.
The primary conidia are formed on the upper portion of
the stroma or head by an abstriction of the ends of the
hyphae (pl. 8, f. 7). They are formed continuously, and
as soon as mature, fall away into the mucous mass which
; surrounds the head. The mass is hyaline at first, chang-
ing, as it grows older, to a straw color, and finally toa
dark brown. The stalk is usually hyaline in the first
stages of its growth, but later becomes almost black at the
base, shading off to a light brown near the apex, with a
striated appearance. No chlamydospores or sclerotia are
4 formed. At first the stroma is simple, but by the addition
Be of branches from the base becomes gregarious (pl. 8,f. 4).
The following are the measurements of the mycelial
structures: the hyaline mycelium measures from 1.5 to
2.54 in diameter, averaging 24. The colored portion
averages in diameter about 4u. The secondary conidia
3 measure 3.54 to 4 in length, and 1.2 to 1.5u in width,
Po averaging 3.7¢ by 1.34. The primary conidia average 5u
a by 3. The stalks average in length from 500p to 900u,
: The head, including the mucous drop, measures on an
a average 200u. These do not agree with any previous de-
: scription of species of Graphium and the fungus is here
named Graphium ambrostigerum, owing to the relation of
ae the conidial stage to ambrosia beetles.
Cultures were made on a number of kinds of wood. The
fungus grew readily in the sapwood of pine, gum, oak and
ash, bearing in most cases numerous heads. Secondary
conidia are usually formed in abundance. Cultures on the
heartwood of these trees produce little or no growth of the
fungus. Light colored wood is stained a color varying
from a dirty gray to a light brown, or almost black.
CHROMOGENIC FUNGI WHICH DISCOLOR woop. 87
The mycelium penetrates pine and other similar woods
through the medullary rays, dissolving the starch. In
gum wood, Liguidambar Styraciflua Linn., it penetrates
in addition many of the vessels of the wood. As compared
with Ceratosomella the penetration is not so rapid, neither
isthe stain so intense; but on the other hand it may. in
time stain wood as deeply.
Graphium eumorphum Sace.
A species of Graphium has been found frequenting the
old wood of Rubus strigosus and related species. This
was isolated and grown first-on wood of several kinds which
it was found to stain lightly, then upon agar media. An
examination of the cultural characters of the fungus place
it readily under Graphium eumorphum Sacc. The follow-
ing description gives the principal characters, from both
natural and artificial cultures :— .
MYCELIUM.
Cultures made from both the primary and the secondary
conidia preduce a sparse, white mycelium, which soon
turns a gray-green, and whose filaments measure from lp
to 2u in diameter. These soon bear numerous conidia of
the Sporotrichum-type (pl. 7, f. 2), and about a week
later the stromata or stked heads of Graphium appear
(pl. 7, £. 1). The heads are white, changing to a green
or greenish yellow, and measure with the mucilage drop
from 30p to 100u. The stalks vary from a yellow to a
dark olive or a brown color, and measure from 3800p to
500p in length, and from 10, to 40» in diameter.
SECONDARY CONIDIA.
The secondary conidia are obovate to elliptical in shape,
and average 7.84 by 3.4m (pl. 7, f. 3). They are borne, '
88 MISSOURI BOTANICAL GARDEN.
y unlike Sporotrichum, on simple erect not prostrate hyphae.
This stage nevertheless is apparently identical with Sporo-
trichum vellereum Sacc. and Speg. var. grisea Boul.
a - 2
PRIMARY CONIDIA.
TT? ey ee, Oe eee Se ree
The primary conidia are borne on branching hyphae in
stalked heads (pl. 7, f. 5), and average 7.74 by 3.4.
They are of a light green or yellow tint, and are identical
in shape and appearance with the secondary conidia.
ee ees
Graphium atrovirens n. sp.
C% In addition to Graphium smaragdinum on the red gum
wood a new species was found present in a number of
instances. This discolored the wood with a brown stain,
; similar to the latter species, and at first was not distin-
| guished from it, having the same color and appearance in
mass; but after a more careful study of the colonies on
agar, different forms of secondary conidia were found
pres“nt in two types of colonies, making it necessary to
separate this form from G. smaragdinum and giving good
* reasons to call it a different species. It is now named
| Graphium atrovirens with the following as its cultural
characters :—
MYCELIUM.
The growth of the colonies on agar plates is fluffy, as
compared with that of G. smaragdinum. This is due to the
formation of strands in the mycelium by the union of a
‘a number of parallel filaments, these growing upward and
often forming large branching masses, similar to those
formed by the mycelium of some species of Cerastosto-
mella. The filaments of the mycelium are hyaline at first,
later changing to dark green or olive, in the vicinity of the
stromata or heads. They measure 3u to 4” in diameter.
Those which unite to form the stroma usually measure but
FOOT Ra, a EPP PRP ME MOR eS Ree is ee AN Oa wy ae gt ge. cig oe 7 ron ee ee
CHROMOGENIC FUNGI WHICH DISCOLOR woop. 89
2u in diameter. The color of the stroma is dark green to
black when mature, but in growing colonies all variations
between hyaline and dark green may be found. The
stalks measure from 1.5 mm, to 3 mm. in height, and 8u
to 80u in diameter, being proportionately more slender
than those of every other species of Graphium described
inthis paper (pl. 8,f. 1). In this species, as in others, all
the forms of gradation between a head with a stalk com-
posed of one filament, and a stalk with many filaments may
be found. The difference between those bearing secondary
conidia and those bearing primary conidia consists mainly
in the absence of color in hyphae bearing the former.
SECONDARY CONIDIA.
The secondary conidia are borne in simple open clusters,
of the type of the conidia of Sporotrichum. They are
borne either terminally or along the sides of lengthened
hyphae (pl. 8, f. 2, 3). They are obovate to elliptical in
shape, measuring 42 to 5.54 by 1.6 to Qu. They are
hyaline, becoming guttulate when old. The clusters re-
main open for a number of days after they are mature, but
under very moist conditions fall together in rounded masses
on the hyphae. The conidia of G'. smaragdinum fall to-
gether as soon as mature, the clusters being much closer in
their formation (pl. 9, f. 9).
PRIMARY CONIDIA.
The primary conidia are borne terminally on the ends of
the branching hyphae in the stroma which form large,
mucilaginous heads. These, without the sheath of mucus,
are flattened oval in shape, (pl. 8, f. 1), white at first,
later gray in color. They measure 40, to 600, in their
greatest diameter, the latter being the measure of the
head including the mucilaginous sheath. The primary
conidia are hyaline, obovate to elliptical, and measure
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90 MISSOURI. BOTANICAL GARDEN.
8.52 to 4.54 by 1.44 to 24. The stalks in this species
bearing the primary conidia are sometimes much branch-
ed, but this feature is dependent more upon the culture
medium and physical conditions than upon the inherent
specific tendencies. One difference between the stalks of
this species and those of G. smaragdinum is that in the
case of the latter they tend to be gregarious, and are more
apt to be swollen at the base.
B. Species oF GRAPHIUM WITH A SECONDARY CONIDIAL
STAGE UNLIKE SPOROTRICHUM.
1. Species with secondary conidia borne continuously and
terminally, falling at once into clusters.
Under this subdivision will fall at least two species,
Graphium smaragdinum and G. rigidum.
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Plate 26. — Photomicrographs showing the cross sections of wood
taken at points immediately below the bands of the bagworm. The
bands were situated at the top of both figures. Upper figure, from
a maple constriction. Lower figure, from a red gum constriction.
>< 50.
DIAGRAMS.
Diagram 1.— Showing the number of atmospheres pressure exerted
by the bands of the bag worm, and the frequency with which the vari-
ous pressures are exerted among 413 bands. The abscissae show the
number of atmospheres pressure; the ordinates indicate the number of
individual bands exerting any given pressure.
Diagram 2.—a,. The relation between the varying widths of bands
and the strength of the bands, expressed in terms of number of atmos-
pheres presssure. The abscissae represent the number of atmospheres;
the ordinates the number of individuals. —b. The relation between the
varying diameters of twigs and the strength of the bands found en-
circling them, expressed in terms of number of atmospheres pressure.
The abscissae represent the number of atmospheres; the ordinates, the
number of individuals.
TEXT FIGURES.
Figure 1. — Longitudinal section through an old swelling of the arbor
vitae ( Thuja occidentalis), showing how the upper or outer portion grew
in diameter, while the part below the band stopped growing entirely.
(P. 189).
Figures 2-4. — Longitudinal sections through constrictions of the soft
maple, caused by the band of the bag worm. The figures show succes-
sive stages of healing. Figures 2 and 3 show the top of the bag at the
left of the constriction. (P. 160, 161).
TEXT TABLES.
Table I.—Showing the relation of the diameter of maple twigs to the
width of the bands of the bag worm which were found upon them.
(P. 176).
Table II. — Showing a number of exceptionally strong bands, their
dimensions and the trees they were found on. (P. 178).
a
REptT. Mo. Bot, GARD., VOL. 17.
PLATE 20.
CONSTRICTIONS OF ARBOR VITAE.
REPT. MO. BOT. GARD., VOL. 17.
PLATE 21,
BAG WORMS— FIRST YEAR.
PLATE 22.
Rept. Mo. Bot, GARD., VOL. 17.
CONSTRICTIONS OF SOFT MAPLE.
REPT. MO. BOT. GARD., VOL. 17, PLATE 23.
CONSTRICTIONS OF HARDWOODS.
REP?T. Mo. Bot, GARD., VOL. 17. PLATE 24.
CONSTRICTIONS OF CONIFERS.
- i i eo ce ae
PLATE 26.
REPT. MO. BOT. GARD., VOL. 17.
WOOD BELOW BANDS.
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