MISSOURI

BOTANICAL GARDEN.

SIXTH ANNUAL REPORT.

ST. LOUIS, MO.:
PUBLISHED BY THE BOARD OF TRUSTEES.
1895.

BOARD OF TRUSTEES OF THE MISSOURI
BOTANICAL GARDEN.

President,
RUFUS J. LACKLAND.

Vice-President,
HENRY HITCHCOCK, LL.D.

JOSEPH W. BRANCH. FREDERICK W. BROCKMAN,
President of the Board of Publie
GEORGE S. DRAKE. Schools of St. Louis.*
WINFIELD S. CHAPLIN,
Chancellor of Washington Univer-
sity.*

GEORGE J. ENGELMANN, M. D.

JOHN B. JOHNSON, M. D.
JOHN GREEN, M. D.,}
DAVID F. KAIME. President of the Academy of Science
ot St. Louis.*

GEORGE A. MADILL. DANIEL 8. TUTTLE,

i *
Wiittux H. H. Parrvs. Bishop of the Diocese of Missouri.

Cyrus P. WALBRIDGE,
JAMES E. YEATMAN. Mayor of the City of St. Louis.*

A. D. CUNNINGHAM, Secretary.

* Ex officio.

1 Elected President of the Academy of Science January 7th, 1895, in place of
Henry 8. Pritchett, who had met with the Board for three years prior to that date.

PREFACE.

Under direction of the Board of Trustees, the sixth
annual report of the Missouri Botanical Garden is offered
to the public. This volume differs from those which pre-
ceded it in the omission of the section consisting of
‘‘ anniversary publications,’’ comprising the annual flower
sermon and the proceedings at the two annual banquets
instituted by Mr. Shaw, the Director’s report containing
all that it is considered necessary to state concerning these.

The fifth report was issued April 27th, 1894. Reprints
of papers from the present volume were issued in advance
of its publication as follows: —

Smith, Sagittaria and Lophotocarpus, May 24th, 1894;
Trelease, Leitneria Floridana, May 30th, 1894.

The reports of the Garden are sent regularly to scientific
institutions and journals in exchange for publications or
specimens desirable for the library and herbarium of the
Garden, and, so far as is possible, reprints of the botanical
articles which they contain are sent to botanists to whom
these papers are directly useful.

Regular agents for the sale of Garden publications are
Dr. A. E. Foote, of Philadelphia, W. Wesley & Son, of
London, and R. Friedlander & Sohn, of Berlin.

WiLuiaM TRELEASE.

St. Louis, March 1, 1895.

(3)

CONTENTS.

PAGE.

1. REPORTS FOR THE YEAR 1894:—
a. Report of the Officers of the Board ake 7
b. Sixth Annual Report of the Director. . ... . rt

2. SCIENTIFIC PAPERS :—
a. Revision of the North American Species of Sagittaria and
Lophotocarpus —
By Jared G. Smith .
. Leitneria Floridana —
By William Trelease « fg ete eee

c. Studies on the Dissemination and Leaf Reflexion of Yucca

aloifolia and other Species —
pereeroert 3. Webber... s,s sh ve wate re ee
ad, Notes and Observations on new or little known Species —
By Jared G. Smith . <a ee ee
e. Notes on the Mound Flora of Atchison County, Missouri—
ee

oe

(5)

27

65

91

. 113

. 121

LIST OF ILLUSTRATIONS.

Among the Willows ... .. +. +. « « « « Frontispiece.

Yucca glauca Nutt. .. . +... . + © » « « Facingpage 7
COOCMRIA ANUIGUOFUIN 5. 58. ee, se ee ig 11
Me Aaa as 8 es i ee ae a) ee
Opuntias, inthe Cactus House. . ...... = 7 23
Leitneria Floridana, inthe Bog ...... « « ds ies 65
Plates 1-29, Species of Sagittaria . . . . . . Following page 64

‘¢ 630-44, Leitmeria Floridana . . ..... . ” ™ 90

“ 46-47, Yucca aloifolia, etc... . . « « « s ae os TS

48, Sida Palmeri?

49, Fugosia Drummondii,

50, Sedum Texanum,

51, Elytraria and Stenandrium,

52, Zephyranthes,

53, Sagittaria isoetiformis,

54, Eragrostis sporoboloides,

55, Eragrostis grandiflora,

56, Eragrostis Beyrichii . . . . . « « » » Ld “ 120

(6)

‘ELAN ‘'vonWIS Voons

REPORTS FOR THE YEAR 1894.

REPORT OF THE OFFICERS OF THE BOARD.
SUBMITTED TO THE TRUSTEES JANUARY 9TH, 1895.

To the Board of Trustees of the Missouri Botanical Garden:

The financial results for the past year have been very
satisfactory considering the depressed condition of trade.
Although some of the properties of the Board have been
vacant all or a portion of the year, the receipts from
rentals have exceeded those of the previous year over
$1,000.00, and at this date, with fewer vacancies, the
income will undoubtedly be in excess of that of the past
year several thousand dollars.

The properties of the Board have been kept in good
repair and nearly all are occupied by desirable tenants at
fair rentals.

No extensive improvements have been made at the
Garden, but the addition of a plant-house costing about
$2,700.00 and several granitoid ponds for the growing of
the Victoria Regia and other lilies have added greatly to
the beauty and attractiveness of the Garden during the
summer months, as well as offering additional attractions
in the way of blooming plants throughout the winter.

After carrying out Mr. Shaw’s bequests for annual
banquets, flower sermon, and premiums for the flower
show, caring for the Garden and paying all expenses of

(7)

8 MISSOURI BOTANICAL GARDEN.

caring for the properties of the Board, we are enabled to
carry forward a gain of $19,824.18 for the year.

Additions have also been made to the library, herbarium,
and furniture for the library building, and the following
sums have been credited to the stock account, which now
aggregates $1,480,886.18: —

Library (purchases and donations)......sssesseesceeeeceeees $4,361 71
Herbarium (purchases and donations)......+.+.eseeceveecees 908 70
Missouri Botanical Garden permanent improvements........ 3,887 53

$9,157 94

For a full and detailed account of the receipts and
expenditures you are referred to the following state-

ment: —
RECEIPTS.

WRU Wi od eo basen Ahad kd nines eee eee eeeee eeeeee $93,252 87

Ptwek Account salopes ss. seis ces iose es Vive was tee 82 10

Garden pasturage and sales............s0seeeee ws 684 30

Interest and cash discount on taxes.........++.++ 1,859 70

Garden hand-book sales........ccseceseccecsece 71 00
PUDUCHUOMG GMIEG . iin cso cecct cnecesvbcesvescecs 86 50

Loss by fire to buildings.........0-+eeeesees. . 119 50 $95,555 97
Cash balance January Ist, 1894..........0...6 . 15,649 75

OO CONS CANES 6 ONES 06 'b 00 bbs MR $111,205 72

REPORT OF THE OFFICERS OF THE BOARD. 9

EXPENDITURES.
Garden Account —
Labor, including garden pupils....+.+++++eeees
WUGh cs css pec css cecccgoewctovescecsecceectens
Stable and implements... ..sesecee cess seeevee
Repairs and supplieS..-+eeceesseeeee coeeceeee
Scholarship, care of lodge and supplies.......
Plants and S@eds...-sccee cccceccsereecccceacs

Herbarium. ...ee ecccceccesceccecs e peccevesess
Library, books, subscriptions, etc....+++eees+s
Garden office, salaries, supplies, etc..........-.

PROGGUTON cclcc 040.8 0 dvceb boss $660 b6:n.6 oclceenaee

Garden Improvements Account —
Greenhouse, buildings, etC.....- ss. seeeeeees °

Publication Account —
Annual VOlUME.....cccccccerscsccerccsccsceee
Property Expenses —
Commission for collecting rents....+++++++++
Taxes, State, school, city and sprinkling..... .
Streets, pavements and SCWETS.-+-+++se+eeeees
Insurance,...++.+6-. Soe HES GO none oo.
Repairs ..ceeeccccccccccccccccsecccres cece aie

Office Account —
IGN) chee Ge vob kes piccceccocensscesseess
Pe OE CG 6 60 nc ccc ccc cs cccces ves sece
Printing, postage, telephone and furnishing...

Sundry Accounts —
Legal expenses . eee ss vine “Petes.
Repairs to buildings deeked by leben. slsaes eves

Bequests —
Premiums to flower show*........ ee ee
Flower sermon...... PEP oe ene ees 's Kebnah seek
Trustees’ banquet..+...ecee cece seer reece ee,
Gardeners’ ida pie tives bac sot ¥04-rbiwy

Washington Tatversite ( School of Stans eves

TUWRUGEL koe cc cceee e@eeeeneeeeevee e@eereeeeeeee ee
Cash on hand December 31st, 1894.... +--+ eeeeee

Total... eeeeseseanes eeeree ee eeesaeeses

$14,096 62

1,582 87
604 44
1,849 37
767 44
284 08
860 51
2,614 90
5,362 54
475 50

5,185 25

1,849 87

1,070 45
20,980 86
353 99
3,977 438

5,041 62

2,821 38
815 00

1,872 18

72 65
119 50

577 00
200 00
905 10
336 59

1,678 60

$28,498 27

5,135 25

1,849 87

31,424 35

5,008 51

192 15

8,692 29

15,000 00
20,405 08

$111,205 72

* Of this sum $100 (toward cost of medal) was properly chargeable to
the year 1893, in which only $400 appears as having been paid for

premiums — see Fifth Report, p. 9.

10 MISSOURI BOTANICAL GARDEN.

The books of the Board have been closed after showing
the operations for the year ending December 31, 1894,
and the receipts have been disposed of as follows: —

Rent account... .eeee cece sccceccecccseevcees erry
Interest Peters reser sreeseees se eeeee nesses reeseeses
Garden Ts 's'v 0-0-0 06408 6 cb 05 es Coes eave

CONTRA.

$93,252 87
1,359 70
71 00

$94,683 57

Garden CXPeCNsSe..osserececccerecceccecsessesses $20;069 98
Office expense ......eeeeeeeee ce cccceree coesccecee 5,008 51
Commission ...--ccrccsccecccccercecscccsecceses 1,070 45
Repairs. .cccaccccccccccccccccsecsccssccescccsce §,041 62
THAUPANCE. 00 bse cee ses Fe ee Pees ee coovceee 8,977 48 .
TAXCS.- cece cccesessscceecccsesece socccccccesese 20,980 86
Street pavements and sewers.........- Coveccccee 853 99
Legal CxpenseS.....esseecessececccececccecevecs 72 65
Washington University (School of Botany)..... 1,673 60
Annual flower SETMON....-ececcecceccesecccrece 200 00
Annual Trustees’ banquet........se.eeeeee bivees 905 10
Annual Gardeners’ banquet.........escccccesces 8336 59
Annual flower ShHOW...-+++seeecesees 455946505 he 577 00
PODMOMIOES ceuese 58 Cease’ aehab cea W eee hs iver 1,813 37
Garden improvements......... seccccccccsccsces 4,778 24
Surplus for 1894. .0.sesccccerceccecceseccececess 19,824 18
$94,683 57 $94,683 57

Surplus Dec. 31, 1893.....0.-seeeeccecce cence rece $40,649 75
Surplus for 1894....ccccccccccccsccesccesccccess 19,824 18

Total surplus December 31, 1894.......... $60,473 93

Respectfully submitted,
R. J. LACKLAND, President.

Attest,

A. D. CUNNINGHAM, Secretary.

‘NOUYONOILNV VISVOO'TOO

SIXTH ANNUAL REPORT OF THE DIRECTOR.
SUBMITTED TO THE TRUSTEES JAN. 9, 1895.

To the Board of Trustees of the Missouri Botanical Garden:

The following report on the Missouri Botanical Garden
and the Henry Shaw School of Botany is respectfully sub-
mitted, in compliance with the rules of the Board :—

THE BOTANICAL GARDEN.

During the past year the number of visitors to the Gar-
den is reported by the gate-keeper to have been somewhat
greater than in 1893, and the Head Gardener, who has
been familiar with the Garden for many years, estimates
that about one-third more persons now visit the grounds
than was the case a few years ago, — an increase partly
due to the increased facilities which now exist for reaching
the Garden, and, perhaps, in part to the attractions which
have been added during the last few years. On the open
Sunday afternoon in June, 20,159 visitors were noted, and
on the open Sunday in September, which was showery,
about 15,500. As in previous years, the visitors on these
Sundays were orderly and apparently well disposed. Dur-
ing the year, visitors have purchased 256 copies of the
little handbook of the Garden offered for sale at the gate.

In the second paragraph of the first clause of his will,*
the founder of the Garden directs that the Garden shall
be kept open during such hours, and under such regula-
tions as the Trustees shall prescribe, ‘‘ every day except
Sundays, for the use of the public at large.’’ In the
thirty-fourth paragraph of the fourth clause of the same
instrument,t he adds: ‘‘ In the first clause, and second

* First Garden Report, p. 32.
tlic. p. 51.

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12 MISSOURI BOTANICAL GARDEN.

paragraph of this my last will and testament, I devise and
express my wish that the garden shall be kept open, under
necessary regulations, Sundays and holidays excepted,
every day of the week. Now as this Trust is made for
the use of the citizens of St. Louis as well as the
public in general; it is my wish that for the conven-
ience of said citizens and public, that the Garden shall
be open to visitors two Sundays in each year, viz, the
first Sundays in June, and the first Sunday in September,
from Two P M tosunset.’’ During the later years of Mr.
Shaw’s life, it was his custom to close the Garden on Sun-
day, but for the convenience of strangers in the city unable
to visit the Garden on other days, he issued a limited number
of Sunday cards, each admitting from one to four persons,
and also allowed the occasional visits of a larger number.
Such permits have been given in a few instances since the
death of Mr. Shaw, but it has always been questionable
whether they were strictly possible under the power con-
ferred on the Trustees by Mr. Shaw’s will, which by impli-
cation directs that the public as a whole and also individuals
shall be refused admittance on Sundays and holidays; and,
however much they may regret the necessity for so doing,
the Board of Trustees have decided that under the instruc-
tions contained in the will they have no option except to
close the Garden on Sundays and holidays (except the Sun-
day afternoons mentioned above ), to individual visitors as
well as to the public as a whole.

The general decorative features of the grounds were main-
tained through 1894 on substantially the same lines as in
the preceding year, though somewhat more color was intro-
duced, especially near the main entrance. A collection of
economic fiber plants, in the northwestern part of the Garden
proper, was made a special feature this year. Although
for a time during the extreme drought of the summer, the
lawns suffered greatly, the Garden has been kept in a fairly
attractive condition through the season. The average num-
ber of laborers (including seven garden pupils) was 37, and

SIXTH ANNUAL REPORT OF THE DIRECTOR. 13

the pay-roll for the year aggregates $14,096.62, of which a
considerable sum was expended for necessary repairs to
greenhouses and other structures.

Some years ago an effort was made to cultivate under-
shrubs and other small plants at suitable points in the
arboretum, by the establishment there of a wild garden.*
Owing to the severity of the seasons and the extent to
which the closely set trees of the arboretum have taken
possession of the soil, this feature has not added to the
attractiveness of the Garden as largely as was hoped, but
there is seldom a time during the open season when the wild
garden and bog do not contain a considerable number of
interesting plants in bloom, and it is hoped that their suc-
cess may be increased in the future. The droughts which
have prevailed during the past five summers, and the
impervious subsoil, have continued to weaken the older
trees, especially conifers, so that, as was stated a year ago,
it is probable that few of the latter will survive many years
longer; and a very severe frost, coming when the earlier
species were beginning their vegetation last spring, killed
outright many trees, such as the weeping willows, while it
paved the way for further harm during the exceptionally
hot and dry summer which followed. Where they are
needed, other trees are being planted to make good the
losses from these causes, but it is difficult to successfully
grow a new tree on the spot from which an old one has been
removed, and a more general clearing and tasty replanting
will become necessary within a few years.

The most generally attractive features added to the
grounds this year are a series of new granitoid lily ponds,
one of which, situated south of the Linnean house, was
planned for the growth of the Victoria Regia, or Amazon
water lily. The decorative effect of these ponds was very
considerable, and they were the source of favorable com-
ment by the press of the city and by visitors. A much

* Third Report, p. 13.

14 MISSOURI BOTANICAL GARDEN.

needed plant house, 21 X 97 feet, was erected in the sum-
mer, just south of the Victoria pond, and is adding greatly
to the attractiveness of the Garden through the winter.
Numerous minor improvements have been reported to the
Board in detail, month by month.

The fruticetum, reported on at various times in the past,*
has been further improved this year by the removal of all
of the old and worthless apple trees and grape vines and
the laying of 1,630 feet of drain tile, and is to be largely
replanted in the spring to carefully selected varieties of
fruit. The appearance of this inclosure has been greatly
enhanced by the replacement of the dilapidated picket
fence separating it from the arboretum, by a neat open
wire fence. It is hoped that these horticultural improve-
ments may be extended by the erection in the near future
of a much needed vegetable forcing house, with a grapery
compartment.

During the year, some 950 packets of seeds and 1,000
plants have been received by way of donation or exchange,
and 850 packets of seeds and a few dozen plants distributed.
The largest of the accessions consisted in 952 plants, many
of them of great value, received from South Park, of
Chicago. A limited number, aside from bulbs and other
transient plants, have also been added to the Garden by
purchase. Some 26,900 bedding plants were propagated
at the Garden for out-of-door decoration, — an increase of
some 6,000 as compared with the preceding year. On the
approach of winter, some 1,100 of these were taken from
the ground and potted, and distributed to charitable institu-
tions and poor homes in the city, through the kindness of
the Bethel Association, to whose officers I wish to express
my indebtedness.

While much remains to be desired, the labeling of the
plants of the Garden is being greatly improved each year,
and [ hope that it will not be long before every plant will

* Garden Reports, ii. 18-20, 30; iii. 12.

‘aT UIVOTIONT V

pe ee

SIXTH ANNUAL REPORT OF THE DIRECTOR. 15

be correctly and legibly named. The enamel (granite
ware) tree labels which were at first tried have not proved
satisfactory, for several reasons, and trees of sufficient size
are now receiving zinc alloy labels cast with raised letters,
and affixed by two pins in a vertical line, so as to admit of
the expansion of the tree during growth. For the smaller
plants, celluloid labels, lettered with a special ink, have
been employed very largely, and present a neat appearance
when wired to the plants or fastened before them; but it
has thus far proved to be impossible to letter them so as to
insure permanency, and experiments are being tried in the
hope of securing labels combining durability and attractive-
ness with reasonable cost.

The herbarium has been increased by the incorporation
of 9,307 sheets of specimens, of which 3,567 sheets were
purchased, 126 belonged to the Bernhardi herbarium, and
5,614 were received by donation or exchange. 208 mounted
sheets of duplicates from the Engelmann herbarium, and
12 mounted duplicates from the general herbarium, besides
about 1,000 unmounted duplicates, have been distributed
to correspondents by way of exchange. As now consti-
tuted, the herbarium contains the following collections :—

The Engelmann herbarium (all groups), about........ 97,800 specimens.
The general herbarium of higher plants:
The Bernhardi herbarium.........eeesseeses 61,120
Other SPpeCiMEDS.....- ceeccecccceecceecececs 54,687
115,807“
The collection of Thallophytes :—
The Bernhardi herbarium.........+ssseseeses 126
Other specimens.......cecccsscccccccccccvce 17,794
17,920 «
Making a total of about.... ..-.--.eseeeeeeeeee 231,527 a

During the year $1,720.11 was spent for purchases and
binding for the library, the additions to which consist of
873 books and 166 pamphlets purchased, and 379 books,
valued at $733.95, and 999 pamphlets, valued at $207.65,

16 MISSOURI BOTANICAL GARDEN.

donated or received by way of exchange. Sixty large vol-
umes of Dr. Engelmann’s manuscript notes and sketches
are appraised at $600.00. An indexer has been kept
steadily employed during the greater part of the year on
the card index of plant illustrations in the library; and the
card index of Experiment Station literature published by
the United States Department of Agriculture, and a set of
index cards referring to recently published plant names,
have been added by purchase. It is estimated that the
card indexes at the library now number about 110,000
cards appraised at $1,100.00, exclusive of the authors’
catalogue of books and pamphlets. Exclusive of the Stur-
tevant prelinnean collection of about 460 volumes, which
has not yet been appraised and added to the capital stock
account, the library now contains :—

Books...... 7,631
Pamphlets.. 9,822

Together. 17,458 works, and 110,000 index cards,valued at $29,630.23.

I have been assisted in office and similar work by Mr. J.
C. Whitten, Horticultural Assistant, Jared G. Smith,
Botanical Assistant, Miss Grace E. Johnson, Artist, Eva
M. Reed, Indexer, and C. E. Hutchings, Amanuensis. In
October, Mr. Whitten left the Garden to accept the pro-
fessorship of horticulture at the Missouri State University,
and his place has been filled by the appointment of Mr.
H. C. Irish, of the South Dakota Agricultural Col-
lege. In addition to the necessary routine work, time
has been found for a certain amount of research work
by Mr. Smith and myself, and phenological notes were
also made through the year by Mr. Whitten, and by garden
pupils under his direction. Several papers embodying the
results of such work are now in preparation. The Garden
table at the Wood’s Holl Marine Biological Laboratory
was again used this year by Mr. M. A. Brannon, who in-
forms me that his studies on Grinnellia are now so far

a

We ea gage tng oh A lag. irae ee aa
* RAE OR eR NES

SIXTH ANNUAL REPORT OF THE DIRECTOR. 17
completed that they will soon be ready for publication.
Through the summer, I utilized a four months’ leave of
absence by visiting the Azores, where a large collection
was made, quite fully representing the flora of these
islands, and adding somewhat to what was known of the
distribution of species through the groups of islands.
This collection is now being worked over by me, and may,
perhaps, form the subject for a paper to be published in
the seventh Garden Report. Mr. Smith is at present
devoting a portion of his time to a study of the Capsicums
which have been cultivated and collected during the past
three years, and toward which study Dr. E. Lewis Sturte-
vant contributed his valuable sets of notes, drawings, and
specimens in 1892.*

Four annual events have taken place in the manner
directed in Henry Shaw’s will: namely, the preaching of a
sermon on the wisdom and goodness of God as shown in
the growth of flowers, fruits, and other products of the
vegetable kingdom; the fifth banquet to the Trustees of
the Garden and their invited guests; the fifth banquet to
the gardeners of the institution and invited florists, nur-
serymen and market gardeners; and the award of pre-
miums or prizes to a flower show or exhibition.

The flower sermon was preached in Christ Church Cathe-
dral, St. Louis, on the morning of May 13, by Right
Reverend T. F. Gailor, Assistant Bishop of Tennessse.

The Trustees’ banquet was given at the Mercantile Club
of St. Louis, on the night of May 18, and was presided
over by Professor H. 8. Pritchett, President of the Acad-
emy of Science of St. Louis. Covers were laid for about
seventy-five persons. Among those present were: —

J. C. Arthur, Shepard Barclay,

Purdue University. Jefferson City, Mo.
O. D. Ashley, C. C. Hemenway,

New York. Pritchett Institute.

* Fourth Garden Report, p. 15.

18 MISSOURI BOTANICAL GARDEN.

F. C, Hicks,
University of Missouri.
R. H. Jesse,
University of Missouri.
T. H. McBride,
University of Iowa.
Conway MacMillan.
University of Minnesota.
Oren Root,
Hamilton College.
E. M. Shepard,
Drury College.
W. von Streeruwitz,
Austin, Tex.
F. E. Nipher,
C. R. Sanger,
E. H. Sears,
M. S. Snow,
of Washington University.
George D. Barnard,
Adolphus Boeckeler,
W. W. Boyd,
J. P. Bryson,
M. Dwight Collier,
F. M. Crunden,
Thomas Dimmock,
Daniel Dillon,
Daniel D. Fisher,
S. W. Fordyce,
D. R. Francis,
Fred. Gabel,
W.C. Glasgow,
C. §. Greeley,
John Green,
Henry C. Haarstick,
James Hagerman,
Edwin Harrison,
Charles M. Hays,
E. A. Hitchcock,
W. L. Huse,
F. L. James,
J. E. Kaime,

Henry King,
Jacob Klein,
W. H. Lee,
J. M. Leete,
Geo. E. Leighton,
R. E. MacMath,
W. C. Marshall,
Albert Merrill,
N. O. Nelson,
Byron Nugent,
Charles Parsons,
Julius Pitzman,
Thomas C. Purdy,
P. G. Robinson,
R. E. Rombauer,
O. U. von Schrader,
W. L. Sheldon,
Wm. Short,
F. L. Soldan,
Charles Speck,
E. O. Stanard,
S. 8. Sumner,
Edwards Whitaker,
O. L. Whitelaw,
W. G. Williams,
James Withrow,

of St. Louis.
F. W. Brockman,
W.S. Chaplin,
Geo. S. Drake,
Henry Hitchcock,
D. F. Kaime,
R. J. Lackland,
H. 8. Pritchett,
D. 8S. Tuttle,
Jas. E. Yeatman,

Trustees of the Garden.
A. D. Cunningham,

Secretary Board of Trustees.

Wm. Trelease,
Director of the Garden.

After the dinner, toasts appropriate to the occasion were
gracefully proposed by the chairman, and responded to
fittingly by President R. H. Jesse, of the University of

SIXTH ANNUAL REPORT OF THE DIRECTOR.

19

Missouri, Professor Oren Root, of Hamilton College,
President O. D. Ashley, of the Wabash Railway, and Dr.
John Green and the Reverend William Short, of St. Louis.

The gardeners’ banquet was given at the Mercantile Club
of St. Louis, on the night of November 5, and was presided

over by the Director of the Garden.

About seventy-five

guests were present, among them: —

A. G. Fehr, Belleville, Ill.

E. W. Guy, Belleville, Ml.

A. §. Halstead, Belleville, Ill.
G. E. Meisner, Bushberg, Mo.
A. Nelson, Lebanon, Mo.

E. A. Riehl, Alton, Il.
Homer Riggle, Columbus, O.
QO. C. Simonds, Chicago, II.
J. F. Stinson, Fayetteville, Ark.
A. L. Vaughan, Chicago, Il.
G. A. Washburn, Bloomington, Ill.
J. C. Whitten, Columbia, Mo.
R. W. Ayres,

J. J. Beneke,

H. W. Chandler, Jr.,

N. J. Colman,

C. L. Connon,

Chas, Connon,

J. M. Connon,

J. W. Dunford, Jr.,

Wn. Ellison,

F, J. Fillmore,

R. Frow,

Wm. Hackman,

J. C. Jannopoulo,

J. M. Jordan,

C. A. Juengel,

Wm. Klockenkemper,

John Koenig,

Julius Koenig,

C. A. Kuehn,

Geo. Longman,

Leonard Matthews,

Fred Meister,

E. H. Michel,

F. W. Ostertag,

Henry Ostertag,

Wm. Pape,

Julius Pitzman,
C. C. Sanders,
Carew Sanders,
Wm. Schray,
J. W. Schuette,
R. F. Tesson,
A. Waldbart,
N. J. Wellhouse,
J. F. Windt,
C. W. Wors,
Harry Young,
of St. Louis.
J. W. Branch,
F. W. Brockman,
R. J. Lackland,
D. S. Tuttle,
of the Board of Trustees,
A. D. Cunningham,
Secretary of the Board.
O. L. Simmons,
C. H. Thompson,
of the School of Botany.
Thomas Doss,
J. W. Dunford,
Philip Giebel,
Jas. Gurney,
C. E. Hutchings,
H. C. Irish,
C. I. Paige,
J. M. Paige,
J. P. Pillsbury,
Chas. Schmidt,
J. G. Smith,
Geo. Stockey,
Wm. Trelease,
Mike Zavadil,
of the Botanical Garden.

20 MISSOURI BOTANICAL GARDEN.

In response to toasts proposed by the chairman, short
and apt speeches were made by Mr. O. C. Simonds, of
Chicago, President of the American Association of Ceme-
tery Superintendents, Professor J. C. Whitten, of the Uni-
versity of Missouri, President F. W. Brockman of the St.
Louis School Board, and Messrs. Leonard Matthews, Julius
Koenig, and Julius Pitzman, of St. Louis.

The award of flower premiums for 1894 was again in-
trusted to the Florists’ Club of St. Louis, for the benefit
of a chrysanthemum show held in the Exposition Building,
St. Louis, November 6 to 9. Some of the premiums were
not competed for, and were consequently not awarded, and
the exhibition judges decided that none of the plants com-
peting for the gold medal founded in 1893 were sufiiciently
meritorious to justify the award of the medal.

The object of Mr. Shaw in providing for ‘* premiums or
prizes to a flower show or exhibition * * * established
by amateurs and horticulturists of St. Louis,’’ evidently
was to stimulate public interest in floriculture, to improve
and increase the variety of plants used in St. Louis, and to
stimulate and assist florists in their efforts to create and
supply a demand for decorative plants. In approving the
suggestions of the Florists’ Club for a list of the Shaw
premiums each year, an effort has therefore been made so
to shape the list as to add to the general attractiveness of
the exhibition and at the same time also to familiarize the
public with new or easily cultivated plants not usually to be
seen in the local florists’ windows. It was in the hope of
bringing to these St. Louis exhibitions newly introduced
plants, as well as to stimulate the importation of such
plants, that in 1893 the Board of Trustees of the Garden
founded ‘‘ the Henry Shaw Medal for the introduction of a
valuable plant,’’ open to competition in any line of decora-
tive horticulture, but to be awarded only ‘ for a plant of
decided merit for cultivation, not previously an article of
North American commerce, and introduced to such com-
merce by the exhibitor during the year in which said award

SIXTH ANNUAL REPORT OF THE DIRECTOR. 21

is made,”’ * and it is hoped that importers and originators of

new plants of merit will be encouraged to compete for this
medal by the precedent established this year by the Florists’
Club in refusing an award except in case of very decided
merit, in conformity with the wishes of the Trustees.

Mr. Shaw’s expressed wish for the instruction of garden
pupils has this year borne fruit in the completion of the
gardening course by Mr. Homer Riggle, who was given
a scholarship in the spring of 1890, and who on the com-
pletion of his studies at the Garden, was appointed to a
gardener’s position at the Ohio State University; and Mr.
J. P. Pillsbury, who would have completed the course in
March, 1895, has been appointed assistant to the Horticult-
urist of the Pennsylvania State College, and will probably
apply for examination for a certificate in 1895, after com-
pleting the work required of garden pupils. :

While the number of free scholarships contemplated by
Mr. Shaw, and provided for in 1889, is limited to six, so
many applications have been received for the admission to
the garden classes of paying pupils, that at its meeting of
November 16th, 1894, the Board of Trustees authorized
the Director ‘‘to admit, in addition to those holding gar-
den scholarships, as many suitably prepared garden pupils
as can, in his judgment, be adequately taught without
material increase in the cost of tuition, each pupil so ad-
mitted to be charged $25.00 per year tuition, and to be
entitled to the same certificate as a scholarship pupil on
completion of the required course and examinations.’’ In
November, therefore, a sixth announcement was issued,
stating that the vacancy caused by the withdrawal of Mr.
Pillsbury would be filled in March next, by the appoint-
ment of a scholarship pupil, and that applications would
be considered from persons desirous of entering the Garden
next April as pupils without any of the scholarship grants
or payments, and subject to the payment of a tuition fee
of twenty-five dollars at the beginning of each class year.

To avoid unnecessary duplication of instruction both at

* Fifth Report, pp. 18, 19.

eae

es

22

MISSOURI BOTANICAL GARDEN.

the Garden and the School of Botany, the course of study
for garden pupils has been re-arranged as is shown in the
appended table, so that the work of the third and fourth
years may be given alternately to two combined classes, the
second year’s class work only being repeated each year.

COURSE OF STUDY.

YEAR. TERM. STUDIES. PER WEEK.
: Economic
April ——* Entomol- lh
o ogy. exercises
8 exercises]
June weekly. 1 a weekly.
Economic
July Floricul- Book-
to ture. — Keeping.
Sept. 8 ey 1
SECOND.
. | Economic Element-
= —— Entomol- Surveying Be,
< ogy. otany.
Dec. 1 2 1 3
Twigs of Element-
Jan. Floricul- Orchard |Landscape
Woody ary
fe ture. Plants. Culture. |Gardening Botany.
1 1 1 1 3
April | Vegetable Landscape re =e
to Gardening Gardening Botan
June y:
4 1 2
July Economic| Orchard |Landscape maton of
to Mycology.| Culture. (Gardening
Flowers.
Sept 1 2 1 2
THIRD.
Oct. Economic Botany of | Vegetable
to Mycology. Fruits. {Physiology
Dec 3 1
Jan. Economic Botany of | Vegetable
to Mycology. Weeds. |Physiology
Mar. 3 1 2
April | Orchard Book- | Surveying
Forestry. an
Culture. Keeping. | prainage
June. 1 1 1 3 Be.
Small
July Garden Botany of
to whines Accounts. Vegetables|
Sept ." 1 1
FOURTH.
Botany of
Oct Special Garden
to Gardening | FT€8try- | accounts. jee
Dec. 2 1 1 e
Botany of
Jan. Special Y Of! Botany of
to Gardening | Forestry. ponee Fouas.
Mar. 2 1 x 1

CACTUS HOUSE.

THE

OPUNTIAS IN

SIXTH ANNUAL REPORT OF THE DIRECTOR. 23

The 75 class exercises per week here tabulated (each
extending over three months), may be grouped under
subjects as follows :—

Gardening:
Floriculture. .... ccc cece cece ccccececcesccscccccscese 8
Vegetable gardening........-- ‘glee Vie'w'e bd § ORG OR TREE 4
Wyit. CULWAIG 6 000.6000 cece econ vececccccece seceeces anes 8
FOrestry 1.2 ccccce sees cece eeeree cece eee eeeeeeeeeees 3
Landscape gardening .---<sssseeeeerereeree cree cceees 3
Selected thesis WOrk....2.seesceecccececeescsvececens 4 380
Surveying and drainage.......sesseseeeceeeeeceeeeeeeeeee 6
Bookkeeping and accounts. ..--..-e rece rece rece cee eereee 4
Economic entomology ...---+++-+++eeeeeee Side odedecseeiss 5
Botany in its relation to gardening:
General botany.....escececese sees cenecnceereuse ar
Botany of decorative plantS...- ++... eereeerereces case
Botany of hardy woody plants...---+++++++esseee sees 3
Botany Of fruits..... 0.02 seee cece eee eee sete teen eeeee 2
Botany of vegetables. ......+eeeeee rece cece cece ee rene 1
Botany Of Weeds... see seee cece ceeece cere ceteee sees 1
Economic mycology ---.+ sees cece cece cece cece cece cees 7
Vegetable physiology...+ +--+ +++- sees ceceee sree ceees 3 30 75

Subjects capable of being taught in the laboratory, the
greenhouse, or the field, are so taught, and all of the theo-
retical instruction is expected to be practically tested in
the performance of manual labor, to which the entire first
year, and half of each day through the remainder of the
course, is devoted. Vacations are granted from July 1 to
15, and December 21 to January 4, inclusive.

THE SCHOOL OF BOTANY.

On the opening of the college year 1894-5, the instruc-
tional force of the School of Botany was increased by the
appointment of Mr. Orville L. Simmons as instructor in
cryptogamic botany, Mr. C. H. Thompson continuing to
act as general instructor. In the spring of 1894, a course
of Saturday lectures and demonstrations in pollination and
plant fertilization was given by me at the Garden to 33
persons, laboratory work was given to 3 teachers, and two

24 MISSOURI BOTANICAL GARDEN.

classes, including 45 children, were taught elementary
botany at the Garden by Miss A. I. Mulford, through the
same term. During the past autumn, Miss Mulford has
further conducted two classes, consisting of 23 persons, in
the study of ferns and of the fruits and seeds of flowering
plants.

The appended list of electives now offered by its under-
graduate department is taken from the forthcoming cata-
logue of Washington University.

1. Elementary Morphology and Organography, with
reference to Oecology and Systematic Botany.
Lectures and demonstrations by the Professor, as
a full study through the first term.

2. Elementary Anatomy and Phanerogamic Botany. La-
boratory work under the General Instructor, as a
full study through the second term.

3. Synoptical study of the Cryptogams. Laboratory
work under the Instructor in Cryptogamic Botany,
as a full study through the first term, followed, if
desired, by :—

4. A special study of some group of Cryptogams, as a
full study through the second term.

5. Methods of Vegetable Histology. Laboratory work
under the General Instructor, as a full course
through the first term.

6. Histology and Morphology of the Higher Plants.
Laboratory work under the General Instructor, as
a full study through the second term, accompanied,
if desired, by :—

7. A laboratory study of the minute anatomy of the lower
Cryptogams, under the Instructor in Cryptogamic
Botany, as a full course for the same term.

8. Technical Microscopy of Timbers. Laboratory work
under the Professor, as a half course during the
first term.

SIXTH ANNUAL REPORT OF THE DIRECTOR. 25

9. Economic Botany. Lectures by the Professor, sup-
plemented by laboratory demonstrations by the
Instructors, as a full course for the second term.

10-11. Applied Mycology. Laboratory work under the
Instructor in Cryptogamic Botany, as a full course
extending through the year.

12-13. Garden Botany. Laboratory study of cultivated
plants, at the Botanical Garden, under the Director
and his Assistants, for one or two terms.

14. Vegetable Physiology. Laboratory work under the
General Instructor, as a full course for the first
term.

It is intended that course 1 shall always be followed by
course 2, the two being preparatory to other electives.
For the present, unless special reason to the contrary ex-
ists, courses 1 and 2 only will be given each year, the
remaining electives being offered in alternate years a8
follows: —

First year (as offered for 1894-5), —
First term, courses 1, 3, 12, and 14.
Second term, courses 2, 4, 9, and 13.

Second year (as offered for 1895-6), —
First term, courses 1, 5, 8, and 10.
Second term, courses 2, 6, 7, and 11.

Students who have taken courses 1 and 2, or have had
their equivalent elsewhere, are admitted to any of the other
elementary electives which can be taken without conflict
with other University work; but students who desire to
equip themselves as botanists are advised to take the elec-
tives as nearly as possible in the order in which they are
offered, and on the completion of the elective courses
should expect to devote not less than ten hours per week
through an entire year to some piece of research work,
selected under the advice of the Professor of Botany.

26 MISSOURI BOTANICAL GARDEN.

The instrumental, library and herbarium equipment of
the School of Botany and the Garden are or can easily be
made ample for the proper teaching of these electives.
All of the courses capable of being taught in the laboratory
are so taught, supplemented by necessary lectures and
reading, and the few lecture courses offered are illustrated
whenever possible by specimens of plants and their
products.

In addition to these undergraduate studies, post-graduate
work is planned to suit the needs of candidates for advanced
degrees, and one applicant for the Doctor’s degree is taking
such work through the present year.

Besides the classes already mentioned, two advanced
students have been more or less steadily occupied in her-
barium work at the Garden through the year, several
visiting botanists have made protracted use of the herbarium
and library, and the Garden representatives of several
difficult genera of plants have passed through the hands of
specialists engaged in the revision of these groups; and it
is intended that under needful restrictions all of the facili-
ties of the Garden shall always be available for the use of
any suitably prepared person engaged in botanical work.

Very respectfully,

WiLuiaM TRELEASE,
Director.
January 9, 1895.

2 Ne ere} oi ee a

POLS SPOR Rt dak MRE gy ace

SCIENTIFIC PAPERS.

A REVISION OF THE NORTH AMERICAN SPECIES OF SAGITTARIA
AND LOPHOTOCARPUS.

BY JARED G. SMITH.

The principal published revisions of the Alismaceae are
those of Engelmann,* Buchenau,f and Micheli.t The
present revision has been based on a study of the material
contained in the Engelmann and general herbaria of the
Botanical Garden, and in the herbaria of the Shaw School
of Botany, Harvard University, the United States Depart-
ment of Agriculture, Columbia College, the University of
Nebraska, Kansas Agricultural College, the University of
Minnesota, and in the private herbaria of Mr. Walter
Deane, Dr. Charles Mohr, Mr. P. A. Rydberg, and Mr.
Henry Eggert. My cordial thanks are also tendered to
the many correspondents who have contributed specimens to
the Garden Herbarium. I have been especially aided in this
work by and amlargely indebted to the many and copious
manuscript notes and drawings made by Dr. George Engel-
mann during his life-long study of these genera. Not only

* Dr. George Engelmann, in A. Gray, Man. (1848); Ed. 2 (1856);
Ed. 5 (1867). For complete references see The Botanical Works of
George Engelmann, Collected for Henry Shaw, 496 (1887).

+ Franz Buchenau, Index Criticus Butomacearum, Alismacearum
Juncaginacearumque hujusque descriptarum, in Abh. Nat. Ver. Bremen
(1868).— Nachtraege zu den kritischen Zusammenstellungen der bis jetzt
beschriebenen Butomaceen, Alismaceen und Juncaginaceen, Bremen,
(1871).— Beitraege zur Kenntniss der Butomaceen, Alismaceen und Jun-
caginaceen, in Engler, Bot. Jahrb. 2: Heft 5 (1882).—In Engler und
Prantl, Die Natiirlichen Pflanzenfamilien 2: 1, 227 (1889).

¢ Marc Micheli, in DC. Monog. Phan. 3:29 (1881).

27 1

28 MISSOURI BOTANICAL GARDEN.

did Dr. Engelmann examine the material preserved in all of
the principal American herbaria, but in his visits to Europe
in 1856-7, and 1883, he studied the types and American
collections preserved in the great English and Continental
herbaria. These notes and the types in his own very rich
herbarium have served as a foundation for the work
undertaken.

I have followed Buchenau and Micheli in separating the
species of Lophotocarpus from Sagittaria. Lophotocar-
pus is annual (at least our species), the flowers are perfect
or staminate, and the stamens are hypogynous. Sagittaria is
perennial, monoecious or dioecious with the fertile flowers
never perfect, and the stamens are borne above the recepta-
cle. The separation of Lophotocarpus simplifies the genus
Sagittaria very much, reducing somewhat its cosmopolitan
character. In their present form the genera more nearly
express the genetic relationship.

Sagittaria is a genus consisting of from ten to thirty
species, according as the ideas of the various monographers
differ. A large part are American, the rest extending over
Europe and Northern Asia. Of the twenty-two species
which I have recognized in North America, one is only an
introduced ballast plant, and one extends into Central and
South America.

The American S. latifolia, together with the related
species of the group Sagittifoliae, has been considered as
falling within the limits of the widely distributed European
and Asiatic S. sagittifolia. Micheli, in DC. Monog. Phan.
3: 66, 69, and, following him, a number of American sys-
tematists, have held this opinion. Dr. Engelmann, in A.
Gray, Man. Ed. 2 (1856), first pointed out the differences
which separate the two species. Buchenau, in Engler, Bot.
Jahrb. 2: 486 (1882), supports this view that they are
distinct. In this I have followed Engelmann and Buchenau.
The two species occupy different geographical ranges and
are isolated one from the other, although both undoubtedly
derived from the same stock. The American species are

2

SAGITTARIA AND LOPHOTOCARPUS. 29

more closely related to the Asiatic than to the European
forms of sagittifolia.

Classification according to leaf forms has proved very
unsatisfactory. After examining nearly two thousand
herbarium specimens, besides much living material, I have
found that there is least variation in the form and size of
the achenium, form of the anthers, pubescence or lack of
pubescence of the filaments, relative length of fertile and
sterile pedicels, and within certain limits in the form of the
bracts. As in most aquatics, the leaves vary through wide
limits in the same species, and characters founded on leaf
differences, at least among the Sagittifoliae, are of little
value. Some reliance, however, may be placed on the
relative length of the lobes or divisions of the leaf, except-
ing always in the immature or phyllodial forms. The
phyllodia themselves are too variable to be of much diag-
nostic value, though some of our more recent American
systematists have endeavored to draw specific characters
from them. Phyllodia are not always present. In some
species they seem to offer wider differences than the leaves.
They appear to be formed principally when conditions are
unfavorable for the development of aerial leaves, and
whether they are considered degraded leaf forms, or rever-
sions to a previously existing type, the characters which
they present have little specific value.

Intravaginal squamules as noted by Buchenau in Engler,
Bot. Jahrb. 2: 467 (1882), are present in the Alismaceae
and allied orders. They consist of from one to three rows
of very delicate scales attached in the axils of the leaves to
the dorsal edge of the petiole. They are linear-lanceolate,
2 to 4 mm. long, colorless, from one to three or four cells
thick at the base. They may be easily studied on young
plants of the common VS. latifolia, and will probably be
found in other species.

Many or possibly all of our North American Sagittarias,
in common with the Old World species, form underground
autumn tubers. These have been observed in S. latifolia,

3

30 MISSOURI BOTANICAL GARDEN.

S. graminea, S. heterophylla, S. longiloba and 8. pa-
pillosa, Horizontal or oblique rhizomes are present in S.
lancifolia and S. graminea cycloptera. The tubers of
latifolia, graminea and heterophylla, are edible. They are
used by the Indian tribes of the Northwest, and are called
either Wappatoo, or Wabesipinig, the latter meaning Swan-
potatoes, because of their furnishing food for the numer-
ous wild-fowl of that region. The tubers are formed in
the mud at the depth of 2 to4dm. In the spring they
throw out a long rhizome which forms at its upper end a
corm, and from this corm arise the roots and leaves.

The descriptions given in the following synopsis have
been drawn as far as possible from herbarium specimens or
the living plants. The exceptions were Sagittaria lanci-
folia angustifolia (Lindl.) Griseb., and Lophotocarpus
Guyanensis (HBK. )Mich., both from Mexico. These were
adapted from Micheli, because of the insufficiency of
material in the herbaria examined.

Since contributing a list of the Sagittarias of the Eastern
United States to the Mem. Torr. Bot. Club, Vol. 5: 25
(1894), I have examined fuller material of 8. graminea
platyphylla, and have thought best to raise the variety to
specific rank.

The limited bibliography given includes the principal
works used in the preparation of this paper. It was
believed that its value as a systematic monograph would
not be enhanced if the great mass of minor references were
printed. I have endeavored to make the synonymy com-
plete as far as original descriptions of American species
are concerned. Buchenau and Micheli have given the
fuller bibliography, and I will refer any who are interested
in the study of these genera to their cited works. If
deemed necessary, the more complete bibliography may be
published at some future time.

Saqirraria L. Sp. Plant, 993 (1753). — Flowers all
unisexual, but the staminate frequently with abortive pis-
tils at center; filaments inserted above the receptacle.

4

SAGITTARIA AND LOPHOTOCARPUS. 31

ANALYTICAL KEY.

1. Sepals of the pistillate flower reflexed or spreading, not accrescent.
* Fertile pedicels slender, ascending, neither much thickened nor
reflexed in fruit.
+ Leaves sagittate; filaments glabrous, not dilated.
++ Basal lobes equaling or shorter than the middle one.
= Beak more than one-fourth the length of the achenium.
Fertile pedicels much shorter than the bracts; leaves ample;

beak of the achenium stout, erect. S. longirostra.
Fertile pedicels longer than the bracts; leaves with linear
lobes; beak of the achenium erect. S. Engelmanniana.

Beak of the achenium lateral, horizontal or oblique. VS. latifolia.

= = Beak of the cuneate obovate achenium very short, erect, re-
trorse; basal lobes very short.

Petioles short and curving; bracts long, scarious margined;

terrestrial, or emergent aquatic. 8S. arifolia.
Petioles long and slender; bracts short; submerged aquatic
with flowers at the surface of the water. S. cuneata.

++ ++ Basal lobes more than half the length of the leaf.
Basal lobes 2 or 3 times as long as the middle one; stamens

less than 24; seed coat punctate. S. longiloba.
Basal lobes less than twice as long as the middle one; stamens
80 or more; seed coat not punctate. S. Greggii.

+ + Leaves entire or hastate.
++ Filaments slender, not dilated; leaves pseudo-penninerved.
Filaments glabrous; bracts short, densely papillose; achenium

short-obovate. S. papillosa.
Filaments glabrous; achenium falcate; fertile pedicels longer
than the sterile. 8. ambigua.

Filaments cobwebby pubescent; achenium falcate. S&S. lancifolia.
++ ++ Filaments dilated, pubescent; veins free to the base of the leaf.
a. Fertile flowers sessile; scape geniculate at the lowest verticil;

achenium stoutly beaked. S. rigida.
b. Fertile pedicels shorter than the sterile; achenium 3 mm. long,
broadly winged, the sides smooth.— In South Carolina.
S. macrocarpa.
c. Fertile pedicels about as long as the sterile.
Leaves terete and mostly bladeless.—Species of the Atlantic

coast. S. teres.
Leaves rigid, elliptical-linear, somewhat triquetrous. S. cristata.
Leaves neither rigid nor terete. S. graminea.

** Fertile pedicels much thickened, reflexed in fruit.
a. Filaments dilated, pubescent. S. platyphylla.

b. Filaments dilated, glabrous.
Stamens 7 or 8.
Scape simple. S. subulata.

4)

32 MISSOURI BOTANICAL GARDEN.

Scape branching, longer than the linear submerged

leaves. S. filiformis.

Stamens 12 or 13. S. demersa.

c. Filaments slender, glabrous; leaves entire; achenia cuneate
quadrate. S. Sanfordit.

* ** Fertile pedicels much thickened, ascending; filaments slender, glab-
rous; scape shorter than the leaves. S. Mexicana.

2. Sepals of the pistillate flower erect and accrescent; petals with a
brownish purple spot at the base. S. Montevidensis.

SYNOPSIS OF THE SPECIES OF SAGITTARIA NORTH OF
MEXICO.

A. Sepals of the pistillate flower refiexed after flowering, not enlarg-
ing in fruit; petals white.

§ Sagittifoliae. Leaves sagittate or rarely when submerged entire, or
reduced to bladeless phyllodia; filaments glabrous, not enlarged at the
base; sterile pedicels longer than the fertile.

* Achenium obovate cuneate, the beak minute, erect, retrorse; fila-
ments shorter than or equaling the anthers; basal lobes less than half
the length of the leaf; bracts slightly connate.

S. arrrouia Nutt. in herb.— S. sagittifolia minor Pursh,
Flora Amer. Sept. 2: 395 (1814).* WS. variabilis
var. hastata forms b and ¢ in part, Macoun, Cat. Can.
Pl., 4: 77, 78 (1888).

Terrestrial, or emergent aquatic, weak, ascending, 2 to 4
dm. high; petioles rather stout, usually curving outward ;
blade of leaf 6 to 12 or 18 cm. long, arrow-shaped, acute,
the margin mostly straight or arcuate, basal lobes diverg-
ent, acute or acuminate; scape weak, ascending, simple or
rarely branched; bracts lanceolate, acute, 8 to 20 mm.
long, scarious margined and obscurely veined, often
reflexed; 1 to 3 lower verticils pistillate; fertile pedicels
ascending, 15 to mostly 25 mm. long, or sometimes almost
wanting; fruiting head round, 8 to 15 mm. in diameter;
achenium 2 mm. long, tumid winged on both margins, the

* Pursh gives the range as ‘‘Pennsylvania to Carolina;’’ but Dr.
Engelmann in his manuscript notes says that Pursh’s plant is from the
Rocky Mountains and that it ‘‘seems to be exactly like’’ a specimen
collected by Nuttall.

6

SAGITTARIA AND LOPHOTOCARPUS. 33

sides smooth, or often with a vertical subepidermal resin
passage. Phyllodia of two forms, either long, slender,
petiole-like, or flattened, linear-lanceolate, 2 to 5 dm. long
and 10 to 15 mm. wide.— Along streams or sandy mar-
gins of lakes and ponds, in the mountains from British
Columbia to California, Nevada and New Mexico, and from
western Kansas and Nebraska to Minnesota and Quebec.
Phyllodial or immature specimens are difficult to dis-
tinguish from plants of S. latifolia form c of similar
habit. The species perhaps approaches most closely the
Northern European and Asiatic S. sagittifolia, from which,
however, mature specimens are easily separated. The
achenia are much smaller, and cunate-obovate instead of
nearly orbicular as in the European species. — Plate 1.

Specimens examined from British Columbia (Macoun, Kamloops, 1879,
and 1889, S. variabilis hastata form c, Sicamous, 1889); Washington
(Dr. Lyall, 1860; Suksdorf, 1318, Sept. 1893, Klickitat Co.); Oregon
(Kellogg and Harford, 952, 1869; Howell, Sauvies Island, 1887; Nuttall) ;
California (Bolander, Sierras, 1870; Vasey, 1875; Mrs. Ames, 1876, Plu-
mas Co.; Mrs. Austin, 1877); Nevada (Stevens, Pac. R. R. Surv.; Torrey,
506, 1865, Lake Washoe; Bailey, 1151, 1867, Truckee Valley; Palmer,
1876); Utah (S. Watson, 1151, 1869, Salt Lake Valley; Jones, 1071, Salt
Lake City, 1879) ; Colorado (Jones, 1878, Denver) ; New Mexico (Wislize-
nus, 27 and 225, 1846, the latter with irregularly approximate whorls; Fend-
ler, 837 and 839, 1847; Edwards, 1849, Santa Fé, the “ 8. Mexicana’’ of
Herb. Torrey but not of Engelmann); Northwest Terr. (Parry; Macoun,
1751, 1879, Eagle Hills “ 8. variabilis gracilis’’, —— Land Hills, “ 8,
variabilis hastata form b,—— and 1883,Moose Mountain Creek) ; Manitoba
(Bourgeau, 1857, Winnipeg Valley); Idaho (Leiberg, 526, 1891, Lake Pend
d’Oreille, ‘‘in water 3-12 dm. deep, or in simply wet places,” “‘on sand-
bars, shallow bays and mouths of rivers, abundant,’”? many forms);
Montana (Notestein, 1893, Great Falls); Geyer, Nicollett’s Northwest
Expedition, 1839; S. Dakota (Williams, 16, 1892, Big Stone Lake and many
other localities. Specimens from Big Stone Lake have branched scape
with whorls irregularly approximate, lobes of leaf linear, much elongated,
and two kinds of phyllodia on the same plant) ; Nebraska (H. Engelmann,
Platte Bottoms, 1852, 1856 and 1858, the latter with bracts 25 to 30 mm.
long; Smith and Pound, 235, 1892, Sand Hills; Bates, Sand Hills); Iowa
(Hitchcock, 1891); Minnesota (Minneapolis, 1877, ““N. H. W.,’? with
broadly ovate leaves, submerged scape, the irregularly approximate
fertile verticils each with 3 to 6 flowers; Bailey, 154, 1886, Vermillion
Lake; Sandberg, 1179, 1891, Lake Itasca) ; Michigan (Davis, 1890, Alma;
Kofoid, 1890 and 1891, Cheboygan Co., and “‘ sandy margins of Black

3 7

34 MISSOURI BOTANICAL GARDEN.

Lake)’ ; Ontario (Macoun, 1884, Nipigon River, S. variabilis hastata form
c). An immature specimen from Macoun, 1882, Grand Vallée, Gaspé, S.
variabilis hastata form c, probably goes here.

S. ARIFOLIA STRICTA 0. var.

Slender, erect, 3 to 4 dm. high; blade of leaf 2 to 5
cm. long; scape simple, erect; bracts ovate, acute, 6 to 8
mm. long; fertile pedicels 8 to 15 mm. long; fruiting
head 1 cm. in diameter; achenium smooth or laterally uni-
costate.— Boggy meadows and slow streams, Falcon Valley,
Washington, W. N. Suksdorf, 1319 (674), Aug. 1, 1885.—
Plate 1.

S. cunnata Sheldon, Bull. Torr. Club, 20: 283, pl. 159,
(1893).

Submerged aquatic, rooting in the sand; leaves long
petioled; blade floating, small, 5 to 8 cm. long, with linear
lobes; scape simple, slender, terete, 6 to 9 dm. long, bear-
ing verticils of flowers at the surface of the water; bracts
ovate-lanceolate, acute, 4 to 6 mm. long; stamens few (?);
fruiting head small, about 1 cm. in diameter ; achenium 1
mm.long. Phyllodia of two forms, either linear attenuate,
petiole-like, reaching nearly to the surface of the water,
or lanceolate, 6 to 12 cm. long, 4 to 8 mm. wide, clustered
at the base.— In shallow ponds, or margins of lakes, from
British Columbia to Minnesota.— Plate 2.

Specimens examined from British Columbia (Dawson, Aug. 1888, Kam-
loops; Macoun, 1889, Sicamous); North West Territory (Macoun, 1751,
1879, Eagle Hills); Washington (Suksdorf, 2262, 1889, Philles Lake,
Spokane Co.); South Dakota (Williams, 15, 1892, Big Stone Lake) ;

Minnesota (Sheldon, 3576, East Battle Lake,—39254, Blanche Lake,—
39264, Mollie Stark Lake, 1892).

** Achenium obovate, the beak prominent, 4 to 4 as long as the ovary;
filaments longer than the anthers; basal lobes not more than half the
length of the leaf; bracts, at least of the lower verticils, free.

S. uatironia Willd. Sp. Pl. 4: 409 (1806). WS. gracilis
Pursh, Fl. Am. Sept. 396 (1814). S. simplex Pursh,
l. ¢., described from staminate scape, and leaf of
Rumex. 8S. variabilis Engelm. in A. Gray, Man. 461

8

SAGITTARIA AND LOPHOTOCARPUS. 33

(1848). SS. sagittifolia of Amer. authors, not Linn.
S. sagittifolia, var. variabilis Micheli in DC. Monog.
Phan. 3: 69 (1881), in part, not 8. longiloba Engelm.
8. variabilis var. angustifolia Engelm. in A. Gray,
Man. Ed. 5, 493 (1867). &. variabilis var. diversi-
folia, Engelm. in A. Gray, Man. Ed. 5, 493 (1867).
S. variabilis var gracilis Engelm. in A. Gray, Man.
Ed. 5, 493 (1867). SS. hastata Pursh, 1. c. &. va-
riabilis var. hastata Engelm. in A. Gray, Man. Ed. 5,
493 (1867). WS. variabilis var. hastata form a, and
in part forms 6 and c, Macoun, Cat. Can. Pl. 4: 77, 78
(1888). S. obtusa Willd. Sp. Pl. 4: 409 (1806). S. sag-
ittifolia var. macrophylla, and var. vulgaris Hooker Fl. Bor.
Amer. 2: 167 (1840). S. Chinensis Parish, Zoe, 1: 122
(1890). WS. Sinensis Brandegee, Zoe, 4: 217 (1893),
not Sims.

Monoecious, with the lower verticils fertile, or dioecious;
scape 1 to 12 dm. high, angled, simple or branched; flow-
ers large, 2 to 4 cm. wide, the petals white; stamens
numerous, 25 to 35; fertile pedicels shorter than the ste-
rile; bracts sometimes connate in the upper verticils, acute,
acuminate, or obtuse, not scarious; achenium broadly
winged on both margins, 2.5 to 3.5 or rarely 4 mm. long,
with a lateral horizontal or curving beak } to } its length,
sides usually smooth, or with a costate angle curving down-
ward from the base of the beak, rarely with a subepidermal
resin passage on each face.— An extremely variable species,
extending from Nova Scotia to British Columbia, south
along the Pacific Coast to California, and throughout the
region east of the Rocky Mountains to Mexico and
Florida. —Plates 3 to 7.

The species is a complex one on account of its wide geo-
graphical distribution and varying habitat. After an ex-
amination of nearly all the available material, the species
seems best divided into forms instead of varieties, of which
I recognize five. These merge into one another, so that it

is difficult to draw hard and fast lines between them. The
9

36 MISSOURI BOTANICAL GARDEN.

first represents, as far as I am able to determine, the
species in its most typical form.

8. latifolia proper.— Monoecious or subdioecious ; large,
erect, 3 to 6 dm. high; leaves longer than broad, glabrous
or rarely pubescent, 15 to 40 cm. long, with lobes from
broadly ovate, acute, to linear lanceolate, acute or acu-
minate; scape erect, simple or branched; bracts glabrous,
acute or acuminate, 1 to 5 cm. long; verticils rarely ap-
proximate, mostly more than five; fertile pedicels 2 to 4 or
sometimes 6 or 8 cm. long; fruiting head 15 to 30 mm. in
diameter; achenium about 3 mm. long.— This is the com-
mon form from Massachusetts and New York west to
Colorado, and south to Florida and Louisiana.

Specimens examined from Prince Edward Island (Macoun, 1888);
Ontario (Macoun, 1884; Britton, 1889); Massachusetts (Deane); New
York (Torrey; Morong); New Jersey (Dr. Gray, 1833; Bernhardi; Brit-
ton, 1889); Pennsylvania (Durand, 1848; Porter); Ohio (Riehl, 1838) ;
S. Carolina (M. A. Curtis); Alabama (Mohr); Indiana (Case, 1878; 8.
Watson, 1890, Crawfordsville, has a branched scape with 6 lower whorls
fertile, fruiting head 20 to 25 mm. in diameter on pedicels 15 to 20 mm.long,
bracts 3 cm. long, acuminate, achenium crested, 2.5 mm. long, the beak
oblique); Illinois (Engelmann and many collectors); Missouri, common;
Michigan (Farwell, 729, 1889; Schneck, 1881; Davis, 1890); Minnesota
(Sandberg, Red Wing, 1885, 622, 1891; Taylor, 907, 1891, Glenwood) ; |
Iowa, common; S. Dakota (Williams, 1892, numerous localities) ; East-
ern Nebraska and Kansas, common; Colorado (Denver, Fritchey, 1886);
Louisiana (Dr. Hale, specimens with ovate bracts 1 cm. long); Florida
(Dr. Chapman, 1863, with the upper part of the petiole and the under
surface of the leaves pubescent, fertile pedicels and acuminate bracts 1
cm. long, bracts and sepals glabrous).

This form includes a part of communis, and all of acuta
and subdioica of the Engelmann Herbarium, and a part of
Macoun’s var. Aastata, Cat. Can. Pl. 7. c. The common
form in Indiana, Illinois, lowa and Missouri, has a dorsally
crested obliquely beaked akene 2 to 2.5 mm. long, fertile
pedicels 12 to 15 mm. long, shorter than the acuminate
bracts. This passes into the form with longer (3 to 3.5
mm.) akenes with horizontal beaks. Iam unable to sep-
arate them except geographically. The dorsally crested

10

SAGITTARIA AND LOPHOTOCARPUS. 37

form is confined to the Mississippi Valley. The horizontal
beaked form is found throughout the range.

Two specimens in the Engelmann Herbarium have a
bract of the lowest whorl foliaceous; that of a specimen
collected in Iowa by Wood is linear-lanceolate, acute, fal-
cate, 8 cm. long; the other, collected by Engelmann in St.
Louis, is 45 cm. long. ‘* Doubled”’ flowers in which the
outer stamens become petaloid have been collected at Lan-
caster and Harrisburg, Pa., by Prof. Porter; at West
Troy, N. Y., by Wibbe; and at Cooperstown, N. Y., by
Miss Keyes, in 1886.

Form a.— Dioecious; leaves longer than broad, 15 to
30 cm. long, from ample, broadly ovate, obtuse, to linear
lanceolate, gradually acuminate; scape simple or branched ;
bracts ovate, obtuse, 8 to 15 mm. long, glabrous. — From
New Brunswick to Minnesota, Louisiana and South Caro-
lina. It includes S. obtusa Willd., S. variabilis obtusa,
Engelm., and S. hastata Pursh.

Specimens examined from New Brunswick (Macoun, 1876, Cambell-
ton) ; Ontario (Macoun, 1873, Hastings Co.; 1877, Belleville; 1884, Nation
River); New Hampshire (Deane, 1889) ; Connecticut (Eaton); New York
(Torrey; Gray; Trelease) ; Pennsylvania (Canby); New Jersey; Delaware
(Durand); South Carolina (Ravenel); Kentucky (Short); Louisiana
(Hale); Missouri (Engelmann; Bush, 1891); Illinois (Engelmann; Hitch-
cock); Indiana; Michigan (Engelmann, Ontonagon; Schneck, 1881);
Wisconsin (Trelease; Ballard, 666, 739, 1891, Waconia); Minnesota
(Dewart, 1888; Dr. Mearns, 673, 674, 1888, Ft. Snelling; Sandberg, 1881,
Vasa; Bailey, 151, 1886, Vermillion Lake; Oestland, 1886, Minnehaha;
Ballard, 1891, 607, 731, 808, 830; Sheldon, 1892, 3061, 3256); S. Dakota
(Williams, 1892); Iowa (Arthur; Hitchcock, 1890); Nebraska (Rydberg,
1887, 1890, Wahoo); Kansas (Pellet, 1890, Johnson Co.; Hitchcock,
1892, Medicine Lodge).

Form b.— Leaves large, 1 to 4 dm. long, the lobes from
lanceolate to broadly ovate, apex obtuse, or abruptly acute,
basal lobes acuminate, divaricate ; scape simple or branched ;
bracts scarious, 5 to 10 mm. long, ovate, obtuse; achenium
3 mm. long with rather tumid dorsal wing, and long hori-
zontal beak, subepidermal resin passages usually present
on each face.— From Washington to Southern California.

11

38 MISSOURI BOTANICAL GARDEN.

Plants from central to southern California with very
large leaves about as wide as long, and smooth, usually
branched monoecious scape, form the S. Chinensis or
Sinensis of Parish and other collectors, but are not syn-
onymous with the Asiatic S. Sinensis Sims. The edible
tubers are used by the Chinese.

Specimens of this examined were: 1367, Wilkes Exploring Expedition,
1838-1842, Sacramento Valley; Brewer, 2189, 1861, Pitt River, among the
tules; Kellogg, near San Francisco; Jepson, 1891, Grand Island, Lower
Sacramento R.; Parish, 2091, 1889 and 1893, San Bernardino, the latter
probably introduced from farther north.

From northern California to Washington the common
form has narrower rather coriaceous leaves, and dioecious
scapes. The edible tubers,* (called Wappatoo), were
formerly staple articles of commerce among the Indian
tribes along the Columbia river and its tributaries.

Specimens of this examined from California (Moore, Grass Valley;
Mrs. Austin, 1878, Plumas Co.); Oregon (E. Hall, 502, 1871; Collier,
Eugene City); Washington (Drake and Dickson, 1883; Suksdorf, 673,
1885, — 2261, 1890, Kirkland; C. V. Piper, Seattle, 1892).

Form c.— Monoecious or dioecious; leaves 5 to 10 or 15
em. long, the middle lobe ovate lanceolate or lanceolate,
rounding to the acute apex, basal lobes narrower and often
shorter, lanceolate, acute or acuminate and widely diver-
gent; verticils few, 2 to 5; the short fertile pedicels about
} the length of the sterile; bracts ovate, acute, 5 to10 mm.
long; fruiting head about 15 mm. in diameter; achenium
2.5 to 8 mm. long. Phyllodia frequently present, either
petiole like or flattened and linear lanceolate. This includes
the varieties of variabilis: hastata, angustifolia, gracilis
and diversifolia of most collectors.— Most abundant in the
region of the Great Lakes and northwards. It ranges from
Prince Edward Island to British Columbia and southward
to New York, Kentucky and Nebraska. Immature or

* See History of the Exped. Lewis and Clarke, Revised and Abridged
Ed. by A. M’Vickar, 2: 85, 108, 107, 171, 8368 (1842). Hooker Fi. Bor.
Amer. 2: 167 (1840).

12

SAGITTARIA AND LOPHOTOCARPUS. 39

phyllodial forms are difficult to distinguish from 8. arifolia
except by the longer style, and lateral beak of the
achenium.

Specimens examined from Prince Edward Island (Macoun, 1888); New
Brunswick (Chadborne, 1883); Quebec (Northrop, 1887, Detour du Lac,
Lake Penniscouta); Vermont (Deane, 1885, Willoughby Lake; Rusby,
1892; Morong, 1885, Ferrisburgh, with lanceolate attenuate phyllodia 1
dm. long, and floating leaves 3 cm. long); New York (Engelmann, St.
Lawrence River below Ogdensburg, with almost sessile staminate
flowers; Sartwell, Penn Yan, with fertile pedicels only 5mm. long; Clin-
ton, Alexander’s Bay, with floating lanceolate leaves 3 to 5 cm. long, and
clustered lanceolate phyllodia; Engelmann, 1856, Table Rock, Niagara) ;
Pennsylvania (Wolle, Bethlehem, the form communis of Engelmann) ;
Ohio (Riehl, 1836); Kentucky (Rafinesque; Short, 1840); Michigan
(Farwell, 419, 1886, and 460, 1890); Ontario (Macoun, 1873, Bay of
Quinte,‘ hastata form c.;’’ Bell, 1878, Missinaiba River, ‘‘ hastata
form c’’.); Manitoba (Bourgeau, 1858, Lake Winnipeg; Macoun, 1884,
Lake Winnipeg, hastata form b.); Minnesota (Pitcher; Wood, 1889;
* Ballard, 163, 831, and 897, 1891, and 1074, 1892; Sheldon, 921,
1891, Sleepy Eye, and East Battle Lake, 1892; Taylor, 642, 1891,
Minnesota Lake); S. Dakota, (Duffey 1889); Nebraska (Rydberg, 1887,
Wahoo, 1890, Lodge Pole Cr., 1891, Scotts Bluff and Deuel Counties,
and 1533, 1812, 1898, Hooker Co.); Saskatchewan (Bourgeau, 1858) ;
British Columbia (Macoun, 1889, mouth of Fraser River, “ hastata
form b.;”’ Lyall, 1858, 49th parallel, Oregon Boundary Survey).

Form d.— Monocecious, very slender, 20 to 25 cm. high;
leaves 8 to 10 cm. long, the lobes narrowly linear, diver-
gent; fertile pedicels 10 to 15 mm. long; verticils rather
remote; fruiting head 1 cm. in diameter.— In the mount-
ains, New York to New Hampshire.

Specimens examined from New York (Mrs. C. Van Brunt, 1886, Balsam
Lake, Catskill Mts.; Torrey); Vermont (Dr. Chapman, Lake Dunmore) ;

New Hampshire (Deane, 1883 and 1884, Shelburne, the variabilis angusti-
folia of S. Watson, but not of Engelmann).

Form e. — Monoecious, erect, 4 to 10 dm. high, with the
habit of typical /atifolia; lower leaves sagittate, the upper
ovate-lanceolate, acute at both ends, the largest 10 cm.
long and 4 to 5 cm. wide; achenium 3.5 mm. long, dorsally
crested, with the beak somewhat recurved at the tip. —
Collected by Dr. Engelmann on the American Bottoms,
opposite St. Louis, in 1856. This form is the S. variabilis

13

40 MISSOURI BOTANICAL GARDEN.

diversifolia of Engelmann. Specimens called diversifolia
by collectors are, without exception, phyllodial or imma-
ture plants, from deep or running water, and belonging
mostly to form c.

S. LATIFOLIA PUBESCENS (Muhl.) J. G. Smith, Mem. Torr.
Bot. Club, Vol. 5: 25 (1894). 8. pubescens Muhl. Cat.
86 (1813). W&. sagittifolia var. pubescens Micheli in
DC. Monog. Plan. 3: 69 (1881). WS. variabilis var.
pubescens Engelm. in A. Gray, Man. Ed. 5, 493
(1867).

Monoecious or dioecious, erect, 3 to 5 dm. high; leaves 7
to 15 or 25 cm. long, broadly ovate, rounding to the
abruptly acute apex; bracts and sepals ovate, obtuse, pubes-
cent or woolly ; leaves, and especially the scape, from pube-
rulent or minutely hirsute to densely pubescent; scape
often very slender; achenium 2 to 3 mm. long; fruiting
head 8 to 15 mm. in diameter; fertile pedicels as long as
or shorter than the sterile. — Outasio to Florida, from the
mountains to the coast. There are three quite well marked
forms. — Plate 8.

form a. — Habit of form a of the species; leaves large,
obtuse; achenium costate-angled, 3 mm. long. This un-
doubtedly represents Muhlenberg’s type.— From Delaware
northward.

Specimens examined from Ontario (Macoun, Bay of Quinte, 1877, very
close to form a of the species, but with pubescent bracts); Pennsylvania
(Darlington; Canby, Chester Co., 1863); Delaware (Canby, 1863).

form b. — Leaves smaller, 7 to 15 em. long, light green,
sub-coriaceous ; bracts and sepals densely pubescent ; achenia
2.5 mm. long, smooth on the sides. — The common form
in the mountains from Virginia southward.

Specimens examined from Virginia (A. H. Curtiss, 1871; Small, 1892;
Britton and Vale, 1892); North Carolina (Rugel, 1842; J. D. Smith,
1881; McCarthy, 101, 1885; Boynton, 1888; Kofoid and Beardslee, 1891) ;
South Carolina (Ravenel, 1869).

Form c.— Seape simple, sooner with the fertile pedi-

14

SAGITTARIA AND LOPHOTOCARPUS. 41

cels about equaling the sterile; leaves small, thin and
papery in texture; pubescence less dense; achenia about 2
mm.long. Mature specimens have not yet been collected. —
Georgia, Florida and Alabama.

Specimens examined from Georgia (Dr. Wray, Augusta); Alabama

(C. Mohr, Montgomery and Mobile); Florida (Mary C. Reynolds, 1877,
St. Augustine).

S. ENGELMANNIANA n. sp. J. G. Smith in Mem. Torr. Bot.
Club. Vol. 5: 25 (1894). WS. variabilis var. ( ?) gra-
cilis S. Wats. in A. Gray, Man. Ed. 6, 555 (1889)
in part, not Engelm.

Monoecious, slender, erect or ascending, 2 to 4 dm.
high, glabrous ; leaves 8 to 20 cm. long, the lobes linear, 1
to 5 mm. wide, apex rounded or acute, basal lobes acumi-
nate; scape simple, striate, equaling the leaves; verticils
4 to 6, the lower fertile; bracts lanceolate, acute, 8 to
12 mm. long; flowers 2 to 3 cm. wide; petals white,
rounded, entire, with a very short claw; sepals short,
ovate, acute; stamens 18 to 25; pistils numerous, the style
nearly twice as long as the ovary; pedicels slender, ascend-
ing, the fertile 10 to 14 mm., the sterile 10 to 20 mm. long ;
achenium narrowly obovate, 4 mm. long, the stout erect
beak 4 to } the length of the achenium, entire-winged
on both margins, with 1 to 3 strong lateral wings extend-
ing downward from the base of the beak on each side;
fruiting head globose, 12 to 14 mm. in diameter.— Rare, in
shallow ponds, from Massachusetts to Delaware and
Florida (?). Dedicated to Dr. George Engelmann, by
whom it was collected at Lake George, New York, in
1856.— Plate 9.

Specimens examined from Florida (?) (Chapman, in Herb. Columbia
College); Delaware (specimens labeled S. variabilis graminifolia, in
both the Engelmann and Bernhardi Herbaria); New Jersey (Diffen-
baugh, 1864, Brown’s Mills, labeled S. variabilis gracilis; Britton, 1889,
Ocean Co. “ 8. varabilis gracilis’’); New York (E. 8. Miller, Wading R.,
Long Island); Massachusetts (Robbins, Sept. 1864, Uxbridge, the 8.
variabilis gracilis of many herbaria; Deane, 1888, Hyannis, and Barn-
stable; Sturtevant, 1890, S. Framingham). 15

492 MISSOURI BOTANICAL GARDEN.

S. Lonerrostra (Micheli) J. G. Smith in Mem. Torr. Bot.
Club, Vol. 5: 26 (1894). WS. sagittifolia var. (?)
longirostra Micheli, in DC. Monog. Phan. 3: 69 (1881).

Monoecious, erect, 4 to 8 dm. high; leaves ample,
smooth, 10 to 25 cm. long, middle lobe broadly ovate-
rounded, abruptly acute; basal lobes ovate, acute; scape
simple, exceeding the leaves, 6-angled below, 6 channeled
above in fruit, and minutely scabrous on the angles; bracts

15 to 30 mm. long, triangular, acuminate; fertile pedicels

ascending, 1 cm. long or less; pistils numerous, the curved

style about twice the length of the ovary; achenium 4 mm.

long, broadly obovate with a stout erect recurving beak,

broadly winged on both margins, the ventral entire, the
dorsal irregularly crenate, a single broken wing or crest on

each face; fruiting head depressed globose, 12 to 18 mm.

in diameter, subsessile. — Margins of ponds, from south-

eastern Pennsylvania and New Jersey to Alabama. This
is the S. variabilis echinocephala of the Engelmann Her-

barium.—Plate 10.

Specimens examined from Pennsylvania: (Wm. M. Canby, Aug. 1863,
Delaware Co., and Oct. 1863, Chester Co.; T. P. James, 1848, West
Philadelphia); New Jersey (James, 1848, near Camden); Delaware
(Canby); Kentucky (Kearney, 187, Aug. 1893, Harlan County); Ala-
bama (Drummond, a few flowers in the Engelmann Herbarium from the
type specimen of the Kew herbarium).

* * * Achenium quadrate obovate, almost beakless; filaments longer
than the anthers; basal lobes more than half the length of the leaf;
bracts slightly connate.

S. tonartospa Engelm. in Torr. Bot. Mex. Bound. Surv.
212 (1858). WS. sagittifolia var. Mexicana Mart. and
Gal., Bull. Acad. Roy. Brux. 9: 379 (1842), not S.
Mexicana Steud.

Monoecious, slender, erect, 3 to 5 dm. high; leaves 10
to 18 cm. long, the linear-lanceolate, tapering, acuminate
basal lobes 2 to 3 times as long as the acute, linear to ovate-
lanceolate middle one; scape simple or rarely branched ;
verticils 4 to 8, 1 to 3 lower ones fertile; bracts lanceolate,
acuminate, 6 to 8 mm. long; fertile pedicels slender, diva-

16

TR he Meets MEI lad i

SAGITTARIA AND LOPHOTOCARPUS., 43

ricate, 15 to 35 mm. long, exceeding the sterile; stamens
15 to 21; achenium 2 mm. long, narrowly winged on both
margins and costate from the base of the beak above, sub-
epidermal resin passages usually present; fruiting head 10
to 12 mm. in diameter; seed obovate, 1 mm. long, the
seed-coat punctate. — Margins of shallow ponds, from
western Kansas and eastern Colorado south to Texas, New
Mexico and Sonora. — Plate 11.

Specimens examined from Kansas (Kellerman, Gove Co., 1885; Hitch-
cock, 1892, in Ford, Seward and McPherson counties) ; Oklahoma (Carle-
ton, 262, 1891, Cherokee Outlet); Colorado (Iendler, 850, 1847, Arkansas
R.; Beckwith and Gunnison, 49, Upper Arkansas R.); Texas (Lind-

heimer, June, 1847, Pierdenales; Wright, 680, and 1140, 1849); Mexico
(Palmer, 1889, Lerdo, Sonora, specimens with winter tubers attached).

S. GREGGII n. sp.

Monoecious, stout, erect, 10 to 14 dm. high; leaves 20
to 40 cm. long, the lanceolate, acuminate, widely divergent
basal lobes longer than the lanceolate or ovate and acumi-
nate middle one; scape erect, 5-angled below, paniculately
branched from the lowest verticils, exceeding the leaves;
branches slender, striate, ascending; verticils numerous,
the lower fertile; bracts lanceolate, acuminate, 15 to 25 mm.
long; fertile pedicels 15 to 30 mm. long, about as long as
the sterile; sepals minutely scabrous; stamens 30 or more,
4to 5mm. long; fruiting heads globose, 8 to 15 mm. in
diameter; achenium 2 to 3 mm. long, tumid crested on
both margins, from obovate with the ventral margin nearly
straight to almost circular, edges acute, sides with an irreg-
ular saucer-shaped wing, and more or less tubercled; seed
1 mm. long, smooth.—In ponds and ditches, California and
Mexico. Collected by J. A. Sanford, July 1893, Stock-
ton, Cal.

The species is dedicated to Dr. J. Gregg, whose No, 833,
collected in May 1849, at Zamora, Michoacan, Mexico,
seems to be the same as our Californian plant. — Plate 12.

§§ Fluitantes: Leaves floating or submerged; fertile pedicel or pedicels
thickened and reflexed in fruit, longer than the sterile; bracts at first

17

44 MISSOURI BOTANICAL GARDEN.

connate to the apex, and spathe-like; filaments dilated, bottle-shaped,
not pubescent; anthers suborbicular.

S. susutata (Linn.) Buchenau, Abh. Nat. Ver Bremen,
2: 490 (1871). Alisma subulata L. Sp. Pl. 343
(1753). WS. pusilla Nutt. Gen. 2: 213 (1818).
S. natans var. lorata A. Gray, Man. Ed. 5, 494 (1867),
not Chapm.

Monoecious, with 1 or 2 flowers of the lowest verticil
fertile, or all staminate; low, 5 to 15 cm. high, usually
with rigid, obtuse or acutish phyllodia, or rarely with
linear-lanceolate, subacute blades 2 to 3 cm. long, narrow-
ing at the base into a spongy petiole; scape simple, equaled
or exceeded by the leaves, few flowered; flowers rather
large for the size of the plant, 15 mm. across; sepals ovate-
lanceolate, obtuse; fertile pedicels 10 to 15 mm. long;
bracts acute; stamens 7 or 8, less than 2 mm. long, the
filaments equaling the anthers; acheniarather few, nearly 2
mm. long, 3-crested, with a short and slender, abrupt
lateral beak and a short, fusiform resin passage down the
middle of each face; fruiting head small.— In mud, tide-
water flats from New York to Florida and Alabama. —
Plate 13.

Specimens examined from New York (Sartwell; Torrey; A. Gray,
Albany); Pennsylvania, New Jersey and Delaware, shores of the.Dela-
ware (Martindale; Conrad; Zanzinger, 1844; Durand, 1848; Tatnall,
1860; Austin, Hackensack; Smith, 1867); Maryland (Canby, Saulisbury,
1863; Porter, Port Dupont); District of Columbia (S. Watson, 1871;
Seaman, 1872); Virginia (Dr. Robbins, shores of the Potomac; L. F.
Ward, Custis Spring, 1877 and 1878); North Carolina (M. A. Curtis,
1843); Georgia (Baldwin); Florida (J. D. Smith, 1884, Magnolia, Clay

Co.), Alabama (Charles Mohr, shores of Mobile Bay). Flowering from
July to September.

S. suBULATA NATANS (Michx.).— S. natans Michx. FI.
Bor. Am. 2: 190 (1803). WS. natans var. lorata
Chapm., Flora S. U. S., 449 (1860).

Larger, submerged plant with ovate-oblong, elliptical, or
subcordate, or truncate-hastate, floating leaves, with broader
and longer flattened phyllodia; scape and pedicels elongated,

18

SAGITTARIA AND LOPHOTOCARPUS. 45

the flowers opening at the surface of the water or raised
above it; bracts acuminate; fruiting head larger, 8 mm. in
diameter ; filaments longer; achenia more numerous, 2 mm.
long, 5-to 7-crenately crested.— Flowering from August
to October. In fresh water ponds and brackish tide waters
along the coast from South Carolina to West Florida.
Specimens from the coast of Florida have the petioles,
phyllodia and floating scape much elongated, the latter
often 8 to 10 dm. long. Figured from a specimen col-
lected in West Florida by Dr. Leavenworth with both
the floating leaves ascribed to natans, and the ribbon-like
phyllodia of Jorata. — Plate 13.

Specimens examined from ‘Carolina’? (Bernhardi Herbarium);
“ Southern States’? (Read, 1856; Baldwin); South Carolina (Ravenel,
Santee Canal); Alabama (Mohr); Florida (Croom, 1835, Quincy; Dr.
Chapman, 218, Apalachicola, and many specimens without date or
locality in many herbaria; Dr. Leavenworth, at St. Marks, “‘ where the
water varies in depth with the tide from 2 to 5 or 6 feet,” and Ft. Duane,
East Florida; Keeler, Mayport and Jacksonville; and Curtiss, 2747, 1878,
Jacksonville).

S. SUBULATA GRACILLIMA (S. Wats.) J. G. Smith, in
Mem. Torr. Bot. Club, Vol. 5: 26 (1894). WS. natans
var. (?) gracillima S. Wats. in A. Gray, Man. Ed.
6, 556 (1890). S. natans Engelm. Bull. Torr. Bot.
Club, 9: 4 (1882).

Submerged aquatic; leaves 6 to 12 dm. long, usually
consisting of narrow flexuous phyllodia 2 mm. wide,
indistinctly 1-nerved toward the apex, or widening into a
3-nerved, lanceolate blade 3 to 4 cm. long, 6 to 8mm. wide;
scape simple, terete or flattened, as long as the leaves, bear-
ing 2 to 6 verticils; pedicels 3 to 10 cm. long ; only 1 or 2
flowers appearing at the surface of the water at a time, 15
to 20 mm. across; inflorescence at length 4 to 5 dm. long ;
bracts acute, soon evanescent; the immature achenium
broadly obovate, with a stout oblique beak. The achenia
ripen under water and mature specimens have not yet been
collected.— In deep water of streams in Eastern Massa-

19

46 MISSOURI BOTANICAL GARDEN.

chusetts and Rhode Island. Flowering late in July.—
Plate 14.

Specimens examined from Massachusetts (Hitchings, July 22, 1870,
Milton, and 1877, Readville; Boott, July 28, 1870, Readville; C. E. Faxon,
July, 1877, Charles and Neponset Rivers, and 1882, Dedham; Edwin

Faxon, 1891, Readville; Deane, 1890, Milton); Rhode Island (Collins,
1890, Ten Mile River).

S. FILIFORMIS n. sp.

Submerged aquatic, with linear, filiform phyllodia 1.5
to 6 dm. long, 2 10 3 mm. wide; scape slender, filiform, 6
to 12 dm. long, branching from all but the uppermost
verticils; bracts lanceolate, acuminate, 3 to 5 mm. long,
at length evanescent; pedicels and branches of the scape
filiform, 4 to 10 em. long; 1 or 2 flowers of the lowest
verticil pistillate, the rest all staminate; verticils 6 to 10,
remote; flowers 10 to 12 mm. wide; sepals ovate, obtuse,
scarious margined ; petals white; stamens 7; filaments over
twice as long as the anthers; mature achenia not yet
collected; ovary obovate, equaled by the slender oblique
style. — Floating in still water, Dog River, Mobile County,
Alabama, Mohr, August 14, 1893. One of the specimens
is proliferous at the lowest verticil of the scape, throwing
out roots, phyllodia and flowering pedicels from the axils
of the bracts.— Plate 15.

More or less imperfect specimens of this are: Curtiss 31, Long Moss
Spring, Jackson Co., and Jacksonville, Fla.; plants collected by Chap-

man, 1842, near Apalachicola; and by Keeler near Jacksonville. These
were all distributed as S. graminea.

§ § § Integrifoliae. Leaves entire or hastate, from linear to ovate,
neither sagittate nor floating; bracts free or connate at the base;
stamens 12 or more; filaments slender, or dilated and pubescent.

* Fertile pedicels neither thickened nor reflexed in fruit,

+ Achenium falcate, obovate, with an oblique beak; leaves lanceo-
late, entire, pseudo-penninerved; stamens more than 20; filaments slen-
der as long as or longer than the linear oblong anthers.

20

SAGITTARIA AND LOPHOTOCARPUS. 47

S. nancrroxia Linn. Amoen. 5: 409(1760). WS. lancifolia
var. major Micheli, in DC. Monog. Phan. 3; 73
(1881).

Rigid, erect; leaves 4 to 9 dm. high, the blades coria-
ceous, 30 to 60 cm. long, 5 to 10 or 20 cm. wide, acute or
long acuminate, 5-to 9-nerved, gradually narrowing to the
base and decurrent into the petiole, pseudo-penninerved
(the primary nerves separating from the thickened midrib
above the base), scape erect, striate, branched, exceeding
the leaves, often 1.5to 2m. high; verticils very numerous;
fertile pedicels ascending, shorter than the sterile, 1 to 3
cm. long; bracts glabrous, free, ovate, acute or acuminate,
10 to 25 mm. long, before flowering enfolding the buds at
the apex of the scape, with the free tips turned outward ;
flowers ample; sepals smooth; petals obovate, white ; sta-
mens numerous; anthers oblong, twisted after dehiscence,
about as long as the cobwebby-pubescent filaments; stam-
inodia present in the pistillate flower; achenium 2 to
3 mm. long, winged on both margins, a short vertical resin
passage on each side; beak short.— From Florida (and
Texas, fide Micheli) southward, throughout the West
Indies, Mexico, Central and South America. Very variable
as to its achenial characters, and the length of bracts and
stamens.— Plate 16.

Specimens examined from Florida (Canby, 1869, Hibernia; Powell,

1872; Keeler, Jacksonville; Bates, 1889, Merritts Island; Simpson, 1890;
Hulst, 1891, De Land; Curtiss, 2742, Jacksonville; Buckley; Burrowes).

S. LANCIFoLIA FALCATA (Pursh) J. G. Smith, in Mem.
Torr. Bot. Club, Vol. 5: 25 (1894). WS. falcata
Pursh, Fl. Am. Sept. 397 (1814).

Rigid, erect, smaller than the species; leaves narrower,
nerves 3 to 5; scape less branched, often simple; fertile
pedicels shorter, 8 to 20 mm. long; bracts short, 5 to 8
mm. long, ovate, obtuse or sub-acute, more or less granu-
lar papillose on the back; filaments cobwebby pubescent,
longer than the anthers; achenium about 2 mm. long,

21

48 MISSOURI BOTANICAL GARDEN.

winged on both margins, the resin passages more promi-
nent.— From Delaware to Florida, Texas and Mexico.—
Plate 16.

Specimens examined from Maryland (Canby, 1872, Saulsbury; Rusby,
1889, Stockton); Delaware (Canby, 1887, Seaford); Virginia (Rugel);
North Carolina (McCarthy, 1885, Wilmington); South Carolina
(Ravenel, Santee Canal); Carolina (Pursh, achenia from the type, in
Herb. Engelmann, ex Herb. Hooker); Florida (Deane; Chapman, Apa-
lachicola; Canby, 1879, Hibernia); Alabama (Mohr. 1879); Mississippi
(Hilgard, 1859, sea coast marshes; Tracy, 2175, 1893, Biloxi); Louisiana
(Lindheimer, 1839, Lake Pontchartrain; Dr. Hale); Texas (Lindheimer,
1842, Galveston Bay) ; Mexico (Rovirosa, 675, 1889). Many Mexican and
Central American specimens have connate bracts, but are in other par-
ticulars identical with this variety.

S. ampieua n. sp.— S. lancifolia Kellerman Anal. Flora
Kansas, 178 (1888).

Erect, 3 to 6 dm. high; leaves lanceolate, long-acute,
gradually narrowing to the base, not decurrent, 12 to 20
em. long, 5-nerved; scape erect or ascending, simple or
branching from the lowest whorl, equaling or exceeding
the leaves; verticils numerous, 8 to 12 or more; bracts
lanceolate, acuminate, connate at the base, sparsely
papillose on the nerves, 10 to 15 mm. long; fertile
pedicels ascending, 2 to 4 em. long, longer than the
sterile; stamens 20 to 25, the slender glabrous filaments
longer than the oblong anthers; fruiting head depressed
globose, 12 to 15 mm. in diameter; achenium 2 mm.
long, narrowly falcate, almost beakless, very narrowly
winged on both margins, the sides smooth, with 2 or 3
resin passages.— In ponds. Central Kansas and Oklahoma.
Apparently first collected by Dr. Butler, 1875, in the Indian
Territory. Also collected in McPherson Co., Kansas, by
J. E. Bodin, July, 1887; and by A.S. Hitchcock, July 29,
1892, in fruit.

This species has been distributed as S. lancifolia but
differs in having papillose, connate, lanceolate bracts,
glabrous filaments, and almost beakless and wingless
falcate achenia.— Plate 17.

22

6 Ea eR ier eae AT

SAGITTARIA AND LOPHOTOCARPUS. 49

+ + Achenium narrowly obovate, with a long, stout, erect beak;
leaves hastate, veins free to the base; filaments dilated, pubescent;
anthers short-oblong; fertile flowers subsessile.

S. riema Pursh, Fl. Am. Sept. 397 (1814). S. heter-
ophylia Pursh, |. c. 396 (1814). &. heterophylla var.
angustifolia Engelm. in A. Gray; Man. Ed. 5, 494
(1867), and var. elliptica Engelm. |. c. 494 (1867).

Monoecious, erect or ascending, 1 to 8 dm. high; leaves
5 to 20 cm. long, varying from elliptical, linear or lanceo-
late to broadly ovate, apex acute or abruptly rounded, base
entire, or cordate, or with 1 or 2 narrow divergent lobes,
veins 7 to 9; scape simple, weak, sinuous, at length decum-
bent, shorter than the leaves, usually geniculate at the
lower verticil; bracts ovate, obtuse, 4 to 8 mm. long,
deeply connate; 1 or rarely 2 verticils fertile, the pedicels
not exceeding 1 cm. long; filaments usually exceeding the
anthers; achenium narrowly winged on both margins, 3 to
4 mm. long, crested above; fruiting head echinate with
the beaks of the achenia, 8 to 15 mm. in diameter, almost
sessile.— In stagnant or running water, rooting in the mud.
Quebec to Tennessee, west to Minnesota and Nebraska.—
Plate 18.

Specimens differ greatly in size and form of leaf, differ-
ences depending largely on the habitat of the plant. When
growing in deep pools or running streams, the petioles
become thick, rigid and elongated, with long narrowly
lanceolate spongy blades, or the tapering attenuate
phyllodia are bladeless. This is the S. heterophylla rigida
of the manuals and collectors. When growing in shallow
ponds or in simply muddy places, the petioles are weaker,
and the blade elliptical ovate and usually smaller, and
the habit erect or ascending. This form is the 8S.
heterophylla elliptica of collectors. Depauperate plants
from shallow water or simply muddy places, with linear
elliptical oblong leaves, are the S. heterophylla angustifolia
of collectors. Toward the southern limit of its range, the
plants are usually of ranker growth, with larger more often

4 23

50 MISSOURI BOTANICAL GARDEN.

hastately lobed leaves. In all these conditions there is
much variation in form and size of achenium, stamen
length, and in the size of the fruiting head. They occur
together throughout the range and seem to be the result
of different habitat, rather than to mark distinct forms or
varieties.

Specimens examined from Vermont (Pringle, 1879; Faxon, 1882;
Morong, 1885); New York (Dr. Pitcher, 1829; Sartwell, Penn Yan;
Ward, 1879; Clinton, in the rapids above Goat Island, Niagara; Engel-
mann, 1840, 1856; Robbins, 1866; A. Gray); Connecticut (Wright) ;
Pennsylvania (Porter; Guttenberg, 1879; James, 1860; Britton, 1889) ;
New Jersey (James, 1848; Porter, 1860; Morong, 1891; Beyrich); Dela-
ware (Tatnall, 1860); Ohio (Riehl, 1835, 1836; Engelmann; Riddell,
1868; Werner, 1887); Kentucky (Rafinesque); Tennessee (Gattinger,
1883); Illinois (Riehl, 1846; Engelmann, many dates and localities;
Brendel, 1873; Eggert, 1877; Hitchcock, 1891; Bush, 1892); Iowa
(Hitchcock, 1888); Nebraska (Williams, 1890; Smith, 1890; Rydberg,
1891; Bates, 1891, 1893, Sand Hills); Minnesota (Sandberg, 1885, 1890,
624 and 638, 1891; Herrick, 1878; Bailey, 542, 1886; Ballard, 273, 588, 814,
1891; Sheldon, 321, 1891, and 3149, 3211, 1892); Wisconsin (Houghton,
1831; Robbins, 1866) ; Michigan (Bigelow, 1865) ; Ontario (Macoun, 1868,
1884); Canada (Pursh).

+ + + Achenium obovate, with a short lateral beak; filament dilated
at the base, pubescent; bracts connate, glabrous.

S. graminea Michx. Fl. Bor. Am. 2: 190 (1803). S. acuti-
folia Pursh, Fl. Amer. Sept. 397 (1814). WS. simplex
Torr. Comp. 356 (1826), not Pursh. S. sagittifolia
var. simplex Hook. Fl. Bor. Amer. 2: 167 (1840).
S. Purshii Kunth, Enum. 8: 160 (1841). SS. stoloni-
fera Engelm. and Gray, Boston Journ. Nat. Hist. 5:
234 (1847).

Monoecious or dioecious, 1 to 6 dm. high; leaves lanceo-
late to ovate elliptical, acute at both ends; or very rarely
truncate or hastate with short divaricate lobes at the base,
1 to 6 cm. wide, 5 to 15 em. long, 3-to 5-nerved, the
nerves free to the base; scape simple, slender, erect, with
few remote, or many verticils sometimes approximate ;
bracts ovate, acute, 3 to 5 mm. long, connate to the middle ;
fertile and sterile pedicels about equal, spreading, 1 to 3 cm.

24

SAGITTARIA AND LOPHOTOCARPUS. 51

long; stamens about 18; anthers exceeding or equaling
the filaments ; achenium 1.5 mm. long, dorsally crested and
obliquely unicostate or winged on the sides, with or without
subepidermal resin passages; fruiting head small, 5 to 10
mm. in diameter. Phyllodia, when present, flattened,
linear lanceolate, acute, 8 to 30 cm. long, 1 to 2 cm. wide.
Plants producing winter tubers and growing in tufts at the
nodes of horizontal stolons.— In shallow ponds and marshes,
or slowly running streams, or in simply muddy places.
From New Foundland westward to the Missouri river, and
south to Florida and Texas. In the southern portion of its
range it passes into the next variety. — Plate 19.

Specimens examined from Canada (Michaux; Macoun, 1883, Cape
Breton; Britton and Timmerman, 1889, L. Muskoka); New Brunswick
(Macoun, Little Tobique River, 1884); New Hampshire (Deane, 1884,
Shelburne); Massachusetts (Robbins, Uxbridge, 1865 and 1866; A. Gray,
Newburyport, 1866; G. E. Stone, Worcester); New York (Dr. Sartwell,
Penn Yan; A. H. Curtiss, Lake Oneida; J. A. Paine, Herkimer Co.,
1854; Ward, South Bend, 1879; Vasey, Lake George, 1882; Morong, New
York City, 1891); New Jersey (Durand; Bergen, 1845; Britton, Branch-
ville, 1886); Delaware (Tatnall, 1860; Durand; Brinton, Townsend,
1890; James, 1867); Pennsylvania (Moser, Bethlehem, 1832; Wolle,
Bethlehem, 1856; Porter, Harrisburg, 1863; James, Philadelphia,
1861 and 1867); West Virginia (Millspaugh, Hinton, 1891); Michi-
gan (Dr. Bigelow, Detroit, 1865); Ontario (Macoun, Petersboro
Co., 1878); N. W. Terr. (Macoun, 208, Eagle Hills, 1879); South
Dakota (Williams, Big Stone Lake, 1892); Iowa (Hitchcock, Consforth,
1890); Nebraska (Swezey, 136, Hardy; Rydberg, Mead, 1890; Smith,
Lincoln, 1890; Bates, Holt Cr., 1892); Kansas (Oyster, Paola, 1891);
Missouri (Engelmann, many dates and collections, 1832 to 1880,— speci-
mens collected at St. Louis in 1857 have the leaves truncate or divari-
cately hastate lobed at the base; Eggert, St. Louis, 1875; Pammel,
Valley Park, 1886; Bush, Courtney, 1891; Jackson Co., 1892); Illinois
(Engelmann, many dates and collections from 1832 to 1880; Meyer, East
St. Louis, 1838; Vasey, Ringwood; E. Hall, Athens, 1861; Eggert, St.
Clair, 1875; S. Watson, Quincy, 1885; Hitchcock, Fish Lake, 1890); Ken-
tucky (Dr. Short, Lexington, 1835 and 1840), North Carolina (Schweinitz) ;
South Carolina (Ravenel, Santee Canal); Georgia (Curtis, Atlanta
1850); Alabama (Mohr, Mobile, 1883, 1884); Southern States (Dr.
Leavenworth; Dr. Walsh) ; Florida (Alden, Ft. King; Harding, Baker Co.
1887; Hulst, DeLand, 1891); Louisiana (Carpenter, 1839; Hale); Texas
(Lindheimer, West of the Brazos, 1839, and 183, 1843 at Houston, S.
stolonifera Engelm. and Gray, plants with linear lanceolate, acute acu-
minate phyllodia 1 cm. wide, 3 to 4 dm. long; Bigelow, San Bois, 1853;

25

52 MISSOURI BOTANICAL GARDEN.

Drummond, 434, 435 and 436; Dr. Bigelow, Whipple’s Exped., Shawnee
villages, 1853).

S. GRAMINEA CYCLOPTERA DN. var.

Slender, erect, 2 to 8 dm. high, from a horizontal or
oblique rhizome; leaves linear lanceolate, tapering gradu-
ally at both ends, or reduced to slender attenuate phyllodia ;
scape simple or branching from the lowest verticil; inter-
nodes longer than the fertile pedicels; bracts and stamens
as in the species; achenium 1.5 mm. long, with an abrupt
dorsal crest, an arched wing, and a median vertical resin
passage on each side, or when maturing under water only
costate or wrinkled.— From South Carolina to Florida and
Louisiana.— Plate 20.

Specimens examined from South Carolina (Dr. Mohr, Nov. 1893,
Ridgeland); Florida (Dr. Chapman, 234, in Herb. Torrey, and Apalachi-
cola; Curtiss, 1876 and 2746, pine barren ponds, Duval Co.; Keeler, near
Jacksonville); Alabama (Mohr, April, 1886, May, 1893, and Aug. 1893,
Mobile); Louisiana (Hale, pine barren ponds, in Herb. Engelmann).

S. GRAMINEA CHAPMANI 0. var.

Three to five dm. high; leaves narrowly lanceolate, acute,
tapering at base into the petiole, 2 to 3 cm. wide, 15 to 20
em. long; scape weak, branched or simple, the fertile
flowers numerous; bracts lanceolate, acute or acuminate,
6 to 12 mm. long, slightly connate at the base; stamens as
in the species; fertile pedicels 1.5 to 3 cm. long; fruiting
head 5 mm. in diameter; achenium almost beakless, 1 mm.
long, with a narrow dorsal crest, the sides not costate nor
winged. Phyllodia oblanceolate, long-acute, 1 to 2 cm.
wide, 10 to 30 cm. long.— In creeks and stagnant ponds,

West Florida and Alabama. Dedicated to the venerable
southern botanist, Dr. Chapman, by whom it was first col-
lected in 1862, ‘‘in a creek on the road to Marianna, 3 or
4 miles from Ochesee, West Florida.’’ Also collected by
Dr. Mohr, 1880, and March, April, May and June, 1884,
in the vicinity of Mobile, Alabama. — Plate 21.

26

SAGITTARIA AND LOPHOTOCARPUS. 53

S. cristata Engelm. in Arthur’s Contributions to the
Flora of Iowa, no. 5, 3 (1882). Proc. Davenport
Acad. Nat. Sci. 4: 29 (1886).

Monoecious, slender, erect, 30 to 75 cm. high, submersed
or in shallow water; leaves linear-lanceolate or rarely
elliptical lanceolate, 6 to 10 cm. long, 1 to 2 cm. wide,
thick, spongy, triquetrous in cross section; petioles long
and slender; scape simple, erect, the flowers at or above the
surface of the water; verticils 4 to 6, the lowest fertile;
bracts 5 to 7 mm. long, acute; fertile pedicels ascending,
1.5 to 3 cm. long; flowers about 2 cm. across; sepals ovate,
obtuse, scarious tipped, rough on the veins; stamens about
24; filaments dilated subulate, pubescent at the middle,
longer than the anthers; achenium 3 mm. long, dorsally
crenately crested, ventral margin straight winged, an inter-
rupted crenate crest on each side, beak short; fruiting head
globose, 15 to 20 mm. in diameter. Often with tufted
phyllodia, flat, linear lanceolate, acute, 1 dm. long. In
deep water, the petioles and scape are spongy and rigid.
The phyllodia grow from the nodes of the stolons along the
muddy bottom, to the depth of 1 or 2 m. or more.— Ponds
and lakes, northern Iowa, southern Minnesota and (?)
western New York. Engelmann’s type specimens were
collected by R. I. Cratty, Emmett Co., Iowa, 1881, —
Plate 22.

Specimens also examined from Minnesota (Sandberg, 623, Minnetonka,
1891; Herrick, 1878; Aiton, Minneapolis, 1891; Ballard, 603, Scott Co.,
1891; Sheldon, 705, White Bear, 1891; Wood, Chub Lake, 1889); New
York (?) Dr. Sartwell in Herb. Gray).

S. MACROCARPA, 0. sp.

Monoecious, slender, 3 to 4 dm. high, suberect; phyl-
lodia flexuous, 25 to 30 cm. long, gradually expanding
above to a linear lanceolate blade 5 to 7 cm. long, 3 to 5
mm. wide; scape slender, equaling or exceeding the leaves,
simple; verticils few, 3 to 5, about the length of the pedi-
cels apart; bracts ovate, acute, 3mm. long, connate; 1 to
3 flowers of lowest verticil perfect, their pedicels slender,

27

54 MISSOURI BOTANICAL GARDEN.

ascending, 6 to 12 mm. long; sterile pedicels 10 to 15 mm.
long; stamens about 12, the dilated pubescent filaments a
little longer than the short oblong anthers; fruiting head
about 12 mm. in diameter; achenia oblong obovate, 3 mm,
long, broadly winged on both margins, smooth or faintly
unicostate, and with 2 or 3 resin passages on each side,
beak short and oblique.—Margin of ponds in South Car-
olina. Collected by M. A. Curtis in South Carolina
(Engelm. Herbarium). Related to éeres and cristata.—
Plate 23.

S. reres S$. Wats. in A.Gray, Man. Ed. 6, 555 (1890).

Monoecious, with acute, attenuate, terete, usually nodose
phyllodia 10 to 30 em. long, very rarely bearing a short
linear blade; scape slender, erect, simple, 15 to 45 cm.
high; bracts ovate, obtuse, 3 mm. long; verticils few, 1
to 8, 1 or 2 flowers of the lowest fertile; fertile pedicels
slender, ascending, 10 to 25 mm. long, exceeding the ster-
ile; flowers small, 12 to 15 mm. across; stamens 12, the
filaments dilated, pubescent, shorter than the anthers;
mature achenium rounded-obovate, 2 mm. long, with a
short, erect beak, the ventral margin wing-angled, dorsal
margin and _ sides crenately several- (7 to 11) crested;
fruiting head 8 to 12 mm. in diameter.—In shallow water,
Massachusetts to South Carolina. Mature achenia collected
only by Walter Deane, at Hyannis, Mussachusetts, August,
1888. The immature achenia are dark brown, and barely
costate laterally. The fruits usually ripen under water,
and do not then assume the same characters as when ripen-
ing above water.— Plate 24.

Specimens examined from Massachusetts (Farlow, 1881; Brewster) ;
New York (E. 8. Miller, Wading River, Long Island, 1871); New Jersey
(Torrey, 1833, pine barrens). Specimens marked A, collected in North
Carolina by M. A. Curtis, are intermediate between this species and
macrocarpa. They have the habit of ¢eres, but with larger achenium
(2.5 mm.) with smooth sides, and more oblique beak.

+ + + + Achenium short obovate, with a short lateral beak; leaves
lanceolate, entire; filament slender, glabrous; bracts connate, densely
papillose.

28

SAGITTARIA AND LOPHOTOCARPUS. 55

S. paprtLosA Buchenau, Abh. Nat. Ver. Bremen, 1: 44
(1868). S. lancifolia var. papillosa (Buch. ) Micheli,
in DC. Monog. Phan. 3: 74 (1881).

Monoecious, slender, erect, 3 to 5 dm. high; leaves
linear lanceolate, acute, gradually narrowing into the slender
petiole, 10 to 15 cm. long, 3 to 95 nerved, pseudo-penni-
nerved toward the base; scape very slender, erect, terete
or obtusely 3-angled, striate, simple or branching from the
lowest whorl; bracts obtuse, 3 to 4 mm. long, densely
granular papillose on the back; fertile pedicels ascending,
1 to 2 cm. long, the sterile 1.5 to 3 cm. long; sepals ovate,
obtuse, papillose; flowers 20 to 30 mm. across; stamens
18, about 3 mm. long, the linear oblong anthers equaled
or exceeded by the slender glabrous filaments; achenium
1.5 mm. long, with a tumid dorsal crest and a single
vertical resin passage on each face; fruiting head globose,
6 to 10 mm. in diameter. —In shallow ponds, Texas and
Louisiana. — Plate 25.

This species was placed by Micheli as a variety of lanct-
folia, but is more closely related to graminea and perhaps
ambigua. Fully matured achenia have not as yet been
collected, or at least are not present in any of the Ameri-
can herbaria examined.

Specimens examined from Louisiana (Dr. Hale, Pine Barren, Alex-
andria); Texas (Drummond, 423; Lindheimer, Houston, 1842, a speci-
men with the foot of the plant bulbous, covered with the bases of the
dead leaf sheaths, and 1843, 182; Elihu Hall, 623, 1872, Houston; G.
C. Nealley, 1884, Beaumont). Lindheimer says of his Houston specimens
that the successive annual tubers are formed one over another, so that a
plant often has two or three of the old exhausted ones attached. The
leaves are sometimes reduced to linear phyllodia which “resemble
flowerless scapes.”’

* * Fertile pedicels thickened or reflexed in fruit.

+ Achenium obovate; leaves hastate or entire; bracts connate to
above the middle, glabrous.

S. PLATYPHYLLA (Engelm.). S. graminea var. platyphylla
Engelm. in A. Gray, Man. Ed. 5, 494 (1867); J. G.
Smith in Mem. Torr. Bot. Club, Vol. 5: 25 (1894).

29

56 MISSOURI BOTANICAL GARDEN.

S. recurva Engelm. Patterson, Check-List, 130
(1887), name only.

Monoecious, erect, 2 to 5 dm. high; leaves subcoria-
ceous, 5 to 7 pseudo-penninerved from near the base, 5 to
15 cm. long, 2.5 to 9 cm. wide, broadly ovate, or ovate
lanceolate, or ovate elliptical, obtuse or acute, abruptly
short acuminate at the apex, rounding or gradually narrow-
ing to the base, or rarely subcordate, auriculate or hastate ;
scape simple, rather weak, ascending, smooth, usually
shorter than the leaves; 2 to 4 lower verticils fertile; fer-
tile pedicels thick, divaricate in flower, weak and reflexed
in fruit, 12 to 25 or rarely 40 mm. long, mostly longer
than the sterile; bracts broadly ovate, 3 to 6 or 8 mm.
long; flowers 2.5 to 3 cm. across; sepals ovate, obtuse,
scarious at the tip; stamens about 21, the dilated pubes-
cent filaments a little longer than the short oblong anthers;
achenium 2 mm. long, obliquely obovate, winged on both
margins, with a narrow dorsal crest, laterally with an
oblique wing-angle extending downward from the base of
the short horizontal or oblique beak; short resin passages
usually present between the lateral angle and the crest;
fruiting head 9 to 12 mm. in diameter; linear oblong or
oblanceolate phyllodia, 10 to 15 mm. wide, 4 to 6 dm. long,
often present.— In swamps and ponds from Texas to
Mississippi and northward to the “‘ sunken lands ’’ of South-
east Missouri.— Plate 26.

Specimens examined from Texas (Lindheimer, 713, New Braunfels,
1847 and 1851; Reverchon, 1875, and “Curtiss, 2746 *’’ 1881, and 939,
1880, from Dallas; E. Hall, 624, Hempstead); Louisiana (Fendler, 1846,
New Orleans; Dr. Hale, Alexandria and Bayou Robert); Mississippi (A.

Wood, 1860); Arkansas (Eggert, 1893, Paragould); Missouri (Bush,
1893, Kennett, Dunklin Co.).

S. Mexicana Steudel, Nom. 2: 491 (1841). &. macro-
phylla Zuce. in Abh. Kén. Bayr. Akad., 289 (1832),
not Bunge; Micheli in DC. Monog. Phan. 3: 71
(1881). See below, p. 33.

This species is said by Micheli to be represented in the herbarium of

30

SAGITTARIA AND LOPHOTOCARPUS. 57

the St. Petersburg Academy, by a specimen collected by Dr. Engelmann
at New Orleans. There is in Herb. Engelmann, a very long-pediceled
specimen of platyphylla from Louisiana which has been mistakenly
called Mexicana. This is undoubtedly the same as the plant in the St.
Petersburg Herbarium. As far as known, S. Mexicana does not occur in
the United States.

++ Achenium quadrate cuneate, beak short and triangular; leaves
entire, pseudo-penninerved; bracts connate at the base.

S. Sanrorpi, Greene, Pittonia, 2: 158, 1890.
Monoecious, 9 to 15 dm. high, rigidly erect or ascend-
ing; petioles rigid, 2 to 4 cm. thick at the base, obtusely
triquetrous; blade linear lanceolate to oblong lanceolate,
acute, tapering into the spongy petiole; scape simple, stout ;
bracts triangular, obtuse, 5 to 7 mm. long; verticils numer-
ous, approximate, 2 to 3 lower ones fertile; fertile pedi-
cels 15 to 20 mm. long, reflexed in fruit; sepals ovate,
acute, 4 to 6 mm. long; stamens about 20, the oblong
anther longer than the dilated glabrous filament; achenia
numerous, 2 mm. long, winged on both margins, the sides
reticulate, with or without a costate lateral angle; beak
short, oblique, triangular; fruiting head 12 to 14 mm. in
diameter. Triangular, alternate, spongy phyllodia several
decimeters long are present when the plant is nearly or
quite submerged. — Growing in the marshes along the
lower San Joaquin River, California. Specimens examined
from Stockton, Cal., collected by J. A. Sanford, 1891, and
1893. We are indebted to Mr. Sanford for fresh material
of this species, and for water color habit sketches drawn
under his direction by Dr. Hudson of Stockton.— Plate 28.
B. Fertile sepals erect and accrescent; pedicels of the pistillate

flowers thickened and reflexed in fruit; petals cream white with a
brownish purple spot at the base. Introduced species.

S. Montevipensis Cham. and Schlecht. Plant. Roman-
zoff. Linnaea 2: 156 (1827). Micheli in DC. Monog.
Phan. 3: 75 (1881).

Monoecious; petioles stout, rigid, erect, ascending;
leaves sagittate, acute or obtuse, 1 to 5 dm. long and
31

58 MISSOURI BOTANICAL GARDEN.

broad, basal lobes acute, acuminate, widely divergent,
glabrous above, sparsely scabrous on _ the nerves
below; scape usually simple, stout, often 6 to 8 cm.
in diameter at the base; verticils mumerous, more
or less approximate, 2 to 4 lower ones fertile; bracts con-
nate at the base, lanceolate, long acuminate, the upper un-
developed portion of the scape appearing comose with their
projecting tips; staminate pedicels slender, ascending, 20
to 30 mm. long, longer than the fertile; flowers large, the
sepals broadly ovate, obtuse, 10 mm. wide, 15 mm. long;
petals obovate, larger than the sepals; stamens very many,
the narrow glandular pubescent filaments longer than the
linear oblong anthers; achenia 2 to 3 mm. long, narrowly
quadrate obovate, winged on both margins, with a short
slender, oblique beak, and a prominent sub-epidermal
resin passage on each side above; fruiting head large, 15
to 30 mm. in diameter.— A South American species estab-
lished as a ballast plant in California and North Carolina.
Specimens examined from California (J. A. Sanford, July
1893, Stockton); North Carolina (Gerald McCarthy,
September, 1888). — Plate 29.

SPECIES FROM MExIco.
§§ Fluitantes: stamens 12 or 18.

S. DEMERSA 0. sp.

Submerged aquatic; the floating phyllodia linear atten-
uate, 2 to 6 dm. long, 10 to 15 mm. thick at the base, taper-
ing gradually to the obtuse apex; scape simple, weak, rising
to the surface of the water, usually 20 to 30 cm. long;
bracts scarious, deeply connate, acute, 3 to 5 mm. long,
soon evanescent; verticils 4 to 6, the lowest fertile; fertile
pedicels thickened, 10 to 35 mm. long, at length reflexed;
sterile pedicels slender, erect-spreading, 10 to 20 mm. long,
the flowers at length deciduous, filaments dilated, glabrous,
twice as long as the sub-orbicular anthers; fruiting head
5 to 6 mm. in diameter, sub-globose; achenium obovate,

p> £
32

SAGITTARIA AND LOPHOTOCARPUS. 59

plump, 1.5 to 2 mm. long, with a very slender lateral
erect beak, margins broadly winged below the middle, sides
smooth; seed 1 mm. long, broadly obovate, striate. —
Ponds near Guerrero, Chihuahua, Mexico, Pringle, 1367,
Sept. 9, 1887. Distributed as S. graminea forma acuti-
folia. It is related to S. subulata and S. filiformis.—
Plate 15.

§ § § Integrifoliae.

* Fertile pedicels neither thickened nor recurved in fruit.

S. LANCIFOLIA ANGUSTIFOLIA (Lindl.) Griseb. Catal. Pl.
Cub. 218 (1866); Micheli, DC. Monogr. 3: 73. WS.
angustifolia Lindley, Bot. Reg. 14: pl. 1141 (1828).

Much smaller than the type in every part; blade of leaf

very narrow or absent; bracts barely 3 mm. long; sepals 3

to 4 mm.; carpels crested,— ex Micheli, 7. c. 73.— Tical-

tepec, Mex., Liebmann, July, 1841, fide Buchenau, Engler’s

Bot. Jahrb. 2: 487.

* * Fertile pedicels thickened or recurved in fruit.

S. Mexicana Steudel, Nomenclator, 491 (1841). S.
macrophylla Zucc. Abhandl. Bayr. Akad. 1832, 289 ;
Micheli, Monogr. 3: 71.—not Bunge, Mem. Say.
Etrang., 2: 137 (1831).— See above, p. 30.

Monoecious, 3 to 10 dm. high; erect, or the petioles
weak and ascending; leaf 10 to 20 cm. long, 1 to 5 cm.
wide, lanceolate, acute at the apex, entire, abruptly nar-
rowing to the petiole, or hastate with small, nearly linear,
divaricate lobes; veins free to the base; scape simple, erect,
about equaling the leaves; verticils 3 to 5, remote ; fertile
pedicels erect-spreading in flower, 5 to 7 cm. long, exceed-

ing the sterile; bracts broadly ovate, obtuse, 8 to 10 mm.

long, connate nearly to the apex; flowers ample, 3 cm.

across; stamens 21 to 30, 4 to 5 mm. long, the slender
glabrous filaments longer than the oblong sagittate anthers.

The mature achenia have not yet been collected. In

immature fruiting heads, the ovaries are broadly winged,

33

hee eed

60 MISSOURI BOTANICAL GARDEN.

with a long horizontal style, and an undulate dorsal crest.—
Plate 27.

Specimens examined from Mexico, collected by L. Hahn, 1869, in
Herb. Engelmann, ex Herb. A. Braun; and a fragment from the type, in
Herb. Engelmann, ex Herb. Monaco, collected by Karwinsky.

Lopuotrocarrus T. Durand, Index Gen. Phan. 627
(1888). Lophiocarpus Miquel, Ill. Fl. Arch. Ind. 1: part
2, 50 (1870), not Turcz.— Fertile flowers with stamens;
filaments hypogynous.

L. catycrnus (Engelm.) J. G. Smith, in Mem. Torr. Bot.
Club, Vol. 5: 25 (1894).— Sagittaria calycina En-
gelm. including var maxima, var media and var. fluitans
Engelm. in Torr. Bot. Mex. Bound. Surv. 212 (1858) ;
S. calycina var. spongiosa, and S. calycina var. gran-
dis Engelm. in A. Gray, Man. Ed. 5, 493, 494 (1867) ;
Lophiocarpus calycinus Micheli, in DC. Monog. Phan.

8: 61 (1881).

Weak; leaves floating or ascending, halberd shaped,
broader than long, or sagittate, hastate or entire, varying
greatly, from 2 to 20 cm. long by 1.5 to 30 cm. wide,
obtuse or acutish, the basal lobes widely divaricate, ovate,
acuminate ; scape simple, 1 to 3 dm. high, weak, at length
decumbent; bracts short, orbicular, obtuse, those at the
base of the staminate verticils often lanceolate, pointed ; fer-
tile pedicels greatly thickened, reflexed, as long as or much
longer than the more slender sterile ones ; filaments slightly
roughened; achenia obovate, 2 mm. long, narrowly winged
on both margins, a very short resin passage at the base of
the beak above, beak about } as long as the achenium, tri-
angular, horizontal. A very variable species according to
the habitat ; petioles and scapes rather spongy, as are the
phyllodia when present; phyllodia sometimes strongly
nodose.— From New Brunswick to S. Dakota and Califor-
nia, and southward.

Specimens examined from New Brunswick (Fowler, Bass R., Kent
Co., July, 1870); Maine (Swan, Kennebeck, Sept. 1859); Massachusetts

34

SAGITTARIA AND LOPHOTOCARPUS. 61

(Boott, Woburn Pond, 1863); New Jersey (Parker, Camden, Oct. 1877;
Austin, Hackensack R., Aug. 1861); Delaware (Tatnall, Wilmington,
1861); Virginia (Coville, Colonial Beach, July 1890); Michigan (Schneck,
Grand Rapids, 1880, 1881); Wisconsin (Hale, Prairie du Chien, 1861);
Illinois and Missouri, many collections, 1842 to 1893; S. Dakota (Will-
iams, 8, 5, 7 and 14, 1892); Nebraska (Williams, Greenwood, 1890);
Kansas (Hitchcock, Manhattan, 1892); Indian Terr. (Palmer, 331, False
Wichita R. 1868; Carleton, 268, Cherokee Outlet, 1891); Louisiana (Hale,
Alexandria); Texas (Wright, 679; Oct. 1849); New Mexico (Wright,
1899, 1852); California (Parish, 1136, Los Angeles Co., Oct. 1881; San-
ford, Stockton, July, 1893).

L. Guyanensis (HBK.) Micheli, Monogr. 3: 62 (1881).
Sagittaria Guyanensis HBK. Nov. Gen. Sp. 1: 250
(1815).

Leaves floating, broadly ovate, deeply cordate, 3 to 5 cm.
long by 4 to 6 cm. wide, obtuse to slightly emarginate ;
scape erect or flexuous procumbent; bracts broadly ovate,
obtuse; pedicels scarcely exceeding the bracts; filaments
somewhat glandular; achenia very numerous, flattened,
deeply notched, winged on both margins, without resin

passage, the beak scarcely surpassing the margin.— Mexico,
etc., fide Micheli, /. c. 63.

EXPLANATION OF PLATES ILLUSTRATING THE NORTH AMERI-
CAN SPECIES OF SAGITTARIA.

The figures were drawn, under the supervision of the
author, by Miss Grace E. Johnson. Nos. 3, 28, and 29, are
from living plants. No. 28 is reduced from a water color
sketch drawn under Mr. J. A. Sanford’s supervision by
Dr. Hudson of Stockton, Cal. No. 2 is from the original
plate published in the Bull. Torr. Bot. Club (1893), kindly
loaned by Dr. N. L. Britton. The remainder are from her-
barium specimens. Detail drawings, unless otherwise speci-
fied, are enlarged ten diameters.

Plate 1, 8. arifolia Nutt.—1, Plant reduced to half size; 2 and 3, lateral
view and cross section of achenium; 4 stamen; 5, plant of S. arifolia
stricta, reduced to half size.

Plate 2, S. cuneata Sheld.— Plant and upper portion of the scape,
natural size; achenium, < 30.

35

62 MISSOURI BOTANICAL GARDEN.

Plate 3, S. latifolia Willd. — 1, Plant reduced one-fourth; 2, portion of
a branching scape, reduced to half size; 8, 4, achenia; 5, cross-section of
achenium; 6, fruiting head, natural size; 7, stamens from the same
flower.

Plate 4, 8. latifolia, Willd., forms a and b.— 1, fertile scape, reduced
to half size; 2, sterile scape, reduced to half size; 8, achenium, form a;
4, achenium, form b, from Washington; 5, achenium, form 0, from §.
California; 6, achenium of S. Sinensis Sims, ex herb. Hooker.

Plate 5, S. latifolia form c.— 1, Plant, reduced to half size; 2, achenium;
3, cross-section of achenium; 4, seed.

Plate 6, 8. latifolia form d.—1, Plant, reduced to half size; 2,
achenium; 38, seed.

Plate 7, S. latifolia form e.—1, Plant, one-fourth natural size; 2,
achenium; 3, cross-section of achenium.

Plate 8, S. latifolia pubescens (Muhl.).— Form a: 1 , leaf, one-half size;
2, achenium; 3, stamen; 4, fruiting head, natural size; details after
Engelmann.— Form b: 5, plant, three-eighths natural size. — Form ¢: 6,
scape, natural size; 7, immature achenium; 8, stamen.

Plate 9, S. Engelmanniana. — 1, Plant, one-half size; 2, achenium; 3,
cross section of achenium; 4, pistil; 5, stamen; 6, leaf, natural size.

Plate 10, 8. longirostra (Micheli) .— Details after Engelmann. 1, plant,
one-third size; 2, leaf, one-half size; 83, achenium; 4, cross section of
achenium; 5, immature achenium from Drummond’s type collection; 6,
young fruiting head, natural size; 7, pistil; 8, stamen; 9, and 10, cross
sections of scape, below and above. Sketches, except no. 4, made from
Canby’s specimens.

Plate 11, S. longiloba Engelm.—1, Plant, one-third natural size; 2,
winter-tuber, with young plant; 8, achenium; 4, cross section of
achenium; 5, seed; 6, stamen.

Plate 12, S. Greggii.—1, Plant, one-sixth natural size; 2, leaf, one-
half natural size; 3, achenium; 4, stamen.

Plate 13, 8S. subulata (L.) Buch.— 1, Plant, natural size; 2, achenium,
lateral and 3, dorsal view; 4, stamen. 8. subulata gracillima (S. Wats.).—
5, Plant, natural size; 6, inflorescence; 7, immature achenium; 8, stamen.

Plate 14, S. subulata natans (Michx.)—1, Plant, one-half size; 2,
achenium, lateral and 3, dorsal view; 4, stamen.

Plate 15, S. demersa.— 1, Plant, one-half natural size; 2, achenium; 8,
stamen; 4, leaf, one-half natural size. S. Jiliformis.— 5, Plant, one-half

’ natural size; 6, pistil; 7, stamen; 8, bracts, « 5.

Plate 16, S. lancifolia L.—1, Plant, one-sixth natural size; 2, achenium.
S. lancifolia falcata (Pursh). —8, Part of scape, natural size; 4, ache-
nium; 5, stamen.

Plate 17, S. ambigua.—1, Plant, one-third size; 2, achenium; 3, sta-
men; 4, bracts, natural size.

Plate 18, S. rigida Pursh.—1, Plant, one-half size; 2, 3, leaves, one-
half size; 4, achenium; 5, cross section of achenium; 6, stamen.

Plate 19, S. graminea Michx.—1, Plant, one-half size; 2, leaf, natural
size; 3, achenium, lateral and 4, dorsal view; 5, seed; 6, stamen.

36

3 RR a aa MISS A

SAGITTARIA AND LOPHOTOCARPUS. 63

Plate 20, S. graminea cycloptera.—1, Plant, one-half natural size; 2,
achenium; 3, cross section of achenium; 4, stamen; 5, bracts, X 2.

Plate 21, S. graminea Chapmani. —1, Plant, one-third natural size; 2,
achenium; 3, stamen; 4, bracts, natural size.

Plate 22, 9. cristata Engelm.—1, Plant, one-third natural size; 2,
achenium, lateral and 3, dorsal view, 4, cross-section; 5, stamen, ripe and
6, effete.

Plate 23, S. macrocarpa. —1, Plant, one-half size; 2, achenium; 3,
stamen.

Plate 24, S. teres S. Wats. —1, Plant, one-half size; 2, stolon, natural
size; 8, achenium, lateral view and 4, in cross section; 5, stamen.

Plate 25, S. papillosa Buch.—1, Plant, one-fourth size; 2, achenium;
8, stamen; 4, bracts, X 6.

Plate 26, S. platyphylla (Engelm.)— 1, Plant, one-half size; 2, ache-
nium; 3, stamen; 4, fruiting scape, one-half size; 5, 6, leaves, one-half
size.

Plate 27, S. Mexicana Steudel.—1, Plant, one-third natural size; 2,
immature achenium; 38, stamen from bud; 4, stamen.

Plate 28, S. Sanfordii Greene.—1, Plant, one-third natural size; 2,
achenium; 3,stamen; 4, leaf, one-half natural size.

Plate 29, S. Montevidensis Cham. and Schl.—1, Plant, one-fourth
natural size; 2, achenium, lateral view and 8, cross-section; 4, stamen;
5, staminate flower, one-half natural size.

37

64

MISSOURI BOTANICAL GARDEN.

INDEX TO SPECIES OF SAGITTARIA.

The references are to the subpagination of the article. Synonyms are in

acutifolia (24)
ambigua 5, 22.
angustifolia (33 )
arifolia 5, 6.
stricta 8.

calycina (34)
Chinensis (9, 12)
cristata 5, 27.
cuneata 5, 8.

demersa 6, 32.

Engelmanniana 5,
15.

falcata (21)
filiformis 6, 20

gracilis (8)
graminea 4, 5 (20)
, 24.
Chapmani 26.
cycloptera 4, 26.
platyphylla (4,
29).
Greggii 5, 17.
Guyanensis (4,35 )

hastata (9)
heterophylla( 4,23)
38

parenthesis.

angustifolia (23)
elliptica (23)
rigida (28)

lancifolia 4, 21( 22)
angustifolia4,33.
falcata 21.
major (21)
papillosa (29)

latifolia 2-5, 7, 8
pubescens 14.

longiloba 4, 5, 16.

longirostra 5, 16.

macrocarpa 5, 27

macrophylla (30,
33)

Mexicana 6, 30, 33

Montevidensis 6,31.

natans (18, 19)
gracillima (19)
lorata (18)

obtusa (9, 11)

papillosa, 4, 5, 29.
platyphylla 5, 29.
pubescens (14)
Purshii (24)

pusilla (18)

recurva (30)
rigida 5, 23.

sagittifolia 7 (2,9)
longirostra (16)
macrophylla(9)
Mexicana (16)
minor (6)
pubescens (14)
simplex (24)
variabilis (9)
vulgaris (9)

Sanfordii 6, 31.

simplex (8, 24)

Sinensis (9, 12)

stolonifera (24)

subulata 5, 18.
gracillima 19.
natans 18.

teres 5, 28.

variabilis (8)
angustifolia (9)
diversifolia (9,

14)
gracilis (9, 15)
hastata (6, 9)
pubescens (14)

REPT. MO. Bot. GARD., VOL. 6. PLATE 1,

SAGITTARIA ARIFOLIA and var. STRICTA.

REPT. Mo. BOT. GARD., VOL. 6. PLATE 38.

'AK,
RY An
aC
‘ ’
WENT
AN AN "f
AN Ne A
4d

7 8 ot
Uf Pass

SAGITTARIA LATIFOLIA.

REPT. MO. BOT. GARD., VOL. 6.

SAGITTARIA LATIFOLIA, a and 8, and 8. SINENSIS.

REPT. MO. BOT. GARD., VOL. 6. PLATE 5.

SAGITTARIA LATIFOLIA, c.

REPT. MO. BOT. GARD., VOL. 6. PLATE 6.

SAGITTARIA LATIFOLIA, @.

REPT. Mo. Bot. GARD., VOL. 6.

PLATE 7.

ff}
' Ah NS

SAGITTARIA LATIFOLIA, e.

REPT. MO. BOT. GARD.

PLATE 8,

» VOL, 6.

SAGITTARIA LATIFOLIA, PUBESCENS.

REPT. MO. BOT. GARD., VOL, 6. PLATE 9. :

SAGITTARIA ENGELMANNIANA.

REPT. MO, BOT. GARD.,, VOL. 6. PLATE 10.

Avil
Fix; m\
: YIN vi >»
; Jf ; HH) ih
a VA, ny / | \
7 Y FI | |
j Hh {|

SAGITTARIA LONGIROSTRA.

REPT. MO. BOT. GARD., VOL. 6. PLATE 1].

a ee

SAGITTARIA LONGILOBA.

REPT. MO. BOT. GARD., VOL. 6. PLATE 12.

iT
i|

SAGITLARIA GREGGIL,

REPT. MO. BOT. GARD., VOL. 6. PLATE 13.

SAGITTARIA SUBULATA and var. GRACILLIMA.

REPT. MO. BOT. GARD., VOL. 6. PLATE 15.

SAGITTARIA DEMERSA and 8, FILIFORMIS.

PLATE 16.

Rept. Mo. Bot. GARD., VOL. (i.

SAGITTARTIA LANCIFOLIA and var. FALCATA.

REPT. MO. BOT. GARD., VOL 6, PLATE 17.

SAGITTARIA AMBIGUA.

PLATE 18.

REPT. MO. Bor. GARD., VOL. 6.

Gee

SAGITTARIA RIGIDA.

PLATE 19,

REPT.

Mo, Bot. GARD., VOL. 6.

SAGITIARIA GRAMINEA.

Rept. Mo. Bot, GARD.» VOL, 6,

PLATE ;

SAGITTARIA GRAMINEA CYCLOPTERA.

oS oe el

REPT. MO. BoT. GARD., VOL. 6.

PLATE 21.

Wit

}
ik
4

i
|

We
YAN
NN
i if

Yih
wi \
W is AX G
WAIN
AK
SAGITTARIA GRAMINEA CHAPMANI.

" mM 1
hs y Ai) Mi
‘ ai

SA

Repr. Mo. Bor. GARD., VOL. 6. PLATE 22,

SAGITTARIA CRISTATA.

REPT. MO. BOT. GARD., VOL. 6, PLATE 23,

SAGITTARIA MACROCARPA.

REPT. MO. BOT. GARD., VOL. 6. . PLATE 24.

SAGITTARIA TERES.

REPT. MO. BOT. GARD., VOL 6. PLATE 25.

SAGITTARIA PAPILLOSA,

REPT. MO. BOT. GARD., VOL. 6. PLATE 26.

SAGITTARIA PLATYPHYLLA.

Rept. Mo. Bor. GARD., VOL. 6. PLATE 27.

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SAGITTARIA MEXICANA.

Reet. MO. Bot. GARD., VOL. 6. PLATE 28,

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SAGITTARIA SANFORDII.

REPT. MO, Bor. GARD., VOL. 6. PLATE 29.

‘NOD WNL NI'VNVGINOTA VIVANLIAT

a a eat ee ae haere ares = ae
Reg 2 et a. %

LEITNERIA FLORIDANA.

BY WILLIAM TRELEASE,

While collecting in the lowlands of southeastern Mis-
souri, in November, 1892, Mr. B. F. Bush discovered a
small tree growing abundantly in the swamp, and collected
specimens of the trunks, branches with staminate catkins,
and a few old leaves. Notwithstanding the incompleteness
of the specimens, Mr. Bush shrewdly located the plant in
Leitneria, a monotypic genus heretofore known with cer-
tainty only from Florida; and a comparison with speci-
mens of L. Floridana in the Garden herbarium, collected
many years ago in Florida by Dr. Chapman and Dr. D. V.
Dean, showed the correctness of the generic determination,
though certain differences in minor points were noticeable.
In April, 1893, he again visited the locality and collected
other material in leaf and with half grown fruit; and
shortly afterward Mr. Henry Eggert gathered it with nearly
mature fruit.

So far as we now know, the tree occurs in the deep
swamps of Butler and Dunklin counties, where it is asso-
ciated with plants of distinctly southern range, such as
Taxodium distichum, Acer rubrum Drummondii, Nyssa
uniflora, Planera aquatica, and Polygonum densiflorum.
As I have convinced myself by personal observation at
Kennett and Neelyville, it grows in rich swamp soil in
sloughs and similar places, which never become dry and
where there is usually from six inches to two or three
feet of water; and Mr. Bush states that along the St. Francis
River it is most frequent in water from three to five feet
deep, where it is rooted in the basal mass of Polygonum
densiflorum, the common swamp smartweed of that region,
which occurs in dense growths often forming a floating

65 5 1

eee

66 MISSOURI BOTANICAL GARDEN,

accumulation on which in places a man may walk. Certain
of these sloughs are almost exclusively occupied by the
Leitineria, which has a spreading root system confined to
the surface layers of the soil. Apparently suckers arise
from some of the roots, as is the case with the Ailanthus
and White Poplar, but I have not been able to actually
trace these young shoots to the older plants, though their
root system is usually developed out of proportion to their
size.* The impression made on one by such a Leitneria
swamp is that of a tangle of coarse bushes from five to ten
feet in height, but on closer observation it is evident that
each stem rises separately from the soil or water, so that
the plant lacks the clustered bushy habit which distin-
guishes a shrub from a small tree, and it not infrequently
attains a height of fifteen to twenty feet and forms a trunk
from three to five inches thick toward the base, where it
gradually increases in thickness as do many other swamp
trees.

Such arborescent specimens have a clear cut trunk below
and are loosely branched above, with the ascending ultimate
twigs commonly as thick as a lead pencil. Its bark is gray
below and rather smooth, usually mossgrown where wet.
The twigs incline to brown, or in the case of suckers are
almost orange colored, and are marked by numerous slightly
prominent lenticels usually of a lighter gray. During the
first year they are densely pubescent with ascending dingy
hairs, some of which persist during the second year, but
ultimately they become glabrous. The leaf scars are five-
ranked, rather uniformly distributed along the twig,
slightly elevated, and of a general crescent form with
obtuse angles and a not infrequent trilobation. Each con-
tains three relatively large bundle scars. No stipule scars
are to be seen. :

The small persistent terminal bud is broadly conical and

* Apparently connected with this mode of spreading, is the preponder-
ance of one sex in each swamp; a fact which Dr. Dean also informs me
he has observed in Florida.

2

LEITNERIA FLORIDANA. 67

protected by about a dozen obtusely triangular gray-tomen-
tose scales, which evidently represent undeveloped leaves
and some of which usually persist for a year or two at the
annual nodes, ultimately leaving rings of narrow transverse
scars marking the site of former winter buds. On mature
plants the upper axillary buds are generally flower buds,
and develop in the autumn into oblong erect subsessile
hairy catkins about half an inch long, surrounded at base
by the bud scales, which pass into the very acute scales of
the inflorescence. The trees are dioecious, and while the
catkin character of the flower shoots is very evident on
the staminate trees, it is much less noticeable on pistillate
trees, the catkins of which are not above half the thickness
of the others, and with correspondingly narrower scales.
The lateral leaf buds are half ovoid, small, appressed to the
stem, and protected by a few scales similar to those of the
terminal bud. So far as has been observed, no super-
numerary buds, either collateral or superposed, occur.

The leaves are lanceolate to elliptical lanceolate, acute at
each end, entire, very narrowly revolute, 3 to 4 inches long,
on half-round petioles about 1 in. long, and densely ap-
pressed villous, with a few interspersed clavate glandular
hairs, especially on the petiole, and evident only on close in-
spection. With age they may become as much as 3 X 7 in.
with petioles 2 to 3 in. long, and are then glabrate and some-
what glossy above except along the midrib and principal
veins, thick and of coriaceous texture, and finally very
rugose on the paler under surface from the prominence of
even the finer veins. Stipules have not been seen.*

The flowers expand before the leaves, early in March,
when Acer rubrum Drummondii is in bloom. The stam-

*In Leitneria Floridana stipules are said to occur by Baillon, Hist.
des Plantes, vi. 241; Van Tieghem & Lecomte, Bull. Soc. Bot. de France,
xxxiii. 184; and Heim, Recherches sur les Diptérocarpacées, — Thesis,
Paris, 1892,—-176. They are not found by Chapman, FI. So. U. S. 428;
Oliver, Hooker’s Icones Plant. ser. 3, i. pl. 1044; and Bentham & Hooker,
Genera Plantarun, iii. 397.

3

68 MISSOURI BOTANICAL GARDEN.

inate catkins then become from one to two inches long,
generally curved outwards, and their scales spread just
enough to expose the stamens and allow the very abundant
and powdery yellow pollen to escape. The soft paren-
chyma of the axis of inflorescence becomes torn in various
directions as the catkins elongate, so that when they have
reached their full development it is loosely fissured through-
out. The fibrovascular bundles at the same time are poorly
developed and almost unlignified, so that it is almost impos-
sible to dissect a catkin without tearing it in pieces. The
same loose texture exists in the basal part of the catkin
scales, where they have increased in length during anthesis,
and as the outer part is considerably longer than the inner, it
assumes a series of characteristic transverse wrinkles below.
This separation of the tissues in both axis and bract, gives
rise to the curious appearance in longitudinal section which
is shown in plate 32, fig. 4-5, for the cleft in each bract is
decurrent down the axis to the point where the firmer fibro-
vascular bundles emerge for the next flower. The staminate
flowers, so far as I have examined them, are glabrous and
quite destitute of a perianth or involucre of any description,
and consist simply of a whorl of about ten short filaments a
little dilated at base and surmounted by slightly versatile but
nearly erect extrorse two-celled anthers dehiscing longitu-
dinally. The pollen grains are nearly globose, smooth,
slightly 3 to 4-grooved with underlying thickening of the
intine, and fall from the dehiscent anther very readily, and
there is no doubt that the species is wind pollinated.*

The pistillate catkins possess the same loose lacunose
structure as the staminate, though the axis is far less torn.
When fully developed they are rarely over half an inch long,
and it requires some little care to detect their presence on

* The structure of the staminate flowers, aside from the lacunose
character of the axis and bract, and the extrorse facing of the anthers, is
well shown by Baillon, J. c. 240, f. 214; Oliver, J. c. f. 1 to 3; and Heim,
i.c. pl. 10, f. 1 to 6. Baillon, J. c. 239, mentions bractlets as being some-
times present.

4

eR a RE Ae te Bs SEP ae nS,

LEITNERIA FLORIDANA. 69

the trees, whereas the staminate catkins are very evident
from a considerable distance. Unlike the staminate flow-
ers, the pistillate, which are limited to the upper axils, are
very short-stalked or with a rudimentary disk, and possess
a rudimentary involucre or perianth of a few small, glandu-
lar-fringed scales, the largest two of which stand nearly
laterally while the remainder are dispersed along the side
next the axis of the catkin.* Only one carpel is present.
The ovary is shortly ovoid, finely pubescent, one-celled, and
contains a single ascending parietal ovule with the micropyle
directed upwards. The green or slightly reddish style is
attached a little at one side and in anthesis curves outwards
and becomes grooved on the stigmatic side, or somewhat
flattened, with the stigmatic surface undulated, possessing
the general characters of wind-pollinated stigmas. The
placenta and stigmatic groove are turned away from the
axis and face the bract, a very unusual position for the
suture in a monocarpellary flower, and one which appears
to indicate that the flower is in reality reduced from a
former state in which there were two carpels radially ar-
ranged with reference to the bract, or perhaps a larger num-
ber; and this inference that the simple flower of Letinera
has been formed by the reduction of an originally more com-
plex flower is further supported by the presence of a rudi-
mentary perianth about the pistil, and by the reported
occurrence of abortive pistils near the end of the staminate
catkins in some instances,f and of one or more stamens
within the scales of occasional pistillate flowers.

The fruit is an erect compressed dry drupe measuring

* Eichler (Blithendiagramme, ii. 42) calls attention-to the large size
of the lateral scales, which, from analogy with Myrica, he regards as
bractlets, considering the others as a perigone, and Van Tighem &
Lecomte (1. c. 184) recognize a calyx as present in the pistillate flower.
Heim (Ass. Frang. J. c. 231) on the other hand speaks of the absence of
both calyx and corolla, in agreement with Baillon, who speaks of the
whorl of scales as a false calyx (1. c. 240, note).

+ Oliver, Hooker’s Icones, J. c. p. 34.

t Baillon, Hist. vi. 240, note.

70 MISSOURI BOTANICAL GARDEN.

about 6X8X22 mm. in the Floridan specimens, though in
the few Missouri specimens that I have seen it was of
scarcely more than half this size. Its surface is coarsely
rugose-reticulated over the firm fibrovascular bundles of
the pericarp. Near the top it is marked by an oblique scar
left by the caducous style, and it contains a single large
seed with a straight embryo and a rather thin layer of
albumen. *

So far as can be made out at present, the Missouri
Leitneria differs from that of Florida only in its larger
size,— the southern plant being described as a shrub 2 to
6 feet high,—in its somewhat larger more coriaceous
leaves rarely more than acute at apex, while those of the
Floridan plant are usually somewhat acuminate, and in its
apparently smaller fruit. Neither of these characters,
however, need of necessity be of specific value, nor repre-
sent more than individual variation due to habitat, climate,
or age.

In an account accompanying his illustrations of. Leitneria
Floridana, Professor Oliver | mentions ‘‘ specimens of per-
haps a second species of Ledtneria from Texas, collected by
Drummond; ’’ and Bentham and Hooker { and Hemsley §
admit this probable second species. In reply to an inquiry,
Professor Oliver writes me that the material of this form
consists of imperfect specimens without original label but
labeled by Sir William Hooker as from ‘Rio Brazos,
Texas, Drummond; ’’ and he adds that so far as these
specimens go, he does not now see any reason why they
should not belong to LZ. Floridana, though the catkins are

* Except for the extrorse placental suture, and the lacunose bracts
and axis, the structure of the pistillate flowers is well shown by Baillon,
f. 215; Heim, pl. 10, f. 7 to 10; and Oliver, f. 6 to 8 — where the position
of the stigmatic groove is correctly shown, but the placenta inverted.
The fruit and seed characters are also shown by Baillon, f. 216; Heim,
pl. 10, f. 11 to 22; and Oliver, f. 9.

¢ Hooker’s Icones Plantarum, 8 ser. i. p. 34.

¢ Genera Plant. iii. 397.

§ Biol. Centr. Amer., Bot. iii. 162, iv. 193.

6

LEITNERIA FLORIDANA. i pi

perhaps slenderer. In a letter accompanying this note,
Mr. Hemsley states that he also suspects that all of the
Leitneria material is of one species.

Dr. Chapman informs me that in Florida the original
stations of the species on the coast have long since been
washed away, but he afterwards found it inland, where,
nevertheless, its distribution is not known to be more than
very restricted. Whether the Texan material referred to
above was actually gathered along the Brazos, or possibly
further east, where Drummond also collected extensively,
the.occurrence of Leitneria in Missouri is, therefore, very
remarkable. But the investigations of Mr. Bush have
shown that this part of our State is a meeting-point for the
floras of the Middle States, the eastern Gulf region and
Texas.*

The reason for the extension of the Gulf Flora noted,
appears to lie in the deep swampy character of much of
the land along the general course of the Mississippi River
in southern Missouri and northern Arkansas as well as
further toward the Gulf. This entire region is noted for
the present fluctuations in its water level, due to artificial
elevation of the immediate bounds of the great river and
its tributaries, some of which nevertheless are destroyed
almost every year during freshets; but presumable natural
changes, and the large measure of success which has been
reached in confining the streams to their channels and
inducing a more rapid flow near the mouth of the Missis-
sippi, have resulted in lowering the maximum average level
of the water in case of inundation, and a far more per-
fect drainage of the lowlands than formerly, during the
season of low water. This seems to be indicated quite
clearly by a study of the cypress trees of the region. I
believe it is generally admitted that the level to which the
root knees of the cypress rise in wet cypress swamps cor-
responds closely with the normal flood water level. This is

* Fifth Garden Report, 140.

TT? MISSOURI BOTANICAL GARDEN.

the case in the deep sloughs of the Varner and St. Francis
rivers near Kennett, though at low water the knees emerge
about two feet. But here and there dead knees, quite
unconnected with standing trunks, or clearly belonging to
the shells of old trees none of which are now living, rise
some three feet beyond the level marked by the knees of
existing trees. These vestiges of an earlier forest growth
appear to indicate that when they were in their prime the
water level stood several feet above the present flood
level, and so far as can be estimated from the trees this
was somewhere from two hundred to five hundred years
ago.

Professor R. Ellsworth Cail, who has made a geological
study of the region in which these plants now appear to
find their northernmost home, writes me that the Mis-
sissippi river formerly without question flowed far to the
west of its course to-day, and in comparatively recent
geologic times cut away the opposing ridge below Cape
Girardeau, and excavated its present channel and the low-
lands around. Meantime it has swung across the valley
several times, but its westward movement has been succegs-
fully resisted by the remains of the great Tertiary plateau
or northward deposit of the ancient Gulf, which extends
from Florida to Mexico and northwards to this region, of
which Crowley’s Ridge is believed to be the last remaining
vestige in the middle valley. He mentions as interesting
in connection with these changes in the Mississippi and its
immediate valley, the fact that the Ohio long preserved its
integrity to a point not far from the present site of
Helena. He also states that there can be no question that
these changes at times have been retarded by depression of
the whole area, and at times hastened by elevation; that
there have been comparatively recent times when all the
region about northeast Arkansas and southeast Missouri
was a veritable marshy waste; and that this condition has
been several times repeated, with a synchronous ameliora-

8

LEITNERIA FLORIDANA. 73

tion of climate favoring the northward extension of a
subtropical flora over the region.*

It seems probable, therefore, that Leitneria and the
species of the Floridan flora which accompany it in the
Missouri swamps, represent the remnants of a warm tem-
perate swamp flora which at one time extended continuously
in the low lands, around the coast and up the river, from
Florida to the upper limits of the present deep swamps;
and that they are now likely to be found in such situations
at any point between the extremes, where the original
conditions have remained little or not at all changed by the
general drainage which has been progressing at least for
the last few centuries. One of the most promising fields
for botanical exploration in the eastern United States, and
one of the least known, is the swamp region of the lower
Mississippi Valley and the Gulf Coast; and I have little
doubt that a fuller knowledge of the flora of this region
will not only confirm the explanation here offered of the
occurence of Leitneria in Missouri, but extend its known
range from this point to its original locality in Florida.
Unless there is an error in the label of Drummond’s speci-
men, it should also be found in similar situations across
Louisiana and around the Gulf in Texas.

Mr. Bush’s attention was first called to the occurrence
of Leitneria in Missouri by the frequent mention in the
swamps of avery light ‘‘ cork wood,’’ greatly surpassing
even young tupelo (Vyssa) in buoyancy, and much used
by fishermen for floats on their nets; and the trunks which
he placed in my hands were of such surprising lightness
that I requested my friend Professor Nipher to determine
the specific gravity of the decorticated wood. An exam-

* The results of Professor Call’s work, here briefly outlined from his
letter, are to be found in the Proceedings of the Iowa Academy of
Science, 1887-9, 52, 85, and more fully, as volume ii. for 1889 of the
Reports on the Arkansas Geological Survey. See, further, a paper by him
in Amer. Journ. Sci. and Arts, 1891, xlii. 394, on the fossil woods of this
region, also considered in his larger report.

9

74 MISSOURI BOTANICAL GARDEN.
*

ination by his assistant, Mr. Brogan, shows it to possess a
density of 0.207, water being taken as unity.* As is often
the case with the roots of light trees, those of this species
are appreciably lighter than the stem, and a similar deter-
mination made by Mr. Timmerman at the request of Pro-
fessor Nipher, gives for its root wood a density of 0.151,
though as a cylinder of 3.79 cc. only could be used, it is
probable that this determination is less accurate than that
for the trunk.

In his treatise on the forest trees of the United States, f
Professor Sargent gives a tabulation of the specific gravity of
the woods of all the North American trees recognized at that
time, as determined at the Watertown Arsenal. From his
tables it appears that the lightest known native wood (that
of Ficus aurea of Florida), has a density of 0.2616, and
the heaviest (also a Floridan species, Condalia ferrea), has
a specific gravity of 1.302, while the density of the great
majority of species lies between 0.400 and 0.800.t

From a set of tables based on the experiments of
Captain Fowke,§ on the woods exhibited at the Paris
Exposition of 1855, and the London Exposition of 1867, it
appears that an East Indian wood, ‘‘ Dedoaf Tha,’ has a
specific gravity of 0.260, very nearly that of Ficus aurea

* Professor Nipher informs me that in the determination, a cylinder
was used, turned as accurately as possible, which was measured at a
sufficient number of points to give average values, and the density was
obtained by calculation from the dry weight of the cylinder, correction
being made for absorption of water between the time of removal from
the drying bath and the completion of the weighing, it having been found
that the gain from the atmosphere was .007 gr. per minute, for the
cylinder used.

t Final Rept. Tenth Census, ix. 249,

{ As a few examples may be mentioned the common hickories, ranging
from 0.810 to 0.887, white oak, 0.747, tupelo, 0.519, the willows and pop-
lars, usually considered very light woods, and lying between 0.363 and
0.607, and white cedar, a favorite material for light boat construction,
with a density of 0.332.

§ Science and Art Department of the Committee of Council on Edu-
cation, London, printed by George E. Eyre and William Spottiswoode,
1867, p. 10.

10

LEITNERIA FLORIDANA. 15

as given in the census tables. Ochroma lagopus, one of a
number of so-called cork woods enumerated by Wiesner,*
is said to have a density of 0.250,f but I cannot place my
hand on any determinations lower than this. Professor
Nipher tells me that the specific gravity of common cork
(the bark of Quercus Suber) is given by Ganot as 0.240.
Although individual variability and the difficulties of deter-
mining the specific gravity of porous and absorbent bodies
like wood make it unwise to accept any of these figures as
invariable, it appears from them that the wood of Leitneria
is very markedly lighter than the bark of the cork oak,
which itself is considerably lighter than any other wood of
which a record can be found.

0.16-0.20
0.46-0.58
0.61-0.65
0.71-0.75
0.91-0.95
1,.26-1.30

q y

FREQUENCY OF TIMBER DENSITIES.

The appended curve, compiled from the census tables,
shows the relative frequency of occurrence of the different
degrees of density in our native timbers, each co-ordinate
representing a difference of .050, and each species falling
within this range being represented by a vertical distance of

* Rohstoffe, 578.
+ Goodale, Physiological Botany, 145.
11

—

76 MISSOURI BOTANICAL GARDEN.

1 mm. on the co-ordinate. Thus, one species has a specific
gravity between 0.16 and 0.20 inclusive; none occur be-
tween 0.21 and 0.25; 46 are found between 0.46 and 0.50;
and 50 lie between 0.71 and 0.75. The majority of species
(242) occur between the maximal points of 0.46 and 0.75,
and it is evident that the fall is much more abrupt from
these common densities to the lowest recorded density than
to the highest, so that any extension in this direction is
more remarkable than a corresponding one in the other.

As the density of wood freed from resin etc., air and
water, is about one-half greater than that of water, the
reason for the extreme lightness of the wood of Lettneria
is to be looked for in connection with its loose structure,
the softness of its tissues, which are easily compressible
under the thumb nail, and the absence, at least in the
largest specimens I have seen, of any heart wood, the tex-
ture being homogeneous throughout.* It was, therefore,
subjected to a rather careful microscopical examination,
with the following results.t

The pith is nearly round in cross section, although where
the bundles of the primary wood join it they project
slightly, giving it a minutely crenulate outline. It is con-
tinuous and of uniform texture, thus differing from the
pith of plants like Paulownia and Lonicera which are ex-
cavated, or Juglans, which is chambered,t and from that of
Liriodendron, etc., which, though solid, is traversed by
firmer transverse plates. The cells, as is usual in pith, are
approximately isodiametric and polygonal, as seen in cross
section, a little smaller near the periphery ; and in longi-

* On this general subject see Nérdlinger, Gewerblichen Eigenschaften
der Holzer, Stuttgart, 1890, 17; and Techn. Eigenschaften der Hélzer,
Stuttgart, 1860, 115.

¢ The substance of this paper was presented orally before the
Academy of Science of St. Louis on May first, 1893, and illustrated by
specimens of the plant and photomicrographs, some of which are here
reproduced in half tone or are made the basis of line engravings, show-
ing the structure of the wood and bark.

t The ‘‘ discoid pith ’’ of Morren, in Ann. of Nat, Hist. 1839, iv. 73.

12

ae ee Lame,

LEITNERIA FLORIDANA. FT

tudinal section they often appear as rectangles, with their
greatest diameter transverse. Their walls, which are very
thin, show, nevertheless, a decided secondary thickening,
and are marked by simple pits, as is usual in pith of this
description. Toward the wood, and especially where the
pith rays pass outward to the branches, occasional cells in
more or less marked vertical rows occur, with large stellate
crystals.

At the margin of the pith a number of layers of cells of
reduced diameter but several times as long as the pith cells,
constitute a pith sheath, and gradually merge into what
are virtually wood parenchyma cells. They have the same
simple pits as the pith cells, but are arranged in vertical
rows in relation with the elements of the xylem, and vary
in length from twice to four or five times their diameter.
In the mass of parenchyma so formed at the margin of the
pith, are found intercellular secretion reservoirs, each of
which is surrounded by a layer of oblong secreting cells
with walls not appreciably thinner than those of the paren-
chyma immediately about them. From one to two dozen
such passages are to be seen in a cross section of the stem.*
Though as a rule they occur singly, it is not uncommon for
two of equal size to stand close together, separated by only
a few layers of cells, or for small ones to stand on one or
both sides of a larger one. The secreting cells are usually

* These were formerly attributed by Van Tieghem (Ann. des Sci. Nat.
7 ser, i. 64, and Van Tieghem and Lecomte, Bull. Soc. Bot. de France,
1886, xxxiii. 182) to the apexes of the primary xylem wedges, but, ac-
cepting as the inner limit of the wood, the position of the innermost
vessels, Professor Van Tieghem now recognizes these secretion pas-
sages as pertaining to the pith (Morot’s Journal de Botanique, v. 384,
385), thus coming into agreement with Miiller (Engler’s Bot. Jahr-
biicher, ii. 449), Burck (Amn. Jard. Bot. Buitenzorg, vi. 151), and Heim
(Ass. Frang. p. l’Avanc. des Sciences, 1891, i. 231; Rech. s. les Diptéro-
carpacées, Thesis, 1892, 175), who have attributed the similarly situated
passages of Dipterocarpeae to the pith. But Reinsch, Engler’s Jahrb. xi.
374, speaking of the Balsamifluae, again calls attention to their close con-
nection with the xylem, and, indeed, it appears extremely difficult so to
define the hadrom bundle as unequivocally \to separate it from the

adjacent pith.
13

78 MISSOURI BOTANICAL GARDEN.

quite convex on the free side, and sometimes protrude into
the passage as papillae or in balloon-like outgrowths. Their
secretion is a yellowish resin, which fills the passages in
sections cut dry and examined in water, but rapidly dis-
solves when this is replaced by alcohol drawn in at one side
of the cover glass, leaving a residue of fine emulsion drops
similar to those formed when an imperfectly anhydrated
preparation is mounted in Canada balsam. In sections ex-
amined in water after being cut dry, the contents of the
secretion passages are traversed by fractures, sometimes
regularly curved, and their high refractive power gives rise
to the appearance of a limiting membrane about the indi-
vidual masses, suggestive of a raised cuticule of the secret-
ing cells, beneath which, according to Tschirch,* the
formation of the resins of schizogene passages occurs. In
observing the solution of the resin on the addition of alco-
hol, however, the apparent membranes are seen to com-
pletely disappear at once, and I have not been able to detect
either cuticular blisters or vestiges of the bases of such
blisters in these sections after the solution of the resin, nor
in any of the numerous sections of alcoholic material which
I have examined. ft

Just outwards from the situation of this ring of secretion
passages, each wedge of the hadrom or xylem begins with

* Berichte Deutsch. Bot. Ges. xi. 201, and Pringsheim’s Jahrb. xxv.
375.

+ Some difference of opinion exists as to the nature of the secretion
within these passages. Van Tieghem and Lecomte (1. c. 182) speak of it
as a resin, while Heim (Thesis, 176) calls it a balsam. The entire
group of terpenes, ethereal oils, resins, and balsams, is a difficult one
chemically, and I do not venture to pronounce on the one now in ques-
tion further than to say that it is insoluble in water, soluble in cold
alcohol except for the emulsion residue referred to above, and that Mr.
Bay, who at my request tested it with the Unverdorben-Franchimont
reaction to acetate of copper (Poulsen, Mikrochemie, 73; Zimmermann,
Bot. Mikrotechnik, 89), found that it assumes with this reagent the green
color characteristic of resins and terpenes. It may be noted that the
same reagent gives an abundant brown precipitate throughout the bark,
indicative of the presence of tannin. On the resins see further Tschirch,
Pringsheim’s Jahrb. xxv. 370.

14

Sod (Sg. Sada ha ra
; a, ea cs 5

LEITNERIA FLORIDANA. VAS)

a few tracheides and spiral vessels about 20 » in diameter,
surrounded by wood parenchyma. Aside from this, except
where the bundles pass out to the leaves, no true spirals
occur in the stem. The remainder of the xylem consists
of pitted tracheides and ducts, wood parenchyma, and,
chiefly, libriform cells. Except for the spiral vessels men-
tioned above, the vascular elements of the wood are all
furnished with bordered pits, a scalariform reticulation
marking their contact with the medullary rays.

The pitted vessels are distributed through the stem in a
characteristic way, as is usual in woody plants. Each year’s
growth in the secondary wood begins with a greatly inter-
rupted row of vessels large for the plant, and measuring
from 50 to 95 » in diameter. In addition to these vernal
ducts, each year’s zone of wood contains a number of
groups of similar but smaller vessels, which are arranged
either approximately parallel to the circumference or with
an oblique direction outwards, often strongly accentuated
in the older wood where it may become quite radial, and,
as acommon thing, grading in size from the middle to each
extremity or from one end to the other, most diameter
measurements lying between 25 and 35». These groups
consist of the elements usual in such duct accumulations,
namely vessels, tracheides, and wood parenchyma. The
vessels show a complete disappearance of the cross septa
which originally divided the vertical rows of rather short
cells from which they arise, except for a narrow ring
around the margin, in this respect differing very markedly
from the similar vessels of Liriodendron, Liquidambar,
etc., where the perforations are formed in such a way as to
leave the original septum as a persistent scalariform plate
running obliquely across the mature duct.

The wood parenchyma, which is not abundant, and stands
in close connection with the ducts and annual rings, does not
differ particularly from the parenchyma already described
about the secretion passages in the pith sheath. The tra-
cheides as arule are similar to the libriform or common wood

15

———— ee

80 MISSOURI BOTANICAL GARDEN.

cells, presently to be described, except that their walls are
somewhat thicker,—sometimes much thicker,—and marked,
even on the tangential sides, with bordered pits very different
from the simple pits of the libriform cells and parenchyma.
Their walls not infrequently show a spiral striation which
in its most marked form consists in a fine acute spiral ridg-
ing of the inner surface of the cell, the ridges, however,
differing from those of the spiral vessels in being wedge
shaped with a broad base, whereas in the vessels they are
of a generally round section, and attached by a narrow
base, which is easily broken, so that they can be uncoiled
from within the lumen.

Except for these masses of vessels with their accompa-
niment, and a layer of cells presently to be described, at
the limits of each year’s growth, the wood consists of
libriform or ordinary wood cells, which differ from the
tracheides in possessing on the radial walls very small,
obliquely crossing, not evidently bordered pits, scattered or
irregularly grouped. They are fusiform or, more accurately,
obliquely truncated at the ends, which overlap in such a
way as to be very evident in tangential sections while much
less obvious in radial sections. Their length is usually
fifteen to twenty times their diameter, the measurements as
a rule lying between 15 X275 and 20X375. Now and then
these libriform cells are found with transverse partitions
running directly across, one or two to each cell, a condition
that has been observed frequently in other woods. While
striation is much less evident than in some of the tracheides,
it is sometimes to be seen.

Rings indicative of the annual (or more accurately
periodic) growth in thickness of the stem, though nearly
invisible to the naked eye, are evident on an examination
of the wood under a lens, being partly caused by the
occurrence of a broken row of vernal ducts, and in part by
a gradual reduction in the radial diameter of a few rows of
libriform cells formed toward the end of the growing sea-
son, whereas those first formed the next spring are of

16

LEITNERIA FLORIDANA. 81

ample size. The most striking feature of these annual
rings, however, consists in the production of a row of
parenchyma cells replacing the libriform as the first series
cut off by the cambium each spring at the beginning of the
year’s growth. These cells are about ten times as long as
broad, with horizontal septa, some of which are evident in
all cross and radial sections of the wood, and, like other
cells of the wood parenchyma, they show simple pits similar
to those of the medullary rays. Exceptionally this annual
layer of parenchyma is locally doubled by tangential
division.

Between the xylem wedges occur the usual medullary
rays. So far as I have seen, these rays, except where they
stand in connection with pith rays, are not more than two
cells in thickness, and it is extremely unusual to discover
more than a single row of cells in their cross or tan-
gential section. As seen in longitudinal section, they
consist of from one to about twenty vertical series of cells,
most commonly about ten. These cells usually measure 18
to 25 » in height, 7 to 10 » in width, and 40 to 75 » (as
a general thing) to above 100 yin radial length. Their
vertical septa frequently are somewhat oblique when
viewed in either transverse or radial section. Their walls
are thin, and simply pitted to correspond with abutting
cells.

All of the elements of the xylem show the customary
middle lamella, with a secondary thickening, which, how-
ever, is very slight, so that the walls of the libriform cells
are rarely over 1.5 » thick, while the medullary ray cells
are thinner, and the tracheides and the wood parenchyma
cells generally range respectively a little thicker and a little
thinner than the libriform, the vessels being, in fact, the
only thick walled cells of the wood, except for occasional
groups of exceptionally thickened tracheides.

From its gross anatomical characters, the wood of Leit-
neria would be compared with Hartig’s group of dicotyle-
donous woods having all of the ducts small, those of the

. 17

82 MISSOURI BOTANICAL GARDEN.

spring growth neither large nor numerous, and the medul-
lary rays invisible to the naked eye.* The occurrence of a
single broken row of vernal ducts somewhat larger than those
of the rest of the year’s growth, the oblique position of the
groups of the latter, and the absence of loose parenchyma-
tous bands, suggest a comparison with Aster argophyllus,
certain species of Ulmus and Celtis, and, particularly,
Daphne Mezereum.t Except for the absence of coarse
parenchyma bands, it also resembles somewhat the wood of
Ailanthus, Hippophae, and numerous Leguminosae, though
in these the vernal ducts are usually larger and more numer-
ous, and the secondary thickening of the medullary rays
and the libriform cells is far more marked.

The cortex of Leitneria, which, as has been stated
above, is rich in tannin, is rather thin, and consists at first
of fundamental parenchyma, which is collenchymatously
thickened, with large often transversely elliptical pits, for
about eight layers of cells immediately below the epidermis,
and passes into a like number of thin walled cells by a tran-
sition through about three layers, while between this primary
cortex and the cambium an abundant secondary cortex is
developed, containing large fan-shaped abundantly crystal-
liferous dilatations of the principal medullary rays, between
which lie broad wedges of bast. Except for a few small
and scattered bundles of hard bast fibers in the pericycle,
at the inner border of the primary cortex close beneath the
collenchyma, none of the bast fibers become thick walled,
but they remain as long wide generally irregularly collapsed
tubes with oblique often clustered simple pits, and destitute
of protoplasm. Traversing these bast wedges are a few
secondary rays, while tangentially they are parted by plates
of thin walled parenchyma cells from two to about five
times as long as broad, with horizontal septa, a few rows
of which contain crystals; and among these cells, rich in

* Hartig, Timbers and How to Know Then, 8.
+ From an examination of Nordlinger’s set of 1100 cross-sections of
woods.— Cf. p. 78 of text accompanying century xi.

18

LEITNERIA FLORIDANA. 83

protoplasm, sieve tubes should be found.* Thetwo systems
thus map out the bundles, in cross-section, into a series of
quadrangles.

Throughout, like the wood, the cortex is destitute of
secretion passages. The cork, which is formed immedi-
ately next the epidermis ¢ during the first summer, cuts off
a bark which does not become very thick, and is inter-
ruptedly stratified by the intercalation of masses of con-
densed cells between layers of more open cells.{ Grit
cells are entirely absent, and I have not satisfied myself that
the primary cortical parenchyma is added to below the
cork by the formation of phelloderm from the inner limits
of the latter.

In sections just below the nodes, fibro-vascular bundles
may be found running obliquely upwards through the
cortex, from the xylem to the leaf scars. The tracheary
elements of these bundles are spirally marked as in the
primary xylem next the pith, and they are unaccompanied
by secretion passages. So far as I have observed, their
transit through the cortex is effected in a vertical distance
little greater than the thickness of the latter, so that they

* Van Tieghem and Lecomte (J. c. 181) speak of these tangential bands
as consisting of ‘tubes criblés,’’ and it is clearly in them that the sieve
tissue should be found; but notwithstanding repeated examination of
sections from fresh, dry and alcoholic material, which had been subjected
to treatment which renders the sieve tubes of Tilia, Magnolia, Ulmus and
other trees very evident, I have quite failed to demonstrate sieve plates
in the cortex of this species.

+ So far as can be judged from specimens still retaining the epidermis,
but with fifteen or twenty layers of cork cells, the first subepidermal
layer of cortical cells becomes active as phellogen; but as this is formed
in the early summer of the first year, while I have been able to study only
very young shoots and those which had ended the season’s growth, I
have been unable to get a preparation showing the first segmentation,
which would afford conclusive information on this point.

¢t This is to be compared with the annual cork layers described for
Balsamifluae by Reinsch, Engler’s Bot. Jahrb. xi. 367.

§ The occurrence of true cortical bundles in various groups of plants
is discussed by DeBary, Vergl. Anat. 266; Mueller, Engler’s Jahrb. ii.
449; Gilg, Ber. Deutsch. Bot. Ges. xi. 21; Burck, Ann. Jard, Bot. Buiten-
zorg, vi. 156; Heim, Thesis, 18, etc.

19

84 MISSOURI BOTANICAL GARDEN.

really belong to the pulvinus of the leaf, but this does not
prevent them from appearing as distinct cortical bundles in
favorably located sections (pl. 39 and 44f.1-3). Three of
these foliar bundles, corresponding to the three bundle
traces evident on the scars marking the former position of
fallen leaves, pass into each petiole, where they soon unite
to form a closed crescent-shaped ring of bundles. No
isolated bundles are contained within this petiolar ring, the
parenchyma within which includes a series of about twenty
secretion passages similar in structure and contents to those
of the stem, with which, however, they have no direct com-
munication. From the petiole, a group of resin passages
runs through the midrib of the leaf, a single one passing out
into each of the finer veins. Stellate crystals are of rather
frequent occurrence through the petiole and midrib, both
of which contain collenchyma.

The upper epidermis of the leaf blade is smooth walled
except for a few striated cells about the bases of some of
the hairs, and consists of a layer of inconsiderably thick-
ened cells, beneath which lies a layer of quadrate cells each
of which contains a large stellate crystal. A layer of
similar cells is also found between the veins and the lower
epidermis, the cells of which are somewhat smaller and
prominently wrinkled on the outer wall, so as to appear
almost muricate in cross section. The stomata are not
sunken below the general level of the epidermis. Tannin
appears to be abundant in the epidermal layers.

Pubescence consists of two kinds of hairs:— abundant,
slender pointed thick walled hairs, usually with several
transverse septa, especially toward the often bulbously
widened base, and mostly isolated, but occasionally binate ;
and.less numerous clavate hairs, septate both longitudinally
and transversely, their small cells with yellow contents.
These compound hairs are chiefly seen on the young stem,
the sides of the petiole, and the upper surface of the mid-
rib of the leaf. The epidermal cells about the bases of the
hairs are usually divided by a septum parallel to the leaf

20

oe a, ae ee? A oe OP ae POR NO BER AOS

LEITNERIA FLORIDANA. 85

surface. The mesophyll is composed of two or, mostly,
three layers of compactly placed palisade cells only a little
longer than broad, occupying the upper half of the leaf,
and a spongy parenchyma with ample intercellular spaces
below. No spicular cells have been observed.

The structure of the roots is interesting to this extent,
that (at least in lateral roots, which, alone, have been
examined) they are entirely destitute of secretion passages,
which are thus seen to be confined to the pith sheath of the
stem, the intra-fascicular parenchyma of the petiole, and
the parenchyma of the veins of foliar organs, including the
carpels. The elements of the root are essentially similar
to those of the stem, and secondary growth in thickness is
effected in the manner usual in the roots of Dicotyledons.

While I have found comparatively little starch in the
stem of specimens gathered either in November or in the
spring, the medullary rays and cortical parenchyma of the
root contain an abundance of roundish grains, often binary,
and of extremely variable size.

Dr. Chapman, who described the genus Lettneria and its
single species, placed it among the Myricaceae because of its
simple flowers arranged in spike-like catkins;* and in this
he was followed by DeCandolle.t Subsequently Baillon
examined it, and placed it with doubt in his group of Cas-
taneae, beside Myrica, making it, however, the representa-
tive of a series which he named Leitnerieae,t and in which
he also placed with question a Madagascar genus, Didymeles.
This series was raised by Bentham and Hooker § to ordinal
rank, under the name given by Baillon, and placed between
the Platanaceae and Juglandaceae, with an indication of
Urticaceous affinities, Didymeles being excluded. The
order is maintained under the name Leitneriaceae by
Engler, || who places it between Myricaceae and Salicaceae.
~ * Flora So. U. 8. 427.

+ Prodromus, xvi. (2), 154.

¢ Hist. des Plantes, vi. 239, 258; Tison, in Baillon, Dict. de Bot. iii. 215.

§ Genera Plant. iii. 397.

| Engler and Prantl, Pflanzenfamilien, iii. (1), 28.
21

86 MISSOURI BOTANICAL GARDEN.

In 1886, Van Tieghem and Lecomte,* after a_histo-
logical study of the stem and leaf, decided, because of the
stratification of the bast and the occurrence of secretion
passages in the pith sheath and the leaf, that Lettneria may
be placed in or at least joined to the polypetalous order
Dipterocarpeae, its macroscopic characters being also not
unlike those of that family except for its diclinous reduced
flowers. More recently Heim { has reviewed its histological
characters, and reached the conclusion that its affinities are
rather with the group Balsamifiuae or Liquidambareae of
Hamamelideae (likewise a polypetalous group, with which
Van Tieghem and Lecomte had also recognized that it
might be compared in certain respects), near which for
both anatomical and organographic reasons he would place
the Leitnerieae, while allowing it to retain its autonomy.

Many botanists look on the Apetalae as only a provisional
aggregate of orders with reduced inflorescence, which must
be disrupted before a truly natural system of classification
is reached; and efforts have been made from time to time
to distribute all of these orders among the Polypetalae and
Gamopetalae, just as single apelatous genera and species
are universally placed in orders otherwise characterized by
the possession of more complete flowers. This problem is
one that may be expected to receive aid from the many
comparative histological studies now being made, especially
by French and German botanists, though no purely organ-
ographic system dispensing with the Apetalae has yet met
with more than limited approval.

Against the approximation of the Leitnerieae to the
Balsamifluae, may be urged the absence of secretion pas-
sages from the liber, and the non-scalariform duct perfora-
tions, though the floral structure suggests a possible further
simplification of the type of this group. On the other
hand, the absence of secretion passages from the root, the

* Bull. Soc. Bot. de France, xxxiii. 184.
+ C. Rend. Assoc. Frang. pour 1’Av. des Sci., 1891, i. 281; Recherches
sur les Diptérocarpacées, Thesis, Paris, 1892, 175.

22

RS ee a ara ET Lee tay se

LEITNERIA FLORIDANA. 87

narrow medullary rays, scanty wood parenchyma and hard
bast, peculiar duct pattern, simple petiolar bundle arrange-
ment, and the venation of the leaves, count against too
close a union of the Leitnerieae with the Dipterocarpeae,—
a conclusion which is further strengthened by the fact that
no existing or fossil representatives of this order are known
from the New World. For the present, therefore, the
order Leitnerieae will probably be maintained either in the
position it now occupies next the Platanaceae, or, in case of
the dismemberment of the Apetalae, near the Dipterocar-
peae or Balsamifluae among the Polypetalae ; and on this
point no one class of histological characters appears to be
conclusive. *

The peculiar lightness and softness of the Missouri cork
wood, combined with its slight porosity, suggest that it
should find application in the arts if, as appears to be the .
case, it can be procured in suitable quantities for econom-
ical working, and while its small size bars it from very
extended use, it is possible that it may prove a useful sub-
stitute for cork in the manufacture of bottle stoppers for
chloroform and other gummy substances, which cause cork
to tear badly after a little use.

* Attention should be called here to the close affinity which Agardh,
Brongniart and Clarke have thought they saw between the Platanaceae and
Balsamiftuae.— On this point see Baillon, Adansonia, x. 134; Eichler,
Bliithendiagramme, ii. 66, and, on anatomical grounds, Gris, Ann. Sci.
Nat., ser. 5, xiv. 40, and Mémoire sur la Moelle, 267, and Reinsch, Engler’s
Jahrb. xi. 369. But Bentham and Hooker (Genera, iii. 396) do not at
all agree with this conclusion.

23

88 MISSOURI BOTANICAL GARDEN.

EXPLANATION ON PLATES ILLUSTRATING LEITNERIA
FLORIDANA.

The habit illustrations were drawn by Miss Grace E.
Johnson, from living plants or herbarium material. The
details are reproduced directly or redrawn from photo-
micrographs or drawings by the author.

Plate 30,— 1, Leafing twig, in spring; 2, autumnal leaf; 3, staminate
catkin; 4, opening pistillate catkin; 5, branch with partly grown fruit.
All natural size.

Plate 31. —1, Pistillate, and 2, staminate shoots, in winter, natural
size; 3, bud and leaf scar details, 3; 4, stem hair, X150; 5, excep-
tional, substellate arrangement of hairs, 200; 6, base of hair, 250;
7, glandular hair, 180; 8, nature print of young leaf, natural size; 9,
lower surface of old leaf, x2.

Plate 32. — 1, Staminate flower, with portion of base of bract, and 2,
dehiscent stamen, <4; 3, ventral, dorsal, and lateral view of indehis-
cent stamen, <9; 4, enlarged longitudinal section of a portion of a
staminate catkin, showing fissuring of bracts; 5, part of a similar sec-
tion, showing extrorse stamens and fissures of bracts and axis of in-
florescence, 18; 6, pollen grain from one end, and in optical section,
400; 7, cross section of pistillate catkin, 18, showing fissured axis
ata, adnate base of bract with fissure at b, receptacle or disk with
perianth, glands, and subtending bract atc, ovary with ovule and sub-
tending bract at d, lower part of stigma with bract at e, and upper
part of stigma with bract at f, — the sections belonging to successively
lower flowers, in the order of lettering, — from one of a series of cel-
loidin sections, retaining the several flowers and their parts in their
natural position.

Plate 33. — Details of pistillate flower and fruit: 1, fully expanded
pistillate catkin, natural size; 2, young flower with bract and scales from
the inner side, 5; 8, longitudinal section of young flower, showing
disk and scales, 18; 4, scales or segments of perianth, 18; 5, longi-
tudinal section of portion of catkin, 18, showing lacunose axis and
bracts, and pistil at point of separation from disk; 6-7, ovule entire and
in section, 18, — the bract at the left asin fig. 5; 8, fertilized ovary
after fall of style, <5; 9-10, partly grown fruits, with longitudinal and
cross sections, <2; 11, dried mature fruit, <2.

Plate 34. — Cross sections of wood of stem, showing duct patterns,
<60.

Plate 35. — Similar sections, for the annual rings, the upper 60, the
lower 125.

Plate 36. — Tangential sections of stem wood, the upper >60, the
lower, showing septate libriform, 125.

24

ae eas ARE

LEITNERIA FLORIDANA. 89

Plate 87. — Tangential section of stem wood, above, showing dilated
ray going to branch, 125; and, below, partial radial section of stem,
60, the xylem at the left, the phloem at the right,—showing complete
duct perforations, stratification of liber, and serial arrangement of crys-
tal cells in the latter.

Plate 38.— Cross section through wood and pith of stem, showing loca-
tion of secretion passages, X60; and radial section of pith, showing the
greater transverse diameter of its cells, x 125.

Plate 39. — Cross section above, of outer part of wood and inner part
of cortex, showing vernal row of ducts, quadrate liber, and, in the
primary cortex, the small ducts of a foliar bundle; below, for compari-
son, a corresponding section of Tilia Americana, with quadrate bast
wedges,— both 60.

Plate 40.— Cross section of primary cortex of very young stem, show-
ing epidermis and collenchyma; below, cross section of liber wedge,
showing thin walled bast fibers with secondary rays and transverse paren-
chyma bands (the section torn through the cambium), — both 125.

Plate 41. — Cross section of central portion of root, showing pith and
xylem, — the large openings are ducts; below, longitudinal section of
secretion passage in petiole, showing elongated form of secreting cells,—
both 60.

Plate 42. — 1, Enlarged radial section of stem xylem, showing medul-
lary ray below, libriform cells at the right, striated transition to tra-
cheides adjoining these, and striated tracheides with bordered pits at the
upper left; 2, radial section, showing reticulated marking of medullary
ray cells in contact with duct, 400; 3, enlarged longitudinal section of
duct, showing bordered pits and remnant of original transverse septum ;
4, much enlarged cross section of wood, showing secondary thickening of
libriform and medullary ray cells; 5, diagram of radial section of xylem
crossed by medullary ray, X200; 6, cross section, and 7, radial section
of xylem, in diagram, 200, showing parenchyma layer at annual ring,—
the vernal wood at the right, the autumnal growth atthe left; 8, diagram
of tangential section of wood, showing libriform cells and medullary
rays, 200.

Plate 43.— 1, Radial section of xylem, showing libriform at the right,
followed by tracheides, vessels, and wood parenchyma, the vessels sub-
scalariform where adjoining a medullary ray, X200; 2, cross section of
secretion passage in pith sheath, 320, pith at the left, xylem at the
right; 3, radial section through the left hand portion of same, 200, be-
ginning with pith at the left, and ending with the secreting cells at the
right; 4, cross section of outer part of pith (above) with stellate crys-
tals, pith sheath, and innermost part of a xylem wedge, showing two
spiral ducts, 200; 5, radial section of outer cortex of old stem, show-
ing stratified cork, 200; 6, similar enlarged section of stem toward end
of first season, showing phellogen layer and subjacent collenchyma; 7,
tangential section of subepidermal collenchyma of young stem, 200; 8,
cross section, and 9, radial section of secondary bast, showing open bast
fibers and interjacent bands of bast parenchyma, 200.

25

90 MISSOURI BOTANICAL GARDEN.

Plate 44.— 1-3, Diagram of successively descending cross sections of
young shoot at emergence of a branch, showing passage of the foliar
bundles through the pulvinus and their union to form the petiole bundle,
9; 4, collenchyma of petiole, 250; 5, radial section of subjacent paren-
chyma, 250; 6, cross section of secretion passage of petiole, 350;
7, cross section of midrib and adjoining lamina of leaf, 30, showing two
forms of pubescence; 8, cross section of very young shoot, showing col-
lenchyma, epidermis, and base of hair, 200; 9, cross section of young
leaf, and stoma, 200, showing crystal layer of upper epidermis, palisade
and spongy parenchyma, and striated lower epidermis.

By accident, the position of the ovule in the upper figure of plate 33,
f. 8 is shown inverted, the true position being that indicated in the cor-
rected figure here given.

26

REPT. MO. BOT. GARD., VOL. 6. PLATE 30.

i Pie Cm

LEITNERIA FLORIDANA,

bh alae, td oat (a i aaa ee

REPT. Mo. BOT. GARD., VOL. 6. PLATE 31.

—aee

LEITNERIA FLORIDANA.

PLATE 32.

Me

A:

=
SAQES
WN
ISS
Ss
SSESg

4

2

REPT. MO. BOT. GARD., VOL. 6.

LEITNERIA FLORIDANA.

REPT. MO. BOT GARD., VOL. 6. PLATE 33.

LEITNERIA FLORIDANA.

PLATE 34,

REPT. MO Bor. GARD., VOL. 6.

gence ea
Seeekothe

A FLORIDANA

ERI

LEITN

REPT. MO. BOT. GARD., VOL. 6. PLATE 35.

LEITNERIA FLORIDANA.

PLATE 36,

REptT. MO. BOT. GARD., VOL. 6.

A.

LEITNERIA FLORIDAN

PLATE 37.

REPT. Mo. BOT. GARD., VOL. 6.

IDANA.

LOR

AcH

LEITNERI

REPT. MO. BOT. GARD., VOL. 6. PLATE

38.

LEITNERIA FLORIDANA.

REPT. MO. BOT. GARD., VOL. 6. PLATE 39,

é F sig

<
Phe pe

LEITNERIA AND TILIA,

REPT. MO. BOT. GARD., VOL. 6. PLATE 40.

ea sich ts ¥e
we Bed he Tepe
Se dwaknie:.:

LEITNERIA FLORIDANA.

PLATE 41.

REPT. MO. BOT. GARD., VOL. 6.

LEITNERIA FLORIDANA,

PLATE #2.

Rept. Mo. BOT. GARD., VOL. 6.

LEITNERIA FLORIDANA.

REPT. MO. BOT. GARI , VOL. 6. PLATE 43.

a

fC

LEITNERIA FLORIDANA.

PLATE 44.

REPT. MO. BOT. GARD., VOL 6.

LEITNERIA FLORIDANA.

PLATE 46.

Bot. GARD., VOL. 6.

REPT. Mo.

YUCCA ALOIFOLIA,

Bs»)
ae
*

ape Fi Se 8°, Ass Ra

STUDIES ON THE DISSEMINATION AND LEAF REFLEXION OF
YUCCA ALOIFOLIA AND OTHER SPECIES.

BY HERBERT J. WEBBER.

The Yuccas or Spanish bayonets, since the discovery of
their peculiar and unique methods of pollination, have
become objects of very general interest and study. It is,
however, to the long continued and painstaking researches
of Drs. Engelmann,* Riley ¢ and Trelease { that we are the
most indebted for the very complete information regarding
this phase of the ecology of the genus which we now
possess.

The methods of dissemination in the various species of
Yucca, while very interesting in many features, have as yet
been very little studied. The only account that has come
to the writer’s notice is that given by Dr. Trelease in
the closing paragraphs of his paper on the ** Further Stud-
ies of Yuccas and their Pollination.’’ § Here the adaptation
of the different forms of Yucca fruits to different
methods of seed dissemination is mentioned, and interest-
ing observations are given on the dissemination of Yucca
brevifolia and certain capsular species.

* Engelmann, Geo., “The Flower of Yucca and its Fertilization.”
Bull. Torr. Bot. Club, Vol. II. (1872), pp. 33-87.— “‘ Notes on the
Genus Yucca.”? Trans. St. Louis Acad. of Sci., Vol. IIT. (1873), pp. 17-54.

¢ Riley, C. V., ‘‘On a New Genus in the Lepidopterous Family
Tineidae, with remarks on the Fertilization of Yucca.”? Trans. St.
Louis Acad. Sci., Vol. III. (1873), pp. 55-69. — ‘“¢ The Yucca Moth and
Yucca Pollination.”? Mo. Botanical Garden, 8rd Annual Report (St.
Louis, 1892), pp. 99-158.

t Trelease, Wm., “‘ The Nectary of Yucca.’’ Bull. Torr. Bot. Club, Vol.
XIII. (Aug., 1886), p. 135. — “* Notes and Observations. I. Detail Ilus-
trations of Yucca.’? Mo. Botanical Garden, 3rd Annual Report (St.
Louis, 1892), pp. 161-168. — “ Further Studies of Yuccas and their Polli-
nation.”? Mo. Botanical Garden, 4th Annual Report (St. Louis, 1893), pp.
181-226.

§ Trelease, Wm., l.c., p. 223.

(91)

92 MISSOURI BOTANICAL GARDEN.

There are three types of Yucca fruits which illustrate
three distinct methods of dissemination. These types of
fruits were used by Dr. Engelmann to distinguish the three
groups of the true Yuccas.

Sarcoyucca: — Fruits pendent, fleshy, indehiscent;
seeds thick and wingless, with ruminated endo-
sperm.

Clistoyucca: — Fruits pendent, dry and coriaceous,
indehiscent; seeds thinner than in Sarcoyucca, wing-
less; endosperm entire.

Chaenoyucca :— Fruits erect, capsular with septicidal
dehiscence; seeds strongly compressed and thin
with a winged margin; endosperm entire.

The Sarcoyuccas, comprising nearly one-half of the
known species of Yucca, have sweet edible fruits and are
apparently well adapted for dissemination by fruit-eating
animals, especially birds. In this group the seeds are
usually protected by the inner part of the ovarian wall,
which in the development of the fruit becomes hard and
firm, suggesting in texture and functions the core of an
apple. The pulp is easily removed from the core, which is
more or less shaped to the seeds. This structure would
suggest that the pulp only is eaten, the core and seeds
being thrown away. In this case, the fruits, it would
seem, were intended to be carried away by the animals
eating them in order to remove the seeds from the parent
plant. Birds in picking away the tender pulp would
hardly reach the protected seeds but merely remove the
pulp, leaving the core and seeds. This would not serve the
function desired. Certain species of the Sarcoyuccas,
baccata, valida, and G'uatemalensis, having the papery core,
are said by Dr. Trelease f to fall early. It is quite prob-
able that this is a habit developed to aid in their dissemi-

* Trelease, Wm., ‘‘Notes and Observations. 7. Yucca Guatemalensis
Baker.’’? Mo, Botanical Garden, 5th Annual Report (St. Louis, 1894), p.
166.

DISSEMINATION AND LEAF REFLEXION OF YUCCA. 93

nation. They evidently fall as soon as mature while still
soft and tempting to animals. If this is so they may be
gathered regularly by certain small mammals that carry them
about and eat merely the pulp, discarding the core
and seeds as we discard the apple core. Such animals
would be prevented by the reflexed leaves from climbing
the stems of the plants and securing the fruits in this man-
ner. It is thus necessary that the fruits fall in order that
they may be secured and carried off. In Yucca aloifolia,
with which we will have most to do, this core does not
occur. In the still unripe but nearly mature fruits one
may get a suggestion of it in the rather firm, white mem-
brane which immediately surrounds the seeds. As the
fruit matures, bower it becomes soft throughout and
this is doubtless of decided importance in its dissemination
as it thus affords no resistance or hindrance to birds eating
the pulp from obtaining the seeds at the same time.

In regard to the dissemination of Sarcoyuccas, Trelease *
writes ‘* These fruits are well adapted to dissemination by
fruit-eating animals, especially birds, the seeds being thrown
away, but I do not know of any recorded observations on
their dissemination.’’? This was the condition of our under-
standing of their dissemination, when in the fall of 1892 I
came to Florida and was afforded opportunity to study Y.
aloifolia in its native home. One day a mocking bird was
noticed picking at the pulp of the ripe fruits and this led
to an investigation of the subject of its dissemination.

The fruit of Yucca aloifolia (Plate 47, fig. 1) is elon-
gated elliptical in outline, with usually a slight central
constriction, though this is by no means always the case.
They sometimes reach a length of 15 centimeters but are
usually from 8 to 10 centimeters long. In cross section
(Plate 47, fig. 2) the fruit is hexagonal in shape, caused
by the spreading of the nectar grooves in the ripening of
the fruit. The openings of the septal nectaries may be seen

* Trelease, Wm., “Further Studies of Yuccas and their Pollination,’’
l.c., p. 224,

94 MISSOURI BOTANICAL GARDEN.

in the upper portion of the ripe fruit on which the three
lobed style remains plainly distinguishable, merely having
slightly increased in size during the growth of the fruit
(see Plate 47, fig. 1). The fruit is fleshy throughout with
no indication of a core when fully mature.

In ripening, the fruit passes from green to purple and
finally to dark purple or almost black in age. The pulp,
which is very tender, is of a characteristic sweetish-bitter
taste. The bitter principle is not evident for a few seconds
after tasting the fruit; but like that of the Indian turnip,
though not so strong, soon makes itself known and persists
fora considerabletime. Dr. Trelease* quotes Mr. Burbidge
as having characterized the taste as resembling a mixture of
black currant jam and quinine. The fruits vary somewhat
in taste, some being quite pleasant and agreeable. A
careful selection of the fruits for a number of years would,
I think without question, result in developing valuable edible
varieties. The fruits, it is said, are sometimes eaten by
man. They, however, are surely not much sought and
man only aids in their dissemination incidentally, spreading
the plant by cultivating it as a showy garden perennial. It
is popularly reported that people are occasionally made sick
by eating the fruits.

Extended observations through three seasons have fully
confirmed the fact that the mocking bird (Mimus polyglot-
tus) is a very important factor in aloifolia dissemination,
especially in what may be termed long distance dissemina-
tion. The mocking bird is very abundant throughout
the range of alotfolia and may be the only animal dissem-
inator of this species. At least I have not been able to
observe any other bird or animal eating its fruits. It is
not improbable that the fruit of aloifolia has become es-
pecially adapted for dissemination by the mocking bird or
some small bird of similar habits. The core which occurs
in all other baccate fruited Yuccas known to the author,

* Trelease, Wm., “Notes and Observations. 7. Yucca Guatemalensés
Baker,” 1. c., p. 166,

DISSEMINATION AND LEAF REFLEXION OF yuccA. 99

would be fatal to dissemination by birds of this size and
nature.

Numerous fruiting plants of alotfolia have been exam-
ined and, in almost every case, some or all of the fruits
were found to be somewhat eaten. Sometimes only a
small portion of the pulp will be eaten, but again a large
part of the fruit will have been removed.

The seeds of aloifolia are comparatively thick, being
usually from 2 to 2.5 millimeters, and have a narrow
two-edged rim (Plate 47, fig. 4). They are blackish,
nearly the color of the pulp, and lie imbedded in this
tender substance but a short distance below the surface.
The birds in picking off portions of the pulp, which is
apparently a delicate morsel for them, soon come to the
seeds. These they evidently do not want, but in their
greediness they cannot avoid swallowing some of them.
The birds, while usually quite shy, may yet with a little
perseverance be observed in all the details of feeding. As
portions of the pulp are deftly picked off, frequently a
seed will adhere to the bill by the surrounding sticky pulp.
Many times I have seen the birds throw off these seeds by
a sidewise jerk of the head as a chicken does in trying to
remove dirt or hair from its bill. This results in throwing
the seeds to a considerable distance, frequently from 1 to
2 meters. Many of the seeds taken with the pulp are
apparently swallowed. All of the seeds removed from
the fruits through the agency of the birds are not thrown
to a distance in this manner or swallowed, but many of
them are merely loosened and drop down into the compact
rosette of erect leaves below. These either adhere to the
leaves by the sticky pulp surrounding them or, if dry, roll
down the leaves to the stem where they are effectually
lodged. All over the upper portion of the plant in this
stage, portions of the pulp are scattered, which were acci-
dentally thrown off with the seeds or dropped by the birds
in eating. These portions frequently contain seeds.

While the fruits of aloifolia are in this bird-disseminat-

96 MISSOURI BOTANICAL GARDEN.

ing stage the leaves immediately below the fruit cluster are
erect, that is, point upward forming an angle of from 30°
to 60° with the stem. They are linear lanceolate and rigid
with sharp horny end spines, and are usually from 35 to 65
centimeters long by from 3 to 5 centimeters wide. They
are very numerous and suddenly broaden out at the base,
greatly overlapping each other, so that an almost closed
cup is formed at the apex where the fruit cluster is borne.
The most of those seeds which fall, not being twitched off
by the bird, strike some one of the numerous upper leaves
and adhere until the rain washes them down to the stem
where they are held till a later stage of development when
the upper cluster of leaves, which are now erect, have be-
come reflexed. This stage will be described in detail later.

Many of the seeds are evidently swallowed by the mock-
ing birds with the surrounding pulp. In this case, as in
the majority of pulpy fruits, the seeds are swallowed unin-
jured and resist the action of digestion so that they pass
through the alimentary canal and are evacuated in good
condition for germination. Mocking bird dung, which had
fallen on the Yucca plants, was examined in many cases
and frequently found to contain aloifolia seeds apparently
uninjured.

The stomachs of several mocking birds were examined,
but, although in three cases almost the entire contents of
the stomach was composed of the pulp of aloifolia, no seeds
were discovered.

Having at my home a captured mocking bird I tested
him with aloifolia fruits and seeds. He had become quite
tame and had learned to pick flies from one’s finger. I
first took single seeds surrounded by some pulp. These he
greedily picked from my finger and swallowed without diffi-
culty. During the feeding several seeds were snapped
from his bill, and several were thrown off by a twitch of
the head in trying to free the pulp from the seeds. These
actions correspond exactly with those previously ob-
served in the field. The number of seeds swallowed was

DISSEMINATION AND LEAF REFLEXION OF YUCCA. 97

counted. One seed made the passage of the alimentary
canal and was evacuated in fifteen minutes after the first
seeds were swallowed. At the end of an hour all of the
seeds which were swallowed had been evacuated. The
next day I gave the bird fifteen aloifolia seeds from my F
finger at one feeding. In half an hour thirteen of these
had been evacuated and the others were evacuated before
the end of the hour.

I next gave the bird an entire fruit and watched him
pick off portions of the pulp, occasionally getting a seed.
The cage previous to this had been thoroughly cleaned.
During about four hours, the time he was given access to
the fruit, he ate and evacuated fifty-one seeds.

All of the details of the feeding were watched many
times with this tame bird and the important features of the
feeding, snapping and twitching the seeds, etc., were found
to correspond entirely with what had been observed in the
field. 2

The seeds which were evacuated were apparently in good
condition, but to surely determine this a number of them
were planted. <A very fair percentage of these germinated
and formed healthy young plants.

During the fall and winter, from November to February,
when the aloifolia fruits are ripening in greatest abundance
and are suitable, they apparently form the main food of
the mocking birds, judging from the frequency with which
one observes them feeding on the alotfolia fruits, and from
their excrement at this season of the year. In early fall,
during September and October, many alozfolia fruits ripen,
but at this season of the year the poke berry (Phytolacca
decandra) and the wild and cultivated persimmons are
conspicuous rivals as mocking bird food. By the middle of
November. these are principally gone, however, or remain
as dried fragments. The fruits of Duranta Plumieri
Jacq. and Melia Azederach L. play some part in supplying.
food for the mocking bird in late fall and winter.

While the fruits of aloifolia are almost always more or

7

98 MISSOURI BOTANICAL GARDEN.

less eaten by the mocking birds, and many seeds removed,
yet by far the larger portion of the fruit with its contained
seeds usually remains still hanging on the old fruit cluster.
The pulp of the fruit gradually dries up and shortly but
little remains of the heavy pulpy fruit but a light mass of
seeds cemented together by the dried up pulp. The break-
ing of the epidermis and eating here and there on a fruit
by the mocking birds greatly hastens this drying. As the
fruits dry they pass the stage when birds aid in their dis-
semination and now a secondary method of local dissem-
ination begins to act.

As the lower fruits of the cluster in Yucca alotfolia
reach maturity, one or more, seldom two or even three,
lateral buds, start up near the base of the peduncle and pro-
long the growth of the stem. A single bud soon simulates
the continuation of the main axis. Several buds form a
branched trunk.

In every case examined the bud or buds appear to develop
from the axil of a leaf about one series below the inflores-
cence, and in its development the bud quite uniformly splits
the base of the leaf which is directly above it. This is some-
what at variance with the observations of Dr. Mellichamp *
who found the bud of alotfolia to spring from exactly the
uppermost axil, at the base of the inflorescence.

The new rapidly growing shoot, which develops from
this bud, grows up within the circle of mature leaves beside
the old fruit cluster (Plates 45 and 46); which is
pushed to one side by the growth of this secondary branch,
which in turn bears an inflorescence.

Whether the inflorescence of Yucca is truly terminal,
morphologically speaking, and the bud lateral, I have not
yet been able to determine. Engelmann f speaks of the
inflorescence of Yucca as terminal; and Trelease mentions

* Quoted by Dr. Engelmann in his ‘‘ Notes on the Genus Yucca.’’
Trans. St. Louis Acad. Sci. III. (1873), p. 22.

+ Engelmann, Geo., ‘‘ Notes on the Genus Yucca,” p. 25. Reprinted
in “‘ The Botanical Works of Geo. Engelmann,”’ p. 282,

DISSEMINATION AND LEAF REFLEXION OF YUCCA. 99

the inflorescence of Yucca brevifolia as terminal: ‘‘ After
blooming, two or three stout branches usually develop by
the side of the original apex, which now has ended its
growth. When these have reached a length of two or
three feet each forms a terminal inflorescence and branches
in its turn, giving rise to a repeated forking or triparti-
tion.”’ * It does not appear, however, that a morphological
examination has been made to determine this point. In
the lack of definite information, I speak of the inflorescence
as terminal and the buds as lateral in this paper.

The periodical development from the base of the inflor-
escence, of the lateral buds which continue the stem in
aloifolia and some other Yuccas, results in forming a trunk
consisting of a certain number of units or segments which
are quite distinct. These may be designated ecological
phytomers or phytomeroids.t The branches formed at the
side of the old fruit cluster do not usually bloom the next
spring, but spend a year in their development, bloom the
second spring, and fruit the second fall or winter after they
start to develop. A period of two years is thus required for
the completion of one of the phytomeroids. No exceptions
to this have been observed though they probably occur.
Some plants bloom very early in the spring and mature their
fruits early in September. It is quite probable that the
lateral shoots of these early blooming plants may succeed
in making a sufficient growth to bloom the next spring,
probably rather late, and mature their fruits the next

* Trelease, Wm., ‘‘ Further Studies of Yuccas and their Pollination,’’
l.c., p. 194.

+ From phytomera (d¢utoy, plant and pepos, part) and edo¢g, form.
Phytomera or phytomers was used by Gray (Gray’s Botanical Text Book,
6th Ed., Vol. I., Structural Botany or Organography on the Basis of Mor-
phology, p. 7), to designate the units of a stem, each including an internode
and node with its leaf.

Phytomeroid as used here is apparently equivalent to what Gray has
called definite shoots (l.c. p. 49), and is what in German is known as
Langtrieb, Haupttried, etc. (R. Hartig, “‘ Lehrbuch der Anatomie und
Physiologie der Pflanzen,’’ p. 116, Pax, * Allgemeine Morphologie der
Pflanzen,’’ p. 24.)

100 MISSOURI BOTANICAL GARDEN.

winter. I am satisfied, however, that this is not frequently
the case. It is probable that certain branches may also
require more than two years for their development before
fruiting. I am inclined to think this not uncommon though
no specimens under continuous observation have been
known to do so.

One of the most marked and peculiar characters of
Yucca aloifolia and a number of other Yuccas is the
reflexing of the leaves. The reflexing does not take
place gradually as the leaves are formed, but corre-
sponds with the phytomeroids of the stem mentioned
above. Only the leaves belonging to the upper phyto-
meroid are erect, those of the preceding older phyto-
meroids are all reflexed more or less strongly, becoming
more and more closely appressed to the stem the nearer
they are located to the base. All of the leaves belonging
to a certain phytomeroid become reflexed at the same
time, and this reflexing takes place at a certain stage in the
development of the fruit and lateral shoot. About the time
that the lateral shoot or shoots which form at the base of
the old fruit cluster start their development, the leaves of
the upper phytomeroid to which the old fruit cluster be-
longs, begin slowly to reflex. As the lateral shoot develops
they become more and more strongly reflexed till by the time
the lateral shoot is half grown they have in most cases
come to point strongly downward and form an angle of
scme 45° or less with the stem (Plates 45 and 46).
This angle grows less and less each year until the leaves
have become closely appressed to the trunk. They finally
die and dry up, but remain persistent, clothing the lower
portion of the trunk (Plate 46). In cultivated aloifolia
these are usually pulled off to render the plants more
sightly, a practice liable to cause a misunderstanding of
the true habit of the plant unless this is remembered.

The leaves of the lateral shoots which start at the base of
the old inflorescence, remain rigid and erect all through
their growth and during flowering, as long, indeed, as these

oe
DISSEMINATION AND LEAF REFLEXION OF yucca. 101

lateral shoots form the uppermost phytomeroid. As the
fruits which they bear ripen and lateral buds in turn start
at the base of their fruit stems, they cease to be the terminal —
phytomeroids and their leaves begin to reflex.

The leaves of the upper phytomeroid while in the
erect stage are separated by a wide angle of divergence
from the reflexed leaves of the preceding phytomeroid,
the point of divergence being usually very marked (Plates
45 and 46).

The phytomeroids of the trunk are usually from one
to two feet in length and easily distinguishable even in
the old lower growths. A slight swelling in the trunk fre-
quently marks the place where one growth ended and the
next began.

The reflexing of the leaves is evidently due to a strong
epinastic growth of the tissue at the base of the leaf.
The tissue on the under surface at this point collapses
under the pressure exerted and becomes somewhat
wrinkled.

The time occupied in the reflexing of the leaves of a
phytomeroid varies considerably. The first marked reflex-
ing takes place quite rapidly, but the gradual drying
and compressing against the stem is a slow action, running
through several years. On starting to reflex, the leaves
pass below the horizontal and come to point downward in
from one to three months. After becoming reflexed, the
leaves remain fresh and apparently active through the
development of about two other phytomeroids, after
which they begin to dry up and finally die. The length
of time which they remain living apparently depends largely
upon the conditions to which the plant is exposed. Plants
growing on the sterile sand dunes along the coast or in old
neglected yards usually have very short leaves, which after
reflexing live only for a short period. Usually all leaves
are found to be dried up in such cases except those on the
two upper phytomeroids. Occasionally only those of the
uppermost phytomeroid will be found perfectly fresh and

102 MISSOURI BOTANICAL GARDEN.

active, the leaves beginning to dry up as soon as they
become reflexed. In plants grown in rich soils or in
well manured yards, the leaves of two or three phytom-
eroids may frequently be found green and apparently
active.

While the reflexing of the leaves in Yucca alotfolia is
probably not primarily to serve as an aid in seed dissemi-
nation it is yet certain that it is of considerable service
in this direction. The seeds which are dislodged by the
mocking birds in feeding, as described above, being heavy
fall into the crown of still erect leaves below, where they
usually adhere to some one of the numerous leaves or slide
down to the apex of the stem where the almost closed cup
formed by the broad bases of the leaves effectually arrests
and retains them until a later period of development when
these leaves become reflexed. If on the contrary the falling
seeds rattle through the cluster of erect leaves of the upper
phytomeroid, on reaching the upper reflexed leaves of the
lower phytomeroid which point downward and outward,
they are shot outward by these, the force of the fall serving
to throw them to a considerable distance from the parent
plant.

When the leaves of the upper phytomeroid, that bear-
ing the fruit cluster, have become reflexed, the seeds
which have been lodged in the cup formed by these before
their reflexion, are gradually dislodged and are thrown out-
ward by the reflexed leaves as they fall. Many may be
dislodged by rain and washed downward but can hardly
fail to be thrown outward as far as the ends of the leaves
of the parent plant. Those seeds which fall when dry
are frequently thrown outward to a considerable distance.
The majority of them strike the ground from one to one
and one-half meters distant from the base of the parent
plant. This distance depends on the height of the plant
and the angle to which the leaves are reflexed. I have
examined some typical but exceptional cases, where the
seeds could noticeably be seen to form a circle around the

TET, OR CEN ree prem

DISSEMINATION AND LEAF REFLEXION OF YUCCA. 103

plant at a distance of about a meter, having been thrown
there in falling by the action of the reflexed leaves.

While this is not the main method of dissemination, I
think that it may reasonably be claimed to form a second-
ary method of considerable importance.

After reaching maturity the fruits remain fresh and soft
for a few weeks but gradually dry up until nothing remains
but a dried fragment of the pulp which cements the seeds
together into a small light mass. Long before they have
reached this thoroughly dry condition they have passed
the stage when the birds will eat them. Sometimes the
dried fruits are broken off as a whole and these, falling,
strike the reflexed leaves and are thrown outward to some
distance as they fall to the ground. The fruits may now
be knocked about by man or animals or may become disin-
tegrated in the place where they fall. This would also serve
to disseminate the seeds to some extent. Many of the seeds
still remain sound and evidently capable of germination.

A still more peculiar aid to dissemination is found in the
larvee of the Bogus Yucca Moth (Prodoxus decipiens Riley)
which is very common here and probably throughout the
aloifolia region.

The eggs of Prodoxus are deposited principally in the
young flower stem, though it is said by Riley * that
they are also found frequently breeding in the fruits of the
indehiscent Yuccas like aloifolia. I have never observed
them in the fruits of aloifolia, though I have made no
special attempt to find them. The incision on the young
peduncle made by the ovipositor leaves a discoloration
and forms ultimately a sort of cicatrice which remains dis-
tinctly visible and as the stem dries becomes slightly
elevated (Plate 47, fig. 3, b). The larva burrows in
the tissue of the peduncle, molts according to Riley three
times, and acquires full growth in a month. ‘ It pre-
pares for hibernation in the autumn, a cocoon of white

* Riley, C. V., “Yucca Moth and Yucca Pollination,’? Mo. Bot.
Garden, 3d Annual Report (St. Louis, 1892), p. 128.

Cyt ee ee eee ee vee
ad

104 MISSOURI BOTANICAL GARDEN.

silk which is covered on the outside with castings and
which remains protected within the stem. Before making
the cocoon, however, it generally eats a passage way to
the outer covering of the stem and lines this with silk
leaving but a thin cap.’’ During the time that the fruits
are drying up, as described above, the larve of Prodoxus
are particularly active and in many cases their burrowing
succeeds in cutting the peduncle entirely off so that the
fruit cluster falls to the ground. I haveseen many clusters
cut off in more than one place so that they are separated
into several fragments. These portions of the fruit cluster
may each retain attached a number of the old more or less
dried up fruits containing many good seeds. As the frag-
ments of the fruit cluster fall they are not lodged in the
leaves as they undoubtedly would be if these remained
erect; but are directed outward so that they fall, as we
have seen that the seeds and single fruits do, on the ground
a short distance away from the parent plant. The dried
fruits are very light and the cluster with the protruding
pedicels of the old flowers, is easily caught and dragged
about by animals or man. Many of the seeds may be
disseminated in this way.

The lateral branch which shoots forth at the base of the
fruit cluster, starting its growth about the time that the
fruits ripen, grows rapidly and by the time the fruits have
passed the bird stage and approached the stage when the
cluster is commonly cut off by the Prodoxus larve, the
growth of the lateral branch has pushed the fruit cluster
considerably to one side. By this time the leaves of the
phytomeroid which bore the fruit cluster have usually
reflexed below the horizontal so that they allow the old
fruit cluster, when cut off, to fall to the ground without
resistance or hindrance and direct it outward and away
from the parent plant as it falls.

The cut end of the peduncle is usually smooth, showing
that the cut was extended entirely across the stem with the
exception of a very thin portion on the outside. This is

DISSEMINATION AND LEAF REFLEXION OF yucca. 105

evidently partially cut through and that which remains is
dry and brittle and breaks off by the weight of the cluster
(Plate 47, fig. 3, a).

The seeds of the old dried fruits are frequently ana
by the ravages of a beetle which burrows in the dried pulp
and seeds. Many of the seeds, however, remain uninjured
and apparently long retain their power of germination.

It is probable that the other Sarcoyuccas show the same
leaf reflexion that occurs in aloifolia, but the periods of
growth and other details may vary. So far as can be
determined from photographs and drawings of the var-
ious species of Yucca, it appears that of the Sarcoyuccas,
baccata,* Guatemalensis,t Schottii,t australis § and macro-
carpa || show this periodic leaf refiexing. Other species
may show the same phenomena but no data are at hand
from which this can be determined.

Of the Clistoyuccas, brevifolia { evidently shows beauti-
fully this periodic leaf reflexing, the old trunk remaining
covered by the persistent, reflexed, dried up leaves which
in age become closely appressed against the trunk.

Among the Chaenoyuccas, Y. elata, a stemmed species
which grows to a considerable height forming a rather
large tree in Arizona, is particularly interesting from the
fact that here among the capsular Yuccas according to Dr.
Trelease ** we find a counterpart of aloifolia in that the

* Trelease, Wm., ‘“‘Notes and Observations. 1. Detail Illustrations,
of Yucca,’’ 1. c., Plate 2.

+ Trelease, ‘‘ Further Studies of Yuccas and their Pollination,” LiiGes
Plates 1 and 2.

t Trelease, as in last citation, Plate 3.

§ Trelease, as in last citation, Plates 4 and 5.

| Merriam, C. Hart, “‘ Notes on the Geographical and Vertical Dis-
tribution of Cactuses, Yuccas, and Agave, in the Deserts and Desert
Ranges of Southern California, Southern Nevada, Northwestern Arizona
and Southwestern Utah.”? In “The Death Valley Expedition; A Biologi-
cal Survey of Parts of California, Nevada, Arizona, and Utah. Part Hi,”
North American Fauna, No. 7, p. 358, Plate 14.

q Trelease, as above, Plates 6, 7 and 8.

** Trelease, Wm., ‘“ Further Studies of Yuccas and their Pollination,”
l.c., p. 202, Plate 10.— Also “ Detail Illustrations of Yucca,” 1.c., Plate 9.

106 MISSOURI BOTANICAL GARDEN.

leaves become reflexed and persist on the lower portion of
the trunk, becoming dry and closely appressed. The pro-
longation of the trunk is also by lateral buds springing
from a point near the base of the peduncle. This species
has a much longer peduncle than any of the Sarcoyuccas
known to the writer and the inflorescence and fruit cluster
are raised considerably above the leaves so that these appar-
ently do not aid or interfere in any way in the dissemination
of the seeds. If this species is modified for wind dissem-
ination, as is the case with most capsular Yuccas, this
elevation of the fruit cluster on a long peduncle would be
of decided advantage.

The ecological significance of leaf reflexion in aloifolia
and other Yuccas is not very evident. Prof. Lester F.
Ward * has suggested that the reflexing of the leaves in the
succulent fruited Yuccas has for its function the protection
of the sweet pulpy fruits from the ravages of small animals
such as the raccoon, opossum, rodents, etc., which might
injure the seeds if the fruits were eaten by them. Any
animal attempting to climb the trunk of aloifolia or other
species having these reflexed leaves, meets an almost
impenetrable barrier in the numerous sharp spines of the
stiff reflexed leaves which cover the older portions of the
trunk, pointing downward like so many bayonets. Should
they attempt to reach the cluster of fruits by jumping up
from below in the case of low plants, or by jumping down
on them from neighboring trees, they are met by the rigid
points of the erect leaves of the upper phytomeroid, which
remain erect till the fruits begin to dry up and pass their
desirability.

Dr. Trelease f mentions having found fruits of baccata in
the crown of leaves where they had fallen on maturity and
had remained protected from rodents by the reflexed leaves.

* In a paper read before the Washington Biological Club.— Unpub-
lished.
t Trelease, “‘ Further Studies of Yuccas,”’ 1. c., p. 186.

DISSEMINATION AND LEAF REFLEXION OF YUCCA. 107

As the leaves of this phytomeroid become reflexed these
fruits would be freed and fall to the ground.

Dr. Merriam * mentions the reflexed leaves of arborescens
(brevifolia) as effectually preventing most animals from
climbing up the trunk from below. Dr. Merriam informs
me, however, that the wood rat succeeds in ascending the
trunk. I quote the following from a letter of February 6th,
1894: ‘*Mr. Vernon Bailey and I have examined many
trunks of the tree Yucca ( Y. arborescens) which had been
ascended by a small desert mammal known as the wood rat
(Neotoma Mexicana). The wood rats cut off the leaves of
the Yucca at the base and thus form a spiral groove or ladder
around the trunk. The leaves are used by the rats in the con-
struction of their bulky nests which are commonly made up
of spiny materials such as sharp splinters of rock, parts of
cactus, leaves of Yuccas, branches of spiny desert shrubs and
thelike. The heaps of rubbish thus formed over the mouths
of their burrows constitute a protection against coyotes and
other enemies.’’ This wood rat which is the only animal
known to ascend the Yucca trunk, may not have as its
main object the fruits to be found there. Dr. Merriam
informs me that he has found freshly cut leaves at a time
when there were no fruits on the plant: They may merely
‘desire the leaves for their nests or may feed on the tissue
of the base of the leaf.

The habit of certain of the cored species of Sarcoyuccas
in dropping their fruits as soon as mature, which was dis-
cussed in the early part of this paper, must be remembered
here. In these species, baccata, valida, and Guatemalensis,
the reflexed leaves could hardly be considered as a protec-
tion to the fruits when these, as soon as they mature and
become desirable, drop to the ground, — as we suspect in
order to be gathered and eaten by certain of those animals
which cannot climb the trunk, the core serving to protect

* Merriam, C. Hart, “‘ Notes on the Geographical and Vertical Distri-
bution of Cactuses, Yuccas and Agave in the Deserts of Southern
California * * * etc.,” 1. c., p. 353.

108 MISSOURI BOTANICAL GARDEN.

the seeds. In aloifolia, where there is no core and the
fruits remain persistent until they dry up, it may be that
the protection afforded the fruits by the reflexed leaves is of
considerable importance. All indications here point to the
fruit as being particularly modified for dissemination by
small birds. The sweet tender pulp would be a tempting
morsel for many animals, and not having a protecting core
many of the seeds might be destroyed.

The service which the reflexed leaves render in seed dis-
semination has already been discussed. I would not con-
sider, however, that this function has primarily had any
influence in leading to the development of the habit of leaf
reflexion. I am not disposed either to look upon the pro-
tection which this habit affords the fruits of certain species
as being the primary cause. Iam inclined to the opinion
that the primary cause must. be sought for among the
intricate relations between light and growth, coupled with
the habit of growth of the plant, the reflexion being nec-
essary to permit the accumulated seeds and rubbish caught
by the erect leaves to fallto the ground. It may be merely
the common habit shown by certain Monocotyledonous
plants of this nature, such as palms which have tufted tops
at the ends of tall trunks. The lower leaves of such trees
gradually assume a reflexed position, evidently due to the
pressure exerted by the development of other leaves above
and an attempt to assume the most advantageous position
in relation to the light. ‘The old leaves of palms gradually
become strongly reflexed and ultimately dry up and break
off, their bases frequently remaining attached for a time
and then rotting away, leaving a smooth trunk. If this”
habit were coupled with the development of phytomeroids
such as occur in Y. aloifolia one might reasonably expect
this leaf reflexion to become periodic, corresponding to the
phytomeroids.

The Clistoyuccas, having dry indehiscent fruits, include
two species, gloriosa and brevifolia. Of the dissemination
of gloriosa apparently nothing is known. While this plant

ee
Gt

See.
oe

DISSEMINATION AND LEAF REFLEXION OF yucca. 109

is common in cultivation it very seldom fruits and has not
been much studied.

Brevifolia differs from other known species in having a
very thick exocarp similar to that of alotfolia but which on
maturing becomes dry and spongy instead of pulpy. What
is known of the dissemination of this species is from the
observations of Dr. Trelease:* ‘‘ The fruits of this species
fall quickly after ripening either by a distinct disarticulation
or because of the brittleness of the pericarp at the base,
and their rounded form and very light specific gravity
render them well-developed ‘tumble fruits’ and point to
their dissemination over the dry sands of the desert by aid
of the strong winds which prevail there, the seeds being
liberated ultimately by the breaking of the fragile pericarp.”’

The Chaenoyuccas or capsular species, which have dry
erect capsules and light, strongly compressed seeds, are
typical wind-disseminated plants. The capsules open by a
gradual septicidal dehiscence from above downward and by
a dehiscence for acertain distance from the top through the
backs of the carpels ( Plate 47, fig. 5).

The seeds are very thin and flat, ranging in thickness
from .5 to 1.5 millimeters, and have a very slight wing (Plate
47, fig. 6), a flying apparatus of the simplest kind similar
to that in Iris, Tulipa and Agave.f They are arranged in
six rows and are gradually sifted out of the erect capsules
by the shaking of the fruit cluster by the wind or animals
and by the wind dipping into the open top of the cells.
The device is thus to secure the gradual scattering of the
seeds, a few at a time by gusts of wind, the wind serving
to carry the seeds for some distance.

The peduncle of the capsular species is usually quite
long, serving to raise the fruit cluster free of obstruction
to some height above the surrounding vegetation. /ilamen-
tosa and glauca, the most widely known capsular Yuccas,

* + Trelease, Wm., ‘‘ Further Studies of Yuccas and their Pollina -
tion,’’ 1. c., p. 224.
+ Hildebrand, ‘ Verbreitungsmittel der Pflanzen,’’ p. 16.

110 MISSOURI BOTANICAL GARDEN.

grow on prairies or in open fields, and elata is a plant which
frequents the open mesas of the Arizona plains. Those
Yuccas which are dependent on the wind for their dissem-
ination apparently always grow in open places where the
wind has free action upon them. The Sarcoyuccas on the
contrary frequently grow in wooded areas, mountainous
districts, ete. Aloifolia sometimes grows in dense ham-
mocks. Macrocarpa, Prof. Toumey reports as a strictly
mountainous plant, frequenting shady cafions in Arizona.*
Habitat evidently has important connection with the
methods of dissemination employed by the various species.

The peduncles in jilamentosa, some time after the fruits
have ripened, very commonly rot at the base and fall. This
serves to empty the few seeds which may remain in the
capsules at some distance from the parent plant. The
peduncles in this species vary, in Florida, from 3 to 12 feet
in length.

SUMMARY.

In Yuccas the three types of fruits, characterizing the
different groups, correspond to three types of dissemina-
tion.

The Sarcoyuccas, having fleshy fruits, are probably in-
tended for dissemination by fruit-eating animals.

In the fleshy fruits of baccata, valida and Guatemalensis
the seeds are surrounded by a papery core similar to that
of the apple. The fruits of these species drop early, prob-
ably as soon as mature. This seems to be a device to aid
in their dissemination. They are probably gathered by
- small mammals that eat the pulp, the seeds, which are
protected by the core, being discarded.

The fruit of alocfolia, which is fleshy throughout and
persistent, is principally disseminated by the mocking
bird. This is accomplished by the bird swallowing the
seeds while eating the pulp. The seeds remain uninjured

* Toumey, J. W., “‘ Notes on the Tree Flora of the Chiricahua Moun-
tains.’”” Garden and Forest, VIII. (Jan. 16, 1895), p. 22.

DISSEMINATION AND LEAF REFLEXION OF yucca. I11

and are evacuated in good condition for germination.
Many seeds are also scattered by being twitched to some
distance by the bird, in freeing its bill from the adhering
pulp and seeds. A tame mocking bird swallowed and
evacuated 51 seeds in 4 hours. A number of these were
afterwards germinated.

The trunk of aloifolia is prolonged by the lateral branch
or branches which spring from near the base of the old
fruit cluster. This branch prolongs the trunk and in turn
bears an inflorescence and fruit, about two years being re-
quired usually to complete the period of development,
The aloifolia trunk in its development may thus be divided
into definite stages or segments, which in the lack of a
term to designate them are here called phytomeroids. The
lateral branch which continues the stem starts to develop
about the time that the fleshy fruits begin todry up. At
about the same time the erect leaves of the phytomeroid
to which the fruit cluster belongs begin to reflex, and soon
come to point strongly downward. The reflexed persistent
leaves of the older phytomeroids become closely appressed
against the stem and, pointing downward like a series of
bayonets, prevent small mammals from climbing the trunk.
The reflexed leaves aid as a secondary method of dissem-
ination. Seeds and dry fruits falling on these are directed
or thrown outward so that they fall at a distance of a meter
or so from the parent plant.

Old fruit clusters of aloifolia, after the fruits have
passed the stage for bird eating by drying up, are cut off
by the larve of Prodoxus decipiens. The leaves of the
phytomeroid to which these fruit clusters belong, have by
this time become reflexed and thus do not prevent the fall
of the cluster but direct it outward away from the parent
plant.

The reflexed leaves by preventing animals from climbing
the trunk also serve to protect the fruits from the ravages
of those animals that might injure the seeds.

The protection of the fruits and aid in seed dissemina-

112 MISSOURI BOTANICAL GARDEN.

tion are not thought to be the primary factors leading to
the development of the habit of leaf reflexion.

The primary cause which leads to the development of
this habit is probably to be found in the relations between
light and growth, and the necessity for some such habit to
free the plant from rubbish which becomes lodged in the
erect crown of rigid leaves.

Yucca brevifolia, illustrating the Clistoyuccas, is unique
in having a light spongy fruit which falls when mature
and is blown about as a rolling fruit, gradually breaking up
and dropping the seeds.

The capsular Yuccas, like filamentosa and glauca, are de-
veloped for wind dissemination. They have very thin light
seeds which are gradually sifted out of the capsules and
blown to some distance by the wind.

The wind disseminated Yuccas grow in open places where
the wind can have free action upon them.

The rotting off at the base and falling of the old fruit
stems of jilamentosa serves finally to throw out all seeds
which remain in the capsules.

Eustis, Florida. January 12th, 1895.

EXPLANATION OF PLATES ILLUSTRATING THE DISSEMINATION
OF YUCCA.

Plate 45.— Yucca aloifolia, with lateral branch developing from near
the base of the peduncle. The leaves of the phytomeroid which bore
the inflorescence have begun to reflex.—Drawn by Mrs. Webber, from
a photograph ofa plant at Bartow, Florida, January 24, 1894.

Plate 46. — Y. aloifolia, with lateral branches starting at the base of
the old peduncles, and the leaves of the upper phytomeroid starting to
reflex.—From a photograph of plants on Anastatia Island, Fla., Jan. 23,
1895.

Plate 47.— Fig. 1. Ripe fruit of Y. aloifolia — Fig. 2. Cross section of
aloifolia fruit. — Fig. 3. Basal end of old peduncle of Y. aloifolia cut off
at a by Prodoxus decipiens; bb, scars resulting from oviposition of Pro-
doxus female; cc, perforations made by the chrysalis of Prodoxus in
issuing; d, portion of the epidermis cut away by the burrowing of Pro-
doxus; ee, tunnels in the tissue made by Prodoxus larve.— Fig 4. Y.
aloifolia seed, and cross section of seed showing ruminated endosperm.—
Fig. 5. Capsule of Y. jfilamentosa. — Fig. 6. Y. silamentosa seed, and
cross section. — All nat. size, drawn from nature by Mrs. Webber.

REPT. MO. BOT. GARD., VOL. 6.

PLATE 45.

hh : ‘f hf i\

We Ly
i aa
i my :
i!

Oe

1 on BNP els Lire WSS
Fontan "ae
he ly ‘

YUCCA

ALOIFOLIA.,.

REPT. Mo. Bor. GARD., VOL. 6. PLATE 47.

YUCCA ALOIFOLIA AND FILAMENTOSA.

NOTES AND OBSERVATIONS

ON NEW OR LITTLE KNOWN SPECIES.

BY JARED G. SMITH.

Srpa PauMeEr!I, Baker?

Slender, erect or ascending perennial, 4 to 6 dm. high,
sparingly branched from the base, the stems cinereous
stellate pubescent; leaves remote, subcordate at the base,
obtuse at the apex, from ovate-triangular to triangu-
lar oblong, usually with rounded basal lobes, serrate
or entire, sparsely stellate-pubescent above, densely stel-
late-tomentose below, the blade usually longer than the
petiole; stipules subulate; peduncles slender, axillary, 3 to
5 cm. long, geniculate above; calyx tube terete; sepals
ovate, abruptly acute, stellate-canescent, about 4 mm. long,
a third as long as the yellow or purple petals; carpels 7 to
9, dehiscent, sparsely pubescent on the back, the sides
latticed; seed obliquely obovate-triangular, glandular punc-
tate, the hilum obovate, smooth. Collected by G. C. Nealley,
near Corpus Christi, Texas, in the summer of 1894, and
reported by him as being rare.— Plate 48.

This form approaches S. Palmeri, Baker, Journ. of
Botany, 30, 295 (1892), a species of section Malvinda,
founded on Palmer’s No. 1038, and may not be distinct from
it. I have not had the opportunity of examining authentic
specimens, but Nealley’s plant does not fully agree with
Baker’s description.

Fueosra Drummonpi, Gray. Pl. Wright. 1, 23 (1852).

This plant, of which habit sketch and detail drawings are

appended, was collected by G. C. Nealley, near Corpus

Christi, Texas, in 1894. The locality is re-established
8 (113)

114 MISSOURI BOTANICAL GARDEN,

after a lapse of sixty years, the plant not having been found
in the United States since Drummond’s original collection
in 1834, The specimens agree with the published descrip-
tions and with herbarium material of the same species from
South America.—Plate 49.

SEpuM TEXANUM, 0. sp.

Herbaceous perennial; glabrous, ascending, 1.5 to 2 dm.
high, cespitose, sparingly branched from near the base;
leaves sparse, thick fleshy, obovate to ovate lanceolate,
obtuse or mostly acute or acuminate, broad at the base,
sessile, 2 to 2.5cm. long; flowers sessile or short pedicelled,
solitary or two or three together, forming a sub-secund
raceme, or paniculately branched in the larger specimens,
rather remotely flowered below; sepals oblong, acute;
petals 5, rosy yellow, narrowly oblong, thickened and
apiculate at the apex, one-half longer than the sepals;
scales minute, truncate; stamens 10, shorter than the
petals, the filaments flattened; anthers bright yellow; car-
pels 5, inflated, many seeded, with a slender beak tipped
by the very short persistent style, dehiscent down the ven-
tral suture. Collected by G. C. Nealley, in flower, the last
week in October, 1894, growing in the chapparal near Cor-
pus Christi, Texas. It is most closely related to JS.
Wrightii, Gray, but differs in the form of the inflorescence
and in the much larger leaves. The type specimen is in
the herbarium of the Missouri Botanical Garden.—Plate 50.

ELYTRARIA BROMOIDES, Oersted.

This and STeNANDRIUM DULCE, Nees, are two interesting
little Texan plants collected near Corpus Christi by Mr.
Nealley. Habit sketches and detail drawings of flower and
fruit are given in plate 51.

ZEPHRYANTHES PULCHELLA, N. sp.
Bulb globose, 1 to 2 cm. in diameter, covered with dark
brown tunics; neck 3 to 4 cm. long; leaves 3 or 4, appear-

NEW AND LITTLE KNOWN SPECIES. 115

ing with the flower, 1 to 2mm. wide, 1.5 to 2.5 dm. long;
scape slender, 1.5 to 2 dm. long; flower solitary, erect;
spathe 2 to 2.5 cm. long, tubular for half its length, slightly
exceeding the pedicel, entire at the apex; perianth erect, 2
cm. long, the segments oblanceolate, acute, bright yellow;
tube 5 mm. long; stamens inserted at the throat, about
half as long as the perianth segments; style equaling the
stamens; stigma capitate, 3-lobed. Collected at Corpus
Christi, Texas, summer of 1894, by G. C. Nealley. It is
intermediate between Z. Texana, Herb. and Z. Longifolia,
Hemsl. For purposes of comparison I have figured flowers
of these latter species on the same plate. The pedicel
length is a very constant character in all the herbarium
specimens of the two older species which I have examined,
and it is one of the principal specific characters used by
Baker in his monograph of the Amaryllidaceae. Hence, I
have considered it best to separate out this material as a
new species.—Plate 52.

SaGiTTaRIia.

Since publishing the monograph of the North American
species of Sagittaria and Lophotocarpus, I have received
material of the following new species of Sagittaria from
Florida.

§§§ Integrifoliae.
+ ++ Achenium obovate, with a short lateral beak; filament dilated
at the base, pubescent; bracts connate, glabrous.

S. ISOETIFORMIS, n. sp.

Monoecious, 1 dm. high; leaves slender, terete through-
out, or dilating into a short linear blade 1 to 2 mm. wide;
scape simple, slender, erect, equaling the leaves, with
mostly a single verticil; bracts ovate, acute, 2 to 3 mm.
long, connate; fertile pedicels shorter than the sterile, very
slender, 15 to 18 mm. long; sterile pedicels filiform, 2 to 3
cm. long; stamens about 9; anthers equaling the filaments ;
achenium about 1 mm. long, dorsally crested, with an arched

116 MISSOURI BOTANICAL GARDEN.

lateral wing, and with 2 pairs of subepidermal resin pas-
sages; fruiting head small, 6 mm. in diameter; flowers
white, 12 to 14 mm. across. Only 1 or 2 flowers fertile.
Growing in tufts at the nodes of very slender stolons, in
water 3 to 10cm. deep. Collected at Eustis, Lake Co.,
Florida, March 22, 1894, by Geo. V. Nash, and by A. S.
Hitchcock in July. It is common along the sandy lake
margins and forms quite extensive patches in the shallow
water, its slender interlacing stolons bearing tufts of leaves,
and rooting at the nodes. It is more closely related to S.
graminea, Michx. than to any other species. In the arti-
ficial key to the genus it should be inserted after S. macro-
carpa, from which it is easily separated by the smaller
achenium. The type is in the Herbarium of the Missouri
Botanical Garden.— Plate 53.

ERAGROSTIS SPOROBOLOIDES, Smith and Bush, n. sp.

Tall perennial; culms 5 to 7 dm. high, erect, simple or
branching from the base; sheaths longer than the inter-
nodes, pilose, especially at the throat; leaves soft, flat,
becoming involute and filiform above, 5 to 8 mm. wide, 4
to 6 dm. long, scabrous on the upper surface, smooth
below; panicle oblong, not emergent from the upper
sheath, half as long as the culm; branches of the panicle
capillary, single or in pairs, minutely scabrous; spikelets
2.5 mm. long, 2 or 3 flowered, on scabrous capillary ped-
icels 1 to 3 cm. long; empty glumes one-nerved, ovate-
lanceolate, acute, scabrous on the keel, two-thirds as long
as the lowest floret ; flowering glume ovate, acute, 3-nerved,
the nerves scabrous above; caryopsis not seen. Collected
at Sapulpa, Indian Territory, July, 1894, by B. F. Bush.
This grass is closely related to Hragrostis capillaris, Nees,
but is perennial, and has much longer and wider leaves.
Immature specimens appear at first sight only one-flowered,
and might be mistaken for excessively long leaved forms
of Sporobolus asperifolius Thurb., a resemblance which is
heightened by the very pilose sheaths; hence the specific

NEW AND LITTLE KNOWN SPECIES. 117

name sporoboloides. The type specimens are in the her-
barium of the Missouri Botanical Garden.—Plate 54.

ERAGROSTIS GRANDIFLORA, Smith and Bush, n. sp.

Perennial; culms simple, 1 meter high, the nodes much
elongated; panicle large, occupying more than half the
culm, 6 dm. long; panicle branches slender, ascending, 12
to 15 em. long, the lower pilose in the axils; empty glumes
lanceolate, very acute, a little shorter than the lowest
florets, 2.5 mm. long, the first one-, the second three-
nerved; rachilla glabrous, articulate; spikelet 10 to 20
flowered, 1 to 2 cm. long, longer than the scabrous capil-
lary pedicels; flowering glume 3 mm. long, ovate, acute,
glabrous, a little longer than the palet ; caryopsis 1 mm.
long, truncate at each end, furrowed; leaves ample, 3 or
4 dm. or more long, 7 to 10 mm. wide, attenuate and fili-
form at the apex, scabrous above and on the margins ;
sheaths smooth or hirsute above, and pilose at the throat.
This species is most closely related to Hragrostis tenuis,
Gray, but is easily distinguished by its elongated spikelets.
Collected at Sapulpa, Indian Territory, Oct. 1, 1894, by
B. F. Bush. Type specimen in the herbarium of the
Missouri Botanical Garden.— Plate 55.

Eracrostis BeyRICHII, n. sp.

Low cespitose perennial, 1 to 2 dm. high; culms genicu-
late below, ascending or erect, much branched from the
base, smooth, terete, glabrous, mostly with 3 nodes;
sheath striate, smooth, shorter than the internode, ciliate
at the throat; ligule very short; leaves soft, smooth or
very minutely scabrous above, linear, 6 to 8 or 15 cm. long,
1.5 to 2 mm. wide, soon involute and filiform, attenuate
above, usually much shorter than the culms; spikelets
sessile or short pedicelled and densely fasciculate in an
interrupted spike-like or capitate panicle, linear-oblong,
strongly compressed; 15 to 40 flowered, mostly 10 to 15 or
rarely 25 mm. long, 5 mm. broad; panicle branches single,

118 MISSOURI BOTANICAL GARDEN.

ciliate in the axils; empty glumes lanceolate acuminate,
over half as long as the lowest florets, the first one-, the
second three-nerved, tinged with purple; flowering glume
and palet pale straw color; flowering glume lanceolate
acuminate, 3-nerved, 4 mm. long, keel and lateral nerves
scabrous ; palet oblong, 3 as long, 2-nerved, scabrous on the
margins and nerves below, erose at the apex; caryopsis
terete, linear, 1 mm. long. This species is dedicated to
Beyrich, who first collected it in Arkansas in 1834. Speci-
mens have also been examined from Dallas, Texas, collected
by Reverchon, 499, 1874, and by Prof. Geo. W. Letter-
man, July 25,1881. Itis most closely related to Hragros-
tis oxylepis, Torr., but differs in habit and in the very acute
flowering glumes and narrow spikelets. Type in the
Bernhardi Herbarium.— Plate 56.

ERAGROSTIS SECUNDIFLORA, Presl, Reliq. Haenk. 1, 276
(1830). |

A specimen of this, collected by Haenke in Mexico, is in
the Bernhardi Herbarium. It is identical with Hragrostis
oxylepis, Torr., Rep. Bot. Whipple Exped. 156 (1857);
Poa interrupta, Nutt. Trans. Amer. Phil. Soc. (n. ser.) 5,
146 (1837), not Lam. Illustr. 1, 185 (1791); Eragrostis
interrupta (Nutt.), Trelease, Ann. Rep. Geol. Surv. Ark. 6,
237 (1888), not Beauv. Agrost. 71 (1812); and Eragrostis
Verae-Crucis, Rupr. Bull. Acad. Brux. 9, reimp. 9 (1842).
I have examined specimens as follows : — Mexico: Haenke,
without locality or date; and Liebmann 538 (2. Verae-
Crucis, Rupr.), Pochutla, La Galera, Oct., 1842. New
Mexico: Fendler 913, 1847. Texas: Drummond 228, 321 and
325, 1832-1834; Lindheimer 32, Galveston, April 1843,
734, 735, 1847, and Granite Hills of the Llano, 464,
Oct., 1847; Ravenel, Indianola, May 3, 1869; Pringle 1969,
July 28, 1888; Pammel, Brazos Co., Aug., 1888; Letter-
man, Texarkana, Oct. 18, 1894; Heller 1802, Corpus Christi,
May 28-31, 1894. Indian Territory: Torrey, Marcy’s
Expedition, Wichita Mts. 1852, in the Buckley Herbarium ;

NEW AND LITTLE KNOWN SPECIES. 119

Bush 823, 824, Arkansas, July 23, 1894. Arkansas:
Engelmann, Arkansas R. below Little Rock, July, 1835.
Mississippi: Seymour 35, (Tracey Coll. ) Starkville, Sept.,
1891. Florida: Dr. Leavenworth, ‘‘Gulf of Mexico.”
Good Plates are given by Torrey: Marcy’s Red Riv. of
Louis. Exp., plate 19 (1854); and by Dr. Vasey: Grasses
of the Southwest, Part 2, plate 48 (1891).

The Bernhardi Herbarium contains a large number of
Haenke’s grasses, the labels of which are almost certainly
in Presl’s handwriting. These have not all been separated
out, but when they are, further notes may be made con-
cerning them.

EXPLANATION OF PLATES.

Plate 48. Sida Palmeri, Baker (?) — Habit sketch, half size; fruit,
X 5, seed, X 7.

Plate 49. Fugosia Drummondii, Gray.— Habit sketch and section of
flower, natural size.

Plate 50. Sedum Texanum, J. G. Smith.— Habit sketch, natural size;
flower and isolated petal, X 3; apex of petal, < 6.

Plate 51. Elytraria bromoides, Oerst.— 1, habit sketch, natural size; 2,
bracts and calyx, < 3; 3, corolla < 2; 4, capsule, X 5; 5, stamen, >< 5B.
Stenandrium dulce, Nees.— 6, habit sketch, natural size; 7, corolla < 2;
8, stamens, X 5; 9, calyx, X 3; 10, pistil, X 5; 11, capsule and seeds,
x 3

Plate 52. Zephryanthes pulchella, J. G. Smith.—1, habit sketch, natural
size; 2 and 3, styleand stamen, X 3. Z. longifolia Hemsl.— 4, flower and
spathe, natural size; 5 and 6, stamen and style, X 3. Z. Texana Herb.—
7, flower and spathe, natural size; 8 and 9, stamen and style, < 3.

Plate 53. Sagittaria isoetiformis, J. G. Smith.— Habit sketch, natural
size; stamen; and cross section, dorsal and side views of achenium,
10; a cross section of petiole, < 30.

Plate 54. Eragrostis sporoboloides, J. G. Smith and B. ¥. Bush. —Habit
sketch, one-half natural size; spikelet and flowering glume, X 5.

Plate 55. Eragrostis grandiflora, J. G. Smith and B. F. Bush. — Habit
sketch, one-half natural size; offshoot from base of plant with immature
spikelets, natural size; base of spikelet, flowering glume and palet, < 5;
caryopsis, < 15.

Plate 56. Eragrostis Beyrichii, J. G. Smith.— Habit sketch, one-half
natural size; base of spikelet, and flowering glume, X 4; 1, empty
glumes, < 4; 2, face and side view of palet, X 4; caryopsis < 15.

The plates were prepared from drawings made by Miss Grace E.
Johnson, from specimens preserved in the Herbarium of the Missouri
Botanical Garden.

Rept. Mo. Bor. GARD., VOL. 6. PLATE 48.

SIDA PALMERI?

REPT. MO, BOT. GARD., VOL. 6. PLATE 49.

|
i]
: +
“|
Sita

FUGOSIA DRUMMONDII.

PLATE 50.

Mo. Bor. GARD., VOL. 6.

REPT.

ll

SEDUM TEXANUM.

PLATE 51.

REPT, MO. BOT. GARD., VOL, 6.

ELYLRARIA AND STENANDRIUM.

REPT. Mo. Bot. GARD., VOL. 6. PLATE 52.

ZEPHYRANTHES.

Repr. Mo. Bot. GARD., VOL. 6. PLATE 53.

SAGITTARIA ISOETIFORMIS.

PLATE 55.

REPT, MO, BOT. GARD., VOL. 6.

et

a

<j

ee

SES

para
Ss

Ghf

RAGROSTIS GRANDIFLORA.

7

E

vy]

Rept. Mo. Bot. GARD., VOL. 6.

PLATE

56.

ERAGROSTIS BEYRICHII.

NOTES ON THE MOUND FLORA OF ATCHISON COUNTY,
MISSOURI.

BY B. F. BUSH.

Having concluded that Atchison County, Missouri, on
account of its extreme northwestern position, would prove
a fertile field for botanizing, I consulted with Dr. Trelease
upon the advisability of making a thorough investigation of
that part of the State, and he very generously placed me
in shape to begin the work.

About the middle of August, 1893, I made my first trip
to that place, and was struck at once by the remarkable
line of loess mounds running parallel with the Missouri
River in Missouri, from Hamburg in Iowa, down to a few
miles south of Saint Joseph, Missouri, where the last
bald-headed mound appears.

These singular mounds are evidently the result of the
glacial period, when great masses of ice were borne toward
the equator, until they were stranded along the low rocky
bluffs of the Missouri River, and as they melted, the mud
and debris settled to the bottom and thus formed these
mounds.

These mounds are very irregular and jagged, running in
straight lines, or else in fantastic loops and curves, often
abruptly terminating in a very high perpendicular wall,
which is almost unaffected by the action of the weather:
or commonly descending gently to the level bottom which
surrounds all sides except on the east. ;

At a distance these mounds appear to be terraced, as if
furrowed by the plow, but on a closer examination it is
revealed that the apparent terraces are trails made by cattle,
which are nearly always to be found grazing on them.
These trails are about four feet apart and either follow a
horizontal line around the mounds, or else gradually incline

(121)

122 MISSOURI BOTANICAL GARDEN.

upward to the top of them, and a well defined trail leads
from the top of one mound down the side and along the
low backbone to the side and top of the next mound, and
so on to all the mounds.

I have found these trails of considerable advantage in
climbing over and around the mounds, and in going from
one mound to another, and I have never seen one isolated
mound in Atchison County.

The south and west sides are those which are steep and
precipitous, but sometimes they descend gradually to the
bottom lands at their bases. The north and east sides are
usually descending and are covered with a short thick
growth of trees and shrubs near their bases, but which does
not rise as high up on the north side as on the east, as it
sometimes reaches quite to the top on the latter.

The texture of the soil is a thin, slightly sandy loess,
very compact and hard, so much so, that names carved in
the face of the steep sides endure for many years. The
fertility of the soil is very low and supports only such plants
as will thrive in similar conditions, being covered with a
short growth of grasses, such as Bouteloua hirsuta, B. oli-
gostachya and Sporobolus airoides.

There is a belt of heavy timber back of the mounds,
which averages about one mile in width, and which is some-
what lower than the mounds, and back of this is a high
rolling prairie, which is considerably lower than the mounds.
Between the mounds and the Missouri River is an extremely
fertile gombo bottom-prairie which is sometimes several
miles in width, and often not a mile wide, which has a flora
very similar to the general Missouri River bottom flora,
except that a few of the peculiar species of the mounds are
here present and should not be considered as characteristic
of it.

Where the south and west sides are steep and precipitous,
there are to be found in abundance such plants as Yucca
glauca, Gaura coccinea, Psoralea argophylla, Meriola ser-
rulata and Sporobolus airoides; where they descend gradu-

NOTES ON MOUND FLORA OF ATCHISON CO., MO. 123

ally to the bottom below, such plants as Pentstemon grandi-
florus, Astragalus lotiflorus and Parosela enneandra are
found. On the north there are usually pockets running up
the sides of the mounds which are generally covered with
dwarfed trees and shrubs, principally Corylus Americana,
Prunus Virginiana, Ostrya Virginica, Rhus glabra, and
Salix humilis.

The tops of these mounds are the most peculiar feature
of the whole country, for they present from a distance a
very much denuded appearance, as if entirely devoid of
vegetation. They vary considerably in height, some not
being one-third as high as the most elevated, and one of
the chain is said to be the highest point in the State, and
no doubt to this great elevation is due their peculiar flora.

The characteristic species of these mounds are Pents-
temon grandiflorus, Spiesia Lamberti, Castilleja sessiliflora,
Psoralea argophylla, Gaura coccinea, Glycyrrhiza lepidota,
Yucca glauca, Meriola serrulata, Sporobolus airoides,
Anemone cylindrica, Lacinaria punctata, Bouteloua hirsuta,
B. oligostachya, Astragalus lotiflorus, Gerardia aspera,
Parosela enneandra, Lithospermum angustifolium, Lygo-
desmia juncea, Lactuca pulchella, Onosmodium molle, and
Houstonia angustifolia, these being the predominant plants,
and not occurring in abundance elsewhere in the State.

The list here appended is based on specimens now in the
Herbarium of the Missouri Botanical Garden which were
also included in the sets distributed in 1893.

ANEMONE CYLinpRIca A. Gray.

Common on the tops of the mounds in bare ground:
commonly with several peduncles involucellate in the
middle. (Watson No. 3.)

ANEMONE Virainiana L.

Common along the bases of the mounds, and in the hazel
thickets. A.Canadensis L. (Watson No. 4), was abundant
in the bottom just below. (Watson No. 5.)

124 MISSOURI BOTANICAL GARDEN.

DELPHINIUM CAROLINIANUM Walt.

Common on the tops of all the mounds. The pistils are
commonly from three to seven, as I have noticed in the
plants of Jackson County, Missouri. (Watson No. 8.)

AcTAEA ALBA (L.) Mill.
Rarely seen on the wooded eastern slopes of the mounds.

(Watson No. 9.)

CLEOME SERRULATA Pursh.

Abundant on the lower part of the mounds. Has a very
fetid odor, on account of which it is called Stinking Clover
by the country people. (Watson No. 16.)

VIOLA PEDATIFIDA Don.

Common all over the bare parts of the mounds. V.
obliqua Hill (Watson No. 21), was common in the woods
on the east of the mounds. (Watson No. 19.)

Vioua PALMATA L.
Very common along the bases of the mounds. (Watson
No. 20.)

Hypericum Ascyron L.

On the northern sides of several of the mounds. At
one mound, I estimated that there were between ten
and fifteen thousand plants in one patch, about four miles
northeast of Watson. (Watson No. 37.)

Linum sutcatum Ridd.

On the bare tops and sides of the mounds, usually with
Gerardia aspera Dougl., which it not a little resembles.
(Watson No. 42.)

CEANOTHUS OvaTus Desf.

Very common on the sides of the mounds, and appear-
ing quite distinct from the next in appearance and mode of
growth. (Watson No. 120.)

NOTES ON MOUND FLORA OF ATCHISON CO., MO. 125

CEANOTHUS OVATUS PUBESCENS T. & G.
Also quite common in same situations as the foregoing.
(Watson No. 121.)

POLYGALA VERTICILLATA L.
Quite common on the bare parts of the mounds. (Wat-
son No. 184.)

TRIFOLIUM REFLEXUM L.
Common over most parts of the mounds in bare ground.
(Watson No. 187.)

PsoRALEA ARGOPHYLLA Pursh.

Abundant on the steep precipitous faces of the mounds.
One of the most striking species of the whole mound
flora. (Watson No. 211.)

AMORPHA CANESCENS Pursh.

Common on the sides of the mounds. Specimens
nearly two meters in height were observed at several
places. (Watson No. 217.)

PaROsELA ENNEANDRA (Nutt. ) Britton.
Abundant on all the mounds, but mostly on the steep
sides to the south and west. (Watson No. 220.)

Paroseva Dauea (L.) Britton.

Common on the lower part of the mounds, and along the
backbones, and still more common on the bottom prairie
below. (Watson No. 219.)

KuHNISTERA PURPUREA ( Vent.) MacM.

Abundant along the backbones connecting the mounds.
The heads are all very short-oblong or oval. (Watson
No. 222.)

KUuHNISTERA CANDIDA ( Willd.) Kuntze.
With, and as abundant as the last, and heads also much
reduced. (Watson No. 221.)

126 MISSOURI BOTANICAL GARDEN.

ASTRAGALUS LOTIFLORUS Hook.

(A. elatiocarpus Sheld. Bull. Minn. Geol. and Nat. Hist.
Surv. n. 9: 20. 1894, at least as toformabrachypus. A.
lotiflorus formae brachypus and pedunculosus of Gray.)

Abundant on all the gentle slopes and along the back-
bones of the mounds. When I first collected this species
in August, 1893, it was in fruit, with very short or no
peduncles, which led Dr. Trelease and myself to place it
with the forma brachypus of Gray.

On April 20th, 1894, I collected it again, but this time
it was in flower and with very long peduncles. ‘This led
me to write Dr. Trelease the statement that the earlier
flowers were very long peduncled and infertile, while later
on it produced fertile flowers on very short peduncles; for
some last year’s fruits were still on some plants and no
sign of long peduncled fruit had been observed the year
before. On June Ist, I again collected the plant and this
time it had fruit on both long and short peduncles On
July 11th, the long peduncles had mostly dropped off the
plants, leaving only the short peduncled fruit. This
clearly shows that the species has both long and short-
peduncled fruit, and that the formae brachypus and pe-
dunculosus of Gray occur on the same plants. I strongly
suspect that the Astragalus elatiocarpus of Sheldon is an
entirely different species. (Watson Nos. 198, 199, 200
and 203.)

Spres1a Lamperti (Pursh) Kuntze.

Common on the steep precipitous sides of the highest of
the mounds. The roots are exceedingly long and tough,
and often are seen hanging over the bluffs from which the
earth has crumbled away and left them. (Watson No.
204.)

GLycyRRHIZA LEPIDOTA Pursh.

Common at the bases of the mounds in bare and some-

what gravelly ground. (Watson No. 205.)

NOTES ON MOUND FLORA OF ATCHISON CO., MO. 127

PHASEOLUS PAUCIFLORUS Benth.

Common along the lower part of the mounds, and along
the backbones. P. angulosa Missouriensis ( Wats. ), Britt.
(Watson No. 213), was tolerably common in marshy
places on the bottom-prairie near the mounds. (Watson
No. 212.)

Acuan Ixurnoensis (Michx.) Kuntze.
Very common in the bare gravelly ground at the base of
the mounds. (Watson No. 214.)

Prunus Vireiniana L.

Abundant in the pockets on the north sides of the
mounds, and is said by the country people to rarely fruit.
(Watson No. 243.)

Rosa ARKANSANA Porter.

Abundant on the lower part of the mounds and along
the backbones. The receptacles are unusually large, rang-
ing from 12 mm. to 18 mm. in thickness, and the stems of
the season are often 25 mm. thick and exceedingly prickly.
(Watson No. 246.)

AMELANCHIER CANADENSIS (L.) Medic.

Common on the east sides of the mounds, and in the
pockets on the north, often becoming quite arborescent.
( Watson No. 313.)

MERIOLA SERRULATA (Nutt. ) Walp.

Abundant on the tops and sides of all the mounds.
The flowers are from 12 to 15 mm. wide. (Watson No.
330. )

GAURA PARVIFLORA Dougl.
Common near the base of the mounds, and along the
backbones. (Watson No. 334.)

128 MISSOURI BOTANICAL GARDEN.

GAURA COCCINEA Pursh.
Abundant on the steep faces of most of the mounds.
(Watson No. 333.)

SYMPHORICARPUS OCCIDENTALIS Hook.

Abundant at the base of the mounds, and in the pockets
on the north sides. S. Symphoricarpus (L.) MacM.
(Watson No. 378), which flowers a month or more later,
is abundant in the belt of timber to the back of the mounds.
(Watson No. 379.)

HovusTonIA ANGUSTIFOLIA Michx.

Common on the tops and down the sides of the mounds.
This is probably the only locality north of the Missouri
River where this species is found in the State. (Watson
No. 384.)

KUHNIA EUPATORIOIDES L.
Common near the base of the mounds in bare ground.
(Watson No. 389.)

LacinaRia PuNCcTATA (Hook.) Kuntze.

Common on the steep faces on the south and west sides
of the mounds. The other known Missouri stations for
this species are: on a high rocky ridge on the prairie west
of Lee’s Summit in Jackson County, and on a bald knob
in Wright County. L. pycnostachya (Michx.) Kuntze
(Watson No. 388), is common on the bottom prairie close
by the mounds. (Watson No. 387.)

SOLIDAGO speciosa Nutt.
Common on most of the mounds in open ground, but
being a much reduced form. (Watson No. 397.)

SOLIDAGO RUPESTRIS Raf,

Abundant on the tops of the mounds. The plants are
very much reduced, being only about 3 dm. in height.
(Watson No. 398.)

NOTES ON MOUND FLORA OF ATCHISON CO., MO. 129

SoLIDAGO NEMORALIs Ait.

Abundant all over the mounds in bare ground, and also
very much reduced in size like the two last species. ( Wat-
son No. 399.)

SoLrpago RIGIDA L.

Common on the bare open places on most of the mounds.
S. Canadensis L. (Watson No. 395), and S. ulmifolia
Muhl. (Watson No. 396), were common in the belt of
timber at the back of the mounds. (Watson No. 400.)

ASTER OBLONGIFOLIUs Nutt.
Common in bare ground mostly all over the mounds.
(Watson No. 403.)

ASTER SERICEUS Vent.
Abundant on the tops and over the bare parts of the
mounds generally. (Watson No. 404.)

ASTER AZUREUS Lindl.
Common to all the bare parts of the mounds. (Watson
No. 405. )

AstTER Drummonpi Lindl.
Common on the sides of the mounds in the hazel thick-
ets. (Watson No. 406.)

ASTER LAEVIS L.
Common on the open parts of the mounds. (Watson
No. 402.)

Aster Ericorpzs L.
Common on the lower part of the mounds. (Watson
No. 407.)

ASTER AMETHYSTINUS Nutt.
Abundant on nearly all the mounds in bare ground.
Specimens were collected that had simple unbranched
9

130 MISSOURL BOTANICAL GARDEN.

stems one meter in height. Very much resembles A.
multiflorus Ait., and indeed it may yet prove to be an
extreme form of that species, but after a long comparison
with authentic specimens, and a critical examination of all
the specimens collected, Mr. J. G. Smith and I decided
that it must be a form of A. amethystinus. One other
Aster, A. paniculatus Lam. (Watson No. 409), was col-
lected on the bottom-prairie near the mounds. (Watson
No. 401.)

ANTENNARIA PLANTAGINIFOLIA (L.) Rich.
Common on the bare parts of the mounds. (Watson
No. 414.)

SILPH(UM INTEGRIFOLIUM Michx.

Common down the sides of all the mounds. S. lacin-
iatum L. (Watson No. 416), and S. perfoliatum L.
(Watson No. 417), were abundant on the bottom-prairie
below the mounds. (Watson No. 415.)

HE iopsis scaBra Dunal.
Common along the backbones of the mounds. (Watson
No. 418.)

Hevrantaus Maxrmixiani Schrad.

Common down the sides and along the backbones of
most of the mounds. H. grosse-serratus Martens ( Watson
No. 420), which it somewhat resembles, is abundant on the
bottom-prairie below. Since Maximilian collected the
species in Missouri in 1815, it has been found in Wright
County, 1884 (Bush), and in Jackson County at Sheffield,
1892 (Bush No. 413). (Watson No. 419.)

ARTEMISIA GNAPHALODES Nutt.
Abundant around the bases of the mounds, and along the
backbones. (Watson No. 421.)

CacaLia TUBEROSA Nutt.
Common near the base of the mounds. (Watson No. 422).

NOTES ON MOUND FLORA OF ATCHISON CO., MO. 131

Carpuus UNDULATUS Nutt.

On several mounds about half way up their sides.
C. altissimus L. (Watson No. 433), and C. lanceolatus L.
(Watson No. 434), were common on the bottom prairie
below. (Watson No. 429.)

LyGopEsMIA JUNCEA (Pursh) Don.
Abundant all over the mounds, wherever the ground is
bare and free of shrubs. (Watson No. 428.)

LAcTUCA PULCHELLA (Pursh) D. C.

Common near the bases of nearly all the mounds.
Flowers a month earlier than any of our other Lactucas.
(Watson No. 427.)

Lactuca Lupovicrana (Nutt.) D. C.

Common with the last species. Several other species, as
L. scariola L. (Watson No. 424), L. Canadensis L.
(Watson No. 425), and L. sagittaefolia Ell. (Watson
No. 423), were common near the mounds on the bottom-
prairie. (Watson No. 426.)

ASCLEPIAS VERTICILLATA L.

Abundant on all the bare parts of the mounds. A. Sul-
livantii Engelm. (Watson No. 436), and A. Syriaca L.
(Watson No. 437), were common on the bottom-prairie
near the mounds. (Watson No. 435.)

ACERATES VIRIDIFLORA Ives Britton.
Common on all the mounds in bare ground. (Watson
No. 438.)

ACERATES VIRIDIFLORA LINEARIS A. Gray.
Not uncommon with the last. The specific form was
not observed in the county. (Watson No. 439.)

Lapruta Lapruta (L.) Karst.
Common on nearly all the mounds, but most likely intro-

132 MISSOURL BOTANICAL GARDEN.

duced by the cattle which graze over the mounds. (Wat-
son No, 472.)

LITHOSPERMUM CANESCENS (Michx.) Lehm.
Common on the open parts of the mounds. (Watson
No. 474.)

LITHOSPERMUM ANGUSTIFOLIUM Michx.
Common on the tops of all the mounds. (Watson No.
473.)

ONOSMODIUM MOLLE Michx.
Common on the sides and along the backbones of the
mounds. (Watson No. 475.)

PENTSTEMON GRANDIFLORUS Nutt.
Very common on the sides of the taller mounds, but not
observed on the lower ones. (Watson No. 476.)

GERARDIA ASPERA Doug].
Common on the tops and steep sides of the larger
mounds. (Watson No. 479.)

CaSTILLEJA SESSILIFLORA Pursh.
Abundant on the sides of nearly allthe mounds. (Wat-
son No. 481.)

TEUCRIUM OCCIDENTALE A. Gray.

Common down the sides of all the mounds, and spread-
ing out into the bottom-prairie below, where it becomes
abundant. Extends down the Missouri River to Holt
County (Corning No. 482), where it is equally abundant,
and to Jackson County (Courtney No. 485), where it is
uncommon, and it has even been picked up in East Saint
Louis by Eggert, but this last may have been introduced
by means of hay with which it grew when cut. (Watson
No. 483.)

NOTES ON MOUND FLORA OF ATCHISON CO., MO. 133

ScUTELLARIA PARVULA Michx.
Common on the sides of most of the mounds. ( Watson
No. 484.)

EvurHorsiA GLYPTOSPERMA Engelm.
Abundant on the tops of all the taller mounds. (Watson
No. 495.)

EUPHORBIA MARGINATA Pursh.
Abundant along the lower part of all the mounds. Called
Snow-on-the-Mountain by the country people. E. serpens
H. B. K. (Watson No. 497) was abundant on the bottom-
prairie below. (Watson No. 496.)

QUERCUS MACROCARPA OLIVAEFORMIS (Michx. f.) A. Gray.

Common in the pockets on the north sides, and in
thickets at the bases of the mounds. The growth of the
year and the leaves are densely woolly-tomentose, and the
trees are from 3 m. to 6 m. in height. (Watson No. 633.)

SaLIx HUMILIS Marsh.
Abundant in the pockets on the north sides of all the
mounds. (Watson No. 658.)

SisyRincHiuM Bermupranum L.

Common on the tops of nearly all the mounds. This is
what has been called S. angustifolium, and which appears
to me to be very distinct from S. Bermudianum, but I bow
to the decision of the makers of the new Check List.
(Watson No. 671.)

4

Yucca exiauca Nutt.

Common to all the large mounds, on the steep precipitous
sides on the south and west. When observed on June Ist,
1894, it was in full flower, andjthe Pronuba moth was present
in great numbers, and on July 11th, I noticed that there was
an abundance of mature fruits, many of which showed the
insect punctures plainly. It is called Soap-weed by the

—_—— » OS SO ee a SO nanan bt Seer see

134 MISSOURI BOTANICAL GARDEN.

country people, who dig the long saponaceous roots for the
purpose of making soap. (Watson No. 672. )

ANDROPOGON scopaRius Michx.
Common along the bases of the mounds. (Watson No.
773.)

ANDROPOGON PROVINCIALIS Lam.
Common with the last species. (Watson No. 774.)

SPOROBOLUS AIROIDEs Torr.
Abundant mostly all over the mounds, especially on the
steep sides of them. (Watson No. 783.)

Bouretovua oxigostacuya (Nutt. ) Torr.
Common on the tops of all the taller mounds. ( Watson
No. 798. )

BouTELOUA CURTIPENDULA (Michx. ) Torr.
Common on all the mounds in bare ground. (Watson
No. 800.)

BovuTe.ova HirsuTa Lag.
Abundant all over the mounds, and constitutes the prin-
cipal forage for grazing. (Watson No. 799.)

- mie ie 2 Ye

AMONG THE WILLOWS.