

ut 
* - 
7 


THE 


Orl 
1B4rce 
3 | 


BOTANICAL GAZETTE 


EDITORS: 
JOHN MERLE COULTER anp CHARLES REID BARNES, 


* WITH OTHER MEMBERS OF THE BOTANICAL STAFF 
F THE UNIVERSITY OF CHICAGO 


ASSOCIATE EDITORS: 


J. C. ARTHUR, 
Purdue University. 
CASIMIR Beet 
Gen 


I. Be DEG 

University of Padua. 
ADOLF ENGLE 

University of Berlin. 
LEON GUIGNAR 

L’Ec a ‘de Pharmacie. 

RoBErRT A, Ha 

Un veri ani Wisconsin. 


Fritz NOLL 

ousies of Bonn. 
VoLNEY M. SPAL 

Univ versity pig Michigan. 
ROLAND nay 

rvard 4 University. 

nee rot 

Missouri Botanical Garden. 
H. MARSHALL 

Univ ety es "Cambridge. 
EUGEN. WARM 

Un pleas of Copenhagen. 


JINzO — VEIT ise Gary 
Sad Univesity: Tokyo. yal Acad f Sciences, 
“Ssitheha.. 


VOLUME XXxlI 
JANUARY— JUNE, 1901 


WITH TEN PLATES AND SEVENTY-NINE FIGURES IN THE TEXT 


CHICAGO, ILLINOIS 
PUBLISHED BY THE UNIVERSITY OF CHICAGO 
1901 


Mo,Bot.Garacn, _ 
1902. 


BY 


: PRINTED ae 
The University of Chicago Press 


CHICAGO 


SABLE OF CONTENTS: 


.PAG 
New or little known North American trees. II, - - - Charles S. Sargent 


Eriocaulon decangulare L.; An anatomical study (with five 
figur' - - = : - - . - Theo. Holm 
oe ae studies with reference to the hos ase of 
nfungous spores, - B. M. Duggar 
The psig ecology of Chicago and vicinity; a study 
of th n, development, and classification of ie 
socie ate Contribaton from the Hull Bota 


Laboratory. XXIV (with thirty-five figures), pica Chandler Cowles 73, 
mele plants from Guatemala and other Central 
erican republics. XXII (with plate 1), - - John Donnell Smith 
Some recent vighitiGacieis a and the nomenclatorial — “ 
they represent, - - - - - M. L. Fernald 
New or little known North American trees. III, - - Charles S, Sargent 
The ugied of hydrocyanic acid as upon Srent and other 
ds (with six figures), C. O. Townsend 


The tuber-like rootlets of Cycas revoluta. Contributions from 
the Hull Botanical Pere XXVI we ten “a 


ures), A. C. Life 
The O83 age of the forests of northern Michi- 
study in physiographic ecology. Contribu- 
tions feoid the Hull Botanical amen ¢ XXVII 
(with eighteen figures), - - Harry Nichols Whitford 
Mexican fungi. III. - - - - - - - - ££. W. D. Holway 
Ovule and embryo of Potamogeton natans. Contributions 
m the Hull Botanical ee XXVIII (with 
iad lt and 11, and one figure). G. M, Holferty 
A contribution to - life history and cytology of Erythro- 
nium. eae butions from the Botanical Laboratory, 
Ohio State ty: VII (with plates tv-1x) - John H. Schaffner 
Studies on ahi plants. I (with plate x) esi Hi. M. Halt 
Contributions from the Rocky Mountain Herbarium, II. - Aven Nelson 


Studies on the geotropism of stems. II (with three figures), Edwin B. Copeland 
BRIEFER ARTICLES — 
Nitrates as a ce of nit f hytic fungi Mary H. Smith 
Non-sexual propagation in Cpcata: a etic eee - Carleton £. Preston 
Vv 


I 


265 


289 
326 


339 


vi CONTENTS [ VOLUME XXXI 


Observations upon the feeding SRT of Fuligo pees 

septica (with one figure), —- - John W. Harshberger 198 
Swarm-spore formation in Hydrodictyon utriculatum 

Roth, - - - - - * = - - H.G. Timberlake 203 

Walnut bacteriosis, - - - = - - - Newton B. Pierce 272 

Fern variation in Great Britain, - - - - Charles T. Druery 347 

Notes of travel. IV, - - - - - David G. Fairchild 352 

Notes of travel. VI. - - - - - - D. G. Fairchild 423 

The cardinal principles of morphology’ - - - W. F. Ganong 426 

CURRENT LITERATURE — - . - - 67, 129, 204, 274, 355, 435 


For titles see index under author’s name and Re- 
. Papers noticed in “ Notes for Students” are 
ae under author’s name and subjects 


Wa a ee ee ee yo eee eee oe ee 


DATES OF PUBLICATION, 


No. 1, January 21; No. 2, February 23; No. 3, March 16; No. 4, April 15; No. 
5, May 18; No. 6, June 20 


ERRATA. 
P. 2, line 2 from above, for N. S. Plank read E. N. Plank. 
P. 5, line 7 from above, for W. N. Canby read W. M. Canby. 
P. 10, line 10 from above, p. 11, lines 17 and 21 from above, for Brainard read 
Brainerd. 
P. 14, line 13 from below, for nearly read rarely. 
P. 132, line 9 from above, for Schuman read Schumann. 
P. 203, footnote 4, line 5, for Mazijck read Mazyck. 
ve P. 225, line 1, for Crategus Texana read CRATAEGUS TEXANA. 
Pi274, footnote I, line 1, for plant read vegetable. 
P. 328, line 16 from above, for G. read Galphimia. 
P. 328, line 9 from below, for S. read Solanum. 
P. 329, line 6 from above, for Aecidium read aecidium. 
P. 333, line 5 from above, for V. read Viguiera. 
P. 343, footnote 3, line 2, for 1888 read 1898. 
P. 356, line 8, dele American. 
UP; 356, line § from sami ae ene heterospory in 


est 356, line A 1 from b: 


Vol. XXXI JANUARY, 


1901 No. | 


BOTANICAL GAZETTE 


JOHN M. COULTER AND CHARLES R.. BARNES, 


WITH OTHER MEMBERS OF THE pial STAFF 
OF THE UNIVERSITY OF CHICA 


ASSOCIATE EDITORS 


j. & ARTHUR 
Purdue University 


CASIMIR tect DOLLE 
Gen 


J. B. DeTO 
nies of Padua 
ADOLF ENGL 
Un cou of Berlin 
LEON GUIGNARD 
L’ Ecole de Pharmacie, Paris 
ROBERT “AHA 
Oni ice ” ‘isconrte 
JINZO an UR 
Kia naar Takyé 


CHICAGO, I HLLINOIS 


FRITZ NOLL 

University: “é Bonn 
VOLNEY M. SPALDIN 

University oS Michigan 
ROLAND THAXTER 

Harvard University 
WILLIAM TRELEASE 

Missouri Botanical Garden 
H. pias tis WA 

University a Cambridge 
ee WARMING 

University of Copenhagen 


VEE WIT (eae ne 


‘oyal —— s Sciences 


eenenrqaner tiers 
A 


sete enn 
f 4 


ooo 


ener rire 


‘3 _ number following. — 
as All remittances 


Botanical Gazette 


A Monthly Fournal Embracing all Departments of Botanical Science 
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Vol. XXXI, No. 1 Issued January 23, 1901 


CONTENTS 


NEW OR LITTLE KNOWN NORTH AMERICAN TREES. II. Charles S. Sargent - I 
ee DECANGULARE Bo AN ANATOMICAL STUDY ieee FIVE piopiciueer 
. Holm 


17 

Re ae eo STUDIES WITH REFERENCE TO THE GERMINATION OF CER- 

FUNGOUS SPORES. &. M. Duggar 
CURRENT LITERATURE. 
“ BOOK REVIEWS - - - - - “ > i é 67 
AGRICULTURAL BOTANY. A BOTANICAL DICTIONARY. 

MINOR NOTICES . - - - - - 3 a zs rt 69 
NOTES FOR STUDENTS - - - - - 5 Fs . ‘ 71 
NEWS > - - . - - - “ = ‘ a is 72 


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VOLUME XxXxI NUMBER 1 


Bet ANICAL” CGNZETTE 
JANUARY, rgo1r 


NEW OR LITTLE KNOWN NORTH AMERICAN 
TREES. ft. 
CHARLES S. SARGENT. 

Gleditsia Texana, n. sp.— Unarmed. Leaves puberulous, or 
tomentose toward the base of the slender rachis, ultimately becom- 
ing nearly glabrous, 12 to 22-foliolate; leaflets oblong-ovate, 
often somewhat falcate, rounded or acute, and apiculate at the 
apex, obliquely rounded at the base, obscurely crenulate-serrate, 
short-petiolulate, thick and firm, dark green and lustrous on the 
upper surface, pale on the lower surface, % to 1 in. long. 
Staminate flowers orange-yellow, in slender racemes 3 to 4 in. 
long; calyx-lobes ovate, acute, villous, rather shorter than the 
petals; stamens exserted. Pistillate flowers unknown. Legume 
straight, compressed, without pulp, rounded or short-pointed at 
the apex, abruptly rounded at the broad base, thin-walled, dark 
chestnut-brown, puberulous, only slighly thickened on the 
margins, many-seeded, 4 to 5 in. long and 1 in. wide. Seeds 
oval, compressed, dark chestnut-brown and lustrous, nearly % in, 
long. 

A tree 100 to 120 feet in height, with a trunk rarely more 
than 2% feet in diameter covered with pale smooth bark, and 
upright slightly spreading branches. Flowers at the end of 
April. Fruit ripens and falls late in the autumn. 

High rather dry bottom-lands of the Brazos river near 
Brazoria, Texas, in dense woods composed principally of Gie- 
ditsia triacanthos, Platanus occidentalis, and Populus deltoidea. 

I 


2 BOTANICAL GAZETTE [JANUARY 


First made known from legumes picked up from the ground 
several years ago by Mr. N. S. Plank, and later by collections 
made by Mr. B. F. Bush in 1899 and Igoo. 


Gleditsia Texana is an interesting addition to the North American 
silva. From Gleditsia triacanthos, which it resembles in foliage and in the 
staminate flowers, it is distinguished by its spineless branches and smoother 
pale bark. From all other species of the genus it differs in the legume, 
This resembles those of the many-seeded species, of which G/editsta triacanthos 
is the type, in its general form and color, and in its numerous seeds. It differs 
from them in its much smaller size, thin compressed walls with thinner mar- 
gins, and in the absence of the sweet pulp which surrounds their thicker 
lighter-colored seeds. It differs in form from the compressed pulpless 
legume of Gleditsia aquatica, and in its more numerous seeds. Like the 
legume of that species, the legume is frequently seedless. 

nown only in a single grove in a comparatively small number of individ- 
uals, and sharing something of the character of each of the other American 
species which both grow near it, the hypothesis of a natural hybrid between 
those species might be considered, were it not forthe fact that the fruit of Gledz¢- 
sta triacanthos is nearly half-grown in this region before the flowers of G/e- 
ditsta Texana open, while the flowers of G/editsta aquatica do not open until 
ten or twelve days after those of Gleditsta Texana have fallen. 


Crataegus Engelmanni, n. sp.— Leaves broadly obovate or 
rarely elliptical, rounded and often short-pointed at the apex: 
gradually narrowed below into short glandular pilose petioles, 
coarsely glandular-serrate with incurved teeth usually only above 
the middle and generally only at the apex, coriaceous, dark 
green, lustrous and roughened on the upper surface with short 
rigid pale hairs, pale on the lower surface, pilose above and 
below on the slender midribs and on the thin obscure primary 
veins and veinlets, 1 to 1% in. long and ¥% to 1 in. broad; sti- 
pules linear-lanceolate, light red, % in. long, caducous. Flowers 
34 in. in diameter, in 8 to 12-flowered wide slender-branched 
cymes thickly coated with long pale hairs ; bracts linear-lanceo- 
late, tomentose or villous, often % in. long; calyx tomentose, 
villous or nearly glabrous, the lobes narrow, acuminate, entire, 
glabrous on the outer surface and usually more or less pubes- 
cent on the inner surface, reflexed after anthesis, often deciduous 
before the ripening of the fruit; stamens ten; filaments slender ; 


Le i SNP n pushed act cee a eee 


+ SSDI aas op SE ROL ay) oe PE 


i itaicerian: 


1901 } NORTH AMERICAN TREES 3 


anthers small, rose-color; styles two or three. Fruit globose, 
about ¥% in. in diameter, bright orange-red with a yellow cheek 
and thin dry green flesh; tube of the calyx prominent, the cavity 
broad in proportion to the size of the fruit, shallow; nutlets two 
or three, thick, prominently ribbed on the back with high rounded 
ridges, 4% in. long. 

A tree from 15 to 20 feet in height with a trunk 5 to 6 inches 
in diameter covered with dark brown scaly bark, wide-spreading 
usually horizontal branches forming a low flat-topped or a rounded 
head, and slightly zigzag branchlets marked with large scattered 
white lenticels, at first clothed with pale hairs, becoming nearly 
glabrous and reddish-brown during the first season and lighter- 
colored and gray, or gray tinged with red during their second 
year, and armed with remote slender straight or slightly curved 
chestnut-brown spines 1% to 2% in. long. Flowers from the 
end of April (Augusta, Ga.) to the end of May (Missouri). 
Fruit ripens early in November. 

Dry hillsides and slopes, often on limestone; less frequently 
along the low margins of meadows and roadsides. Kimmswick, 
Missouri, Dr. George Engelmann, May 1860; Pacific, Missouri, 
George W. Letterman, 1881; southern Missouri, &. 7. Bush (nos. 
86 and 19), 1898, 1899; West Nashville, Tennessee, 7. G. 
Harbison, May 1899; Tuscaloosa, Alabama, 7. G. Harbison, 
May 1899; Gunterville, Alabama, 7. G. Harbison, June 1899; 
Rome, Georgia, C. Boynton, May 4, 1899; Birmingham, Alabama, 
C. Boynton, January and July 1899; Washington Road near 
Augusta, A. Cuthbert and C. S. Sargent, April 1900. 

Well distinguished from Crataegus Crus-gallz, with which for many years 
it has been confounded, by its smaller thinner leaves roughened above by 
rigid hairs and pilose on the lower surface, more prominent primary veins and! 
villous or tomentose branchlets and cymes, Crataegus Crus-gaédi in all its. 
forms, as I now understand it, being perfectly glabrous, with veins which, 
except in the case of leaves on the most vigorous shoots, are usually almost: 
entirely within the parenchyma; by its smaller brighter red and yellow fruit ; 
and by its less numerous and more slender spines. 

Crataegus Canbyi, n.sp.—Glabrous. Leaves oblong-oval to 
ovate or r rarely obovate, pointed or occasionally rounded at the 


4 BOTANICAL GAZETTE [JANUARY 


apex, cuneate and entire at the base, coarsely and doubly ser- 
rate above the middle with glandular incurved teeth, often more 
or less deeply lobed on vigorous leading shoots with broad acute 
lobes, coriaceous, dark green and lustrous above, pale and dull 
below, 2 to 2% in. long and 1 to 1% in. wide, or on leading 
shoots often 3 to 4 in. long and 2 in. broad, the thick midribs 
and four or five pairs of remote primary veins raised and con- 
spicuous on the lower surface and impressed on the upper sur- 
face; petioles stout, grooved and glandular on the upper side 
with scattered dark red persistent glands, more or less winged 
above by the decurrent base of the leaf-blades, red below the 
middle, % to % in. long; stipules oblong-obovate to linear-lan- 
ceolate, glandular-serrate, 1% in. long. Flowers 5 in. in diameter, 
in loose many-flowered long-branched compound cymes; calyx- 
lobes narrow, acuminate, serrate with slender teeth tipped with 
small red glands, nearly entire, reflexed and closely appressed 
on the fruit, often deciduous before maturity; stamens ten or 
rarely twelve or thirteen; filaments slender, elongated; anthers 
small, rose-color; styles threeto five. Fruits oblong to subglo- 
bose, full and rounded at the ends, depressed at the insertion of 
the stalks, flesh red, thick, juicy, and succulent, % to 5 in. long, 
drooping on the slender elongated peduncles, lustrous dark 
crimson, punctate with occasional large pale dots; calyx cavity 
deep and narrow; nutlets three to five, 4 in. long, light chestnut- 
brown, prominently ridged on the back with broad rounded ridges. 

A bushy much-branched tree 20 to 25 feet in height with a 
short stout trunk often a foot in diameter covered with dark 
brown scaly bark; branches stout, erect, wide-spreading, at 
first dark green and marked with numerous pale raised lenticels, 
slightly zigzag, soon becoming light orange-brown and lustrous, 
gray during their second year, and armed with few stout chest- 
nut brown spines varying from 34 to 1% in. in length. Flowers 
the middle of May; the fruit ripens early in October and falls 
in early winter. 

Hedges and thickets near Wilmington, Delaware; Pennsyl- 
vania, meadows of Tohickon Creek, Quakertown, C. D. Frets, 


1901] NORTH AMERICAN TREES 5 


1899 and 1900; Tinicum island, Delaware county, B. H. Smith, 
1900. 

To be distinguished from the related Crataegus Crus-galii by its oblong 
usually pointed leaves with veins raised and prominent on the lower surface, 
by its short infrequent spines, by the habitual appearance of the flowers ten 
days before those of that species open in the same region, and by the red 
juicy flesh of the fruit. First noticed in October 1898, by Mr. W. N. Canby 
of Wilmington, Delaware, who permits the association of his name with this 
handsome tree. 

Crataegus Peoriensis, n. sp.—Leaves obovate, short-pointed or 
occasionally rounded and sometimes slightly irregularly lobed at 
the broad apex, gradually narrowed below, sharply and often 
doubly glandular-serrate usually only above the middle, glab- 
rous with the exception of a few scattered pale hairs néar the 
base of the upper surface of the midribs, thick and firm, with 
four or five pairs of thin primary veins raised and conspicuous 
below, deeply impressed above and running very obliquely from 
the slender midrib to the end of the leaf, dark green and very 
lustrous on the upper surface, pale on the lower surface, bright 
bronze color as they unfold, % to 2 in. long, 3% to I in. wide, 
and on vigorous leading shoots frequently a third larger and 
occasionally laterally lobed; petioles broad, deeply grooved, 
more or less winged and slightly glandular above the middle, 
and covered early in the season with short white scattered hairs ; 
stipules glandular, linear-lanceolate, or on vigorous shoots 
lunate and ¥% to I in. long. Flowers cup-shaped, % in. in 
diameter when fully expanded, in broad loose slender-stemmed 
many-flowered corymbs, their bracts and bractlets linear, conspic- 
uously glandular, caducous; calyx glabrous, the cup deep and 
narrow; calyx-lobes narrow, acuminate, entire or irregularly 
glandular-serrate with minute stalked dark red glands, pubescent 
below the middle on the upper surface, spreading nearly at right 
angles to the cup when the flowers open, persistent on the fruit ; 
stamens ten; filaments elongated, slender; anthers small, rose- 
color; styles usually two or three. Fruit oblong to obovate, 
full and rounded at the ends, slightly depressed at the insertion 
of the peduncle, bright scarlet, % to 34 in. long, with thick 


6 BOTANICAL GAZETTE [JANUARY 


nearly white firm flesh; nutlets two, or rarely three, thick, 
prominently ridged, about ¥% in. long. 

A tree 20 or 25 feet in height with a trunk sometimes a 
foot in diameter covered with dark brown scaly bark, stout 
spreading branches forming a broad flat-topped head, and 
slightly zigzag, glabrous branchlets green when they first 
appear, orange-brown and lustrous during their first season, 
growing lighter during their second year, and armed with 
straight or slightly curved very slender dull chestnut-brown 
spines 2 to 24% in. long. Flowers from the middle to the end 
of May. Fruit ripens early in October but when gathered 
retains its shape and color until December. 

In open woods along the moist borders of streams or depres- 
sions in the prairie and on hillsides in clay soil. Short and 
Peoria counties, Illinois, where it was discovered by Mr. Vir- 
ginius H. Chase of Wady Petra, Illinois, September 1897 (nos. 
48, 446, 449, 481, 485). 

Crataegus pratensis, n. sp.—Leaves obovate-oblong, pointed 
or rounded at the apex, gradually tapering below and decurrent 
nearly to the base of the short slender petioles, sharply and 
often doubly serrate usually only above the middle with teeth 
tipped with minute dark red caducous glands, more or less 
deeply divided above into short broad acute lobes, with slender 
midribs and narrow oblique primary veins deeply impressed on 
the upper side and raised and prominent below, bright bronze- 
yellow and coated on both surfaces with short pale hairs when 
unfolding; at maturity glabrous, thick and firm, dark green and 
lustrous on the upper surface, pale on the lower surface, % to 2 
in. long, I to 1% in. broad, and on vigorous shoots often 3 in. 
long and 2% in. broad; stipules lunate, stalked, coarsely 
_ glandular-serrate, on vigorous shoots often 1 in. in length. 
Flowers % in. in diameter, in broad loose thin-branched many- 
flowered compound pubescent ultimately glabrous cymes; calyx 
coated on the outer surface with matted white hairs, the cavity 
deep and narrow; calyx-lobes acute, coarsely glandular-serrate, 
spreading or more or less reflexed at anthesis, often deciduous 


ciate wpe eee ees a a a es ee ae ae Se ee eee 


a EE ne 


t 


1901] : NORTH AMERICAN TREES 7 


from the fruit ; stamens ten; filaments slender, elongated ; anthers 
small, rose color; styles two or three. Fruit globose, pendent on 
the elongated peduncles, bright scarlet, ¥ in. in diameter, with 
thin dry yellow flesh; nutlets usually two or three, broad and 
thick, full and ridged on the back, about ¥ in. long. 

‘A tree occasionally 20 feet in height, with a trunk 3 to 7 in. in 
diameter covered with dark brown scaly bark and frequently 
armed with long slender much-branched ashy gray spines, wide- 
spreading branches forming a broad round-topped head, slender 
glabrous branchlets marked with white lenticels, at first green, 
becoming light chestnut-brown and lustrous during their first 
summer, and furnished with slender straight or slightly recurved 
chestnut-brown spines 2 to 2% in. long. Flowers from the 
middle to the end of May. Fruit ripens the first of October and 
falls in November. 

Open woods near the banks of small streams in the prairie 
region of Stark and Peoria counties, Illinois, where it was dis- 
covered by Mr. Virginius H. Chase in May 1898. Rare and 
local. Trees which appear to be of the same species were found 
by Mr. B. F. Bush near Swan, Taney county, Missouri, in Octo- 
ber 1889; and later by Professor William Trelease and myself. 

Crataegus submollis, n. sp. (Crataegus tomentosa Emerson, 
Trees Mass. 435. 1846 [not Linnaeus]; ed. 2, 494, p/.—. 1875.— 
Crataegus coccinea mollis Brunet, Cat. Vég. Lig. Can. 25. 1867 
[in part, not Torrey & Gray]; Watson & Coulter, Gray’s Man. 
ed. 6. 165. 1890 [in part ].—Crataegus subvillosa Macoun, Cat. 
Can. Pl. 1:147. 1888 [in part, not Torrey |.— Crataegus mollis 
Sargent, Silva N. Am. 4:99. p/. 782. 1892 [in part, not Scheele}; 
Koehne, Herb. Dendr. 232.— Crataegus coccinea subvillosa Lange, 
Rev. Spec. Gen. Crataegi 31. fig. —).—Leaves ovate, acute, 
cuneate at the base, sharply serrate with gland-tipped teeth, 
slightly divided above the middle into three or four pairs of 
acute lobes, membranaceous, pale yellow-green and roughened 
on the upper surface with short closely appressed rigid hairs, 
paler and at first coated below with dense hoary tomentum, at 
maturity puberulous only on the prominent midribs and remote 


8 BOTANICAL GAZETTE [JANUARY 


slightly raised veins, 2% to 4 in. long and 2 to 2% in. wide; 
petioles slender, slightly grooved and glandular on the upper 
side, winged above by the decurrent bases of the leaves, tomen- 
tose, often tinged with red below the middle and I to 2 in. long; 
stipules linear-lanceolate or lunate on vigorous shoots, tomen- 
tose, glandular-serrate, 1% to 34 in. long, caducous. Flowers I 
in. in diameter, in broad many-flowered compound tomentose 
cymes, their bracts and bractlets glandular-serrate with dark 
glands; calyx tomentose, its cup deep and broad; calyx-lobes 
acute from a broad base, tomentose, glandular with small stalked 
persistent red glands, usually wide-spreading at anthesis and 
persistent, erect and much enlarged on the fruit; stamens ten; 
filaments slender ; anthers small, pale yellow; styles three to five. 
Fruit pear-shaped with yellow subacid dry flesh, about 34 in. 
long, gracefully drooping on the slender elongated puberulous 
branches of the cymes, bright orange-red, lustrous, marked with 
occasional pale lenticels, puberulous toward the base; nutlets 
usually five, rounded and slightly ridged on the back, a third of 
an inch long. Flowers from the 20th to the end of May. Fruit 
ripens and falls from the first to the middle of September. 

A tree 25 to 30 feet in height with a trunk occasionally a 
foot in diameter covered with light brown scaly bark, and wide- 
spreading branches, or often a tall intricately branched shrub, 
and with branchlets at first green and coated with hoary tomen- 
tum, becoming before midsummer dark orange-brown, and 
during their second year lustrous and pale gray-green to orange- 
brown, glabrous and very lustrous, slightly zigzag and armed 
with slender bright chestnut-brown straight or more or less 
incurved spines 2% to 3 in. long. 

On rich hillsides, along the margins of roads, and sometimes 
in low moist soil, from the neighborhood of Montreal,_/. G. Jack, 
1899, to Orono and Dover, Maine, M@. L. Fernald, May 1887, 
and Aguust 1896; Gerrish island, Maine, J. G. Jack, September 
1899; and to Jamaica Plain and Milton, Massachusetts. 

Confounded by me in the fourth volume of The Silva of North America, 
where it is well figured, with Crataegus mollis of Scheele, a common tree 


1901 } NORTH AMERICAN TREES 9 


from Michigan to Nebraska and Missouri. Crataegus submodé/is differs from 
that species in its smaller and less deeply lobed cuneate leaves, which are 
not truncate or cordate even on vigorous leading shoots; in its ten, not 
twenty, stamens ; smaller, less downy fruit on much longer, more slender and 
drooping peduncles ; in its more zigzag orange-colored branchlets, those of 
Crataegus mollis being of a bright chestnut-brown color during the first sum- 
mer ; longer and much more numerous spines ; and in its smaller size. 

Crataegus dilatata, n. sp.— Leaves ovate from a broad, trun- 
cate, cordate, or slightly rounded base, coarsely irregular glan- 
dular-serrate, unequally lobed usually with two or three pairs of 
acute lateral lobes, membranaceous, with slender midribs and 
four or five pairs of thin primary veins, when the flowers open 
roughened on the upper surface with short ridged white hairs, 
light yellow-green, and conspicuously plicate, at maturity dark 
green, 2 to 2% in. long, usually as wide as long, and on vigor- 
ous leading shoots often 4 to 4% in. long and usually rather 
broader than long; petioles slender, terete, glandular, at first 
more or less villous but soon glabrous, 1 to 1% in. long, dark 
red toward the base; stipules linear-lanceolate and on vigorous 
shoots lunate and foliaceous, glandular with dark red glands, 
% in. long, caducous, Flowers 1 to 1% in. in diameter when 
expanded, in broad, open, nearly glabrous, villous or tomentose 
compound cymes, the bracts and bractlets, like the bud-scales, 
glandular with stalked red glands; calyx villous with soft white 
deciduous hairs, the cup broad and shallow; calyx-lobes broad, 
acuminate, coarsely glandular-serrate, much enlarged and spread- 
ing on the fruit, and bright red on the upper surface below the 
middle ; stamens 20; filaments slender, elongated ; anthers large, 
rose color; styles usually five, surrounded at the base by small 
tufts of white hairs. Fruit subglobose with sweet, yellow, thick 
flesh, 34 in. in diameter, bright scarlet, punctuate with minute 
dark spots; nutlets five, comparatively small, prominently 
ridged on the back, about ¥% in. long. 

A tree sometimes 20 feet in height, with a tall straight 
trunk, wide-spreading branches forming a broad round head, or 
bushy with numerous spreading stems and slender, glabrous, 
slightly zigzag branchlets marked with large pale lenticels, at first 


10 BOTANICAL GAZETTE | JANUARY 


dark green tinged with red, becoming light chestnut-brown and 
very lustrous during their first season, dark dull gray in their 
second year, and armed with stout straight chestnut-brown 
spines 1 to 2 in. long, or sometimes nearly spineless. Flowers 
about May 20. Fruit ripens and falls early in September. 

Rich hillsides and borders of salt marshes. Revere, Massa- 
chusetts, C. &. Faxon, May 1884; Somerset, Massachusetts, 
Miss L. H. Handy, May 1899; Ipswich, Massachusetts, John 
Robinson, May 1900; Tiverton, Rhode Island, C. S. Sargent, 
August 1900; Middlebury, Vermont, Ezra Brainard, 1900. 

It is possible that this is one of the plants cultivated in Europe as Cra- 
taegus coccinea, but | have not been able to find any trace of it as a garden 
plant. 

Crataecus Hormesiana Ashe, Jour. Elisha Mitchell Sci. Soc. 
167: 78. 1900.— Leaves ovate, pointed, rounded, cuneate or on 
‘vigorous leading shoots sometimes truncate or cordate at the 
base, sharply and often irregularly doubly serrate with sharp 
teeth tipped at first with prominent dark red caducous glands, 
usually slightly lobed with from three to five pairs of acute 
lateral lobes, membranaceous, pale yellow-green above, pale 
and glabrous below, scabrous on the upper surface with short 
rigid white hairs when the flowers open, at maturity becoming 
thick, firm, and nearly smooth, 2 to 3 in. long and 2 to 2% in. 
wide, with prominent midribs and five or six pairs of stout pri- 
mary veins deeply impressed above; petioles slender, terete, 
slightly grooved and glandular on the upper surface with per- 
sistent dark glands, and 1 to 1% in. long; stipules lunate, 
glandular-serrate, small, caducous. Flowers cup-shaped, 3% in. 
in diameter when fully expanded, in loose many-flowered com- 
pound glabrous cymes on slender elongated pedicels; calyx 
‘narrow, glabrous, tinged with red, its cup ‘broad and shallow; 
calyx-lobes acute, glandular-serrate or rarely nearly entire, 
persistent and erect on the fruit; stamens five or occasion- 
ally six, seven, or eight; filaments stout, spreading; anthers 
large, dark purple-red; styles usually three. Fruit pendulous, 
oblong, crowned with the erect conspicuous calyx-lobes, crimson, 


PLS Tr ar Tee Ree PRG ele a PSSM Bee ce are ee age meee cee Nee Tolar Oey Sy teem Cera ae 


a ee ae ee a ee ee ee MEE ee ee ee ee ee RE Le RS Nay Rec eee Oe SET a TSR ean Nemes See ee pe RTE pe ee Se ne SaaS eee eee ee | ena at a eT aces Ra Mn ee a ea Sin 
eae aera ets is = sas ae - i asl rei ess i i oa ™ . ene vs e carrie 
2 


igor | NORTH AMERICAN TREES It 


lustrous, % to 34 in. long, with thin dry yellow flesh of disagree- 
able flavor; nutlets prominently ribbed, light chestnut-brown, 
about % in. long. 

A tree occasionally 30 feet in height with a well developed 
trunk 12 to 15 inches in diameter, covered with pale or nearly 
white scaly bark, stout upright branches forming an open irregu- 
lar or more rarely a broad compact head, stout chestnut-brown 
branchlets and straight usually infrequent spines 1% to 2 in. 
long. Flowers from the 5th to the middle of May. Fruit 
ripens and falls early in September. 

From the Province of Quebec, /. G. Jack, and Ontario, D. W. 
Beadle, to Maine, Gardner, Alice M. Richards, Ellsworth, Mrs. 
M. A. Clarke, North Berwick, Mrs. John Parlin, Acworth, R. 
Hoffman; New Hampshire, Holderness, C. Z. and W. Faxon, 
Haverhill, #. Faxon, Troy, Rand and Robinson; Vermont, Char- 
lotte, 7. H. Horsford, Ferrisburg and West Addison, Ezra 
Lrainard, Bennington, Rutland, North Pownal, and Fairhaven, 
W. W. Eggleston, Rutland, C. S. Sargent; Massachusetts, Berlin 
and West Boylston, J. G. Jack, South Lancaster, C. S. Sargent, 
Lanesboro, C. £. Faxon; New York, “‘N. N.Y.” in Herb. Gray, 
without date, Fort Crown Point, Ezra Brainard, Rochester, C. C. 
Laney ; Rhode Island, Tiverton, Miss Alice Sargent; and Pennsyl- 
vania, Sellersville, Dr. C. D. Fretz. 

Crataegus Holmesiana inhabits rich moist hillsides and the borders of 
streams and swamps, and is always easily recognized by the pale bark of the 
stem, the distinctly yellow color of the leaves, the small cup-shaped flowers 
with few stamens, and by the large oblong early fruit. It is very abundant 
in the upland pastures of Worcester county, Massachusetts, and along the 
banks of Branch creek at Sellersville, Pennsylvania. It is one of the largest 
species of New England. I have been unable to discover that this handsome 
tree, which has probably always been confounded with Crataegus coccinea, 
has ever been an inhabitant of gardens, unless the figure of Crataegus coc- 
cimea, published in 1835 in the Botanical Magazine ( pl. 7434) was intended 
to represent this species. 

CRATAEGUS COCCINEA Linnaeus.—The name Crataegus coccinea 
was first used by Linnaeus in the first edition of Species Planta- 
rum 1:476. 1753. His description of this species, ‘Crataegus 


2 BOTANICAL GAZETTE [JANUARY® 


’ 


foliis ovatis repando-angulatis serratis glabris,” had, however, 


appeared in 1737 in Hortus Cliffortianus. In both works a species: 
of Plukenet (Phyt. Bot. fl. 46. f. g) and a species of Miller (Cat.. 


Pl. Hort. Angl. p/. 737. f. 7) were referred by Linnaeus to his 
Crataegus coccinea. Plukenet’s plant is preserved in the British 


Museum. It belongs to the mollis group, but the specimen is so- 


meager that I have been unable to identify it. Miller’s figure 
perhaps represents a species of the mo/lis group, but it is certainly 
not the same plant as the one figured by Plukenet, and I cannot 


identify it. The only representative of Crataegus coccinea in Lin- 


naeus’s herbarium, and so labeled by him, is an entirely different 


plant from either of those represented in Plukenet’s or Miller’s. 
figures which Linnaeus had referred to his species. Morever,. 


the specimen is not glabrous but villous on the leaves, corymb,. 
and young branches, and the leaves can hardly be described as 


‘‘ repando-angulatis serratis.” The Linnaean specimen is not dated,. 


and it is therefore possible that it was not from this specimen 
but from Plunkenet’s or Miller’s figure that Linnaeus drew his des- 
cription of Crataegus coccinea. There seems in this case, there- 
fore, but one of two methods to follow in considering this name; 
either the specimen in Linnaeus’s herbarium must be ignored, 
and the name dropped entirely because it was given to a species 
founded on two distinct plants, neither of which can be satisfac- 
torily determined; or the specimen in the Linnaean herbarium 
labeled Crataegus coccinea by Linnaeus himself must be accepted 
as the type of this species. In view of the fact that the name 
Crataegus coccinea is one of the best known of the names which 
have been applied to American species of the genus, and as the 
plant labeled Crataegus coccinea by Linnaeus is now known to be 
a common and widely distributed species in the north Atlantic 
coast region, it is perhaps best to consider the specimen in the 
Linnaean Herbarium as the type of Crataegus coccinea, which can 
be described as follows: 

Crataegus coccinea Linnaeus.—Leaves elliptical or on vigor- 
ous shoots mostly semiorbicular, acute or acuminate, divided 
above the middle into numerous acute coarsely glandular-serrate 


w90r] NORTH AMERICAN TREES 13 


lobes, cuneate and finely glandular-serrate below the middle and 
often quite entire toward the base, with slender midribs and 
remote primary veins arcuate and running to the points of the 
lobes, at the flowering time membranaceous, coated on the 
upper surface and along the upper surface of the midribs and 
veins with short soft white hairs, at maturity thick, coriaceous, 
dark green and lustrous on the upper surface, paler on the lower 
surface, glabrous or nearly so, 1% to 2 in. long and 1 to 1% in. 
wide, with slender glandular petioles 34 to 1 in. long, slightly 
‘grooved on the upper surface, often dark red toward the base, 
and like the young branchlets villous with pale soft hairs; 
stipules lanceolate to oblanceolate, conspicuously glandular ser- 
wate with dark red glands, % to 34 in. long. Flowers % to 3% 
in. in diameter when fully expanded, in broad many-flowered 
‘compound tomentose cymes; bracts and bractlets linear-lanceo- 
late, coarsely glandular-serrate, caducous; calyx tomentose, the 
lobes lanceolate, glandular-serrate, nearly glabrous or tomen- 
tose, persistent, wide-spreading or erect on the fruit, dark red 
above at the base; stamens ten; anthers yellow; styles three or 
four. Fruit subglobose, occasionally rather longer than broad, 
dark crimson, marked with scattered dark dots, about ¥% in. in 
diameter, with thin sweet dry yellow flesh; nutlets three or four, 
about ¥% in. long, conspicuously ridged on the back with high 
grooved ridges. 

A low bushy tree occasionally 20 feet in height with a short 
trunk 8 to Io in. in diameter, or more frequently shrubby and 
forming wide dense thickets, and with stout more or less zigzag 
branches bright chestnut-brown and lustrous during their first 
year, ashy gray during their second season and armed with 
many stout chestnut-brown straight or curved spines I to 1% in. 
long. Flowers late in May. Fruit ripens and falls toward the 
end of October usually after the leaves. 

Slopes of hills and the high banks of salt marshes usually in 
rich well-drained soil, Essex county, Massachusetts, John Robin- 
son, 1900; Gerrish island, Maine, /. G. Jack, 1899-1900; Bruns- 
wick, Maine, Mrs. Kate Furbish, May 1899; Newfoundland, A. 
C. Waghorne, 1894. - 


14 BOTANICAL GAZETTE [JANUARY 


CRATAEGUS COCCINEA rotundifolia. —With Crataegus coccinea as 
described above there often grow in the same thickets plants which 
differ from it only in the less development of the hairs on the 
leaves, young branches, and corymbs. Some of these plants are 
entirely glabrous with the exception of a few short hairs on the 
upper surface of the young leaves, while others show all degrees 
of variation in the developement of their villous covering. The 
synonymy of this form, which cannot be considered more than a 
variety, is as I understand it as follows: 

Crataegus rotundifolia Moench, Baume Weiss. 29. A/. 7. 1785. 

Mespilus glandulosa Ebrhart, Beitr. 3:20. 1788. 

Crataegus glandulusa Aiton, Hort. Kew. 2: 168. 1789. 

Crataegus horrida Medicus, Gesch. Bot. 1793. 

Mespilus rotundifolia Du Roi, Harbk. Baumz, 2: 607. 1795. 

Crataegus coccinea Lindley, Bot. Reg. 23: ~/. 7957. 1837 (not Linnaeus). 

Crataegus coccinea macracantha Sargent, Silva N. Am. 4:96. 1892, in 
part, not Lindley. 

Crataegus coccinea rotundifolia is one of the commonest New England 
forms, ranging southward to eastern Pennsylvania, Easton, 7. C. Porter, 
1894, Stroudsburg, W. M7. Canby, 1900, and Delaware, Fairhurst, W. M7. 
Canby, 1900; and westward to the region of the Great Lakes. Its northern 
and western range, however, is still imperfectly known, as there are evidently 
some distinct forms of this group which are still confounded with Crataegus 
coccinea and this variety. All the species of the group, which has been 
curiously overlooked by American botanists, have thick coriaceous dark 
green and lustrous mature leaves, flowers with ten or nearly twenty stamens 
and pale yellow anthers, and globose or subglobose scarlet fruit of medium 
size with three or four nutlets. 

Crataegus Jonesae, n. sp. ( Crataegus coccinea macracantha Rand 
& Redfield, Fl. Mt. Desert Island 98. 1894, not Dudley ).— 
Leaves elliptical, pointed, cuneate and decurrent at the base, 
sharply and doubly serrate and usually lobed above the middle 
with numerous small acute lobes, coriaceous, dark green and 
lustrous above, pale and puberulous below, especially on the 
stout midribs and broad remote primary veins, deeply impressed 
above, 3 to 4 in. long and 2 to 3 in. broad, at first coated above 
with soft pale caducous hairs and glandular with small dark red 
deciduous glands on the teeth; petioles stout, more or less 


1901] NORTH AMERICAN TREES 15 


broadly winged above, deeply grooved, villous, tinged with red 
below the middle, frequently twisted at midsummer, thus bring- 
ing the lower surface of the leaf-blades to the light, 1% to 2 in. 
long; stipules linear-lanceolate,.entire or coarsely glandular- 
serrate, % in. long, dark green fading red. Flowers bad- 
smelling, I in. in diameter when fully expanded, in loose lax 
compound many-flowered long-branched tomentose cymes ; 
calyx tomentose, the cup broad and shallow; calyx-lobes Jinear- 
lanceolate, entire, tomentose, elongated, persistent and closely 
appressed on the fruit; stamens ten; filaments slender, elongated ; 
anthers large, pale rose-color; styles two or usually three. Fruit 
pendulous in loose clusters, oblong to oblong-obovate, full and 
rounded at the base, 34 to 1 in. long, 34 in. broad, lustrous, 
bright carmine red, punctate with few dark spots, flesh thick 
mealy sweet and yellow; nutlets three or rarely two, thick, dark- 
colored, conspicuously ridged on the back, about seven-sixteenths 
of an inch long. 

A tree occasionally 20 feet in height with a tall trunk a 
foot in diameter covered with dark brown scaly bark, or in the 
immediate vicinity of the sea often shrubby with numerous stout 
spreading stems forming a broad massive bush, and stout wide- 
spreading or ascending branches zigzag for several years, dark 
green, tomentose and marked with numerous dark red oblong 
lenticels when they appear in the spring, becoming dark orange- 
color and very lustrous in the first season and light gray during 
their second year, and armed with straight or occasionally 
curved spines, 2 to 3 inches in length, dark chestnut-brown 
and lustrous and usually pointed toward the base of the 
branch. Flowers in early June. Fruit ripens and falls early in 
October. 

Borders of streams, where it grows to its largest size, and the 
rocky banks of ocean sounds and bays. Now known only in 
southeastern Maine, where it is distributed from the valley of 
the Penobscot river to the island of Mt. Desert. Orono, M. L. 
Fernald, May 27, 1887; Somesville, Rand & Redfield, June 1889; 
Somesville, EZ. Faxon, June 1890; £.L. Rand, Birch hill, Mt. 


16 BOTANICAL GAZETTE [JANUARY 


Desert island, June 1890, Northeast harbor and Southwest harbor, 
September 1899; Castine, C. E. Faxon, September 1889; Bar har- 
bor, Miss B. Jones, June and September 1899. 

It is a pleasure to commemorate in this handsome and distinct tree the 
name of Miss Beatrix Jones, the distinguished landscape-gardener, who first 
made it possible for me to understand its characters. 


ARNOLD ARBORETUM, 
Jamaica Plain, Mass. 


sae omiaag Toy, en Seatac AES Sia San eee 


ERIOCAULON DECANGULARE L.; AN ANATOMICAL 
o2 UD 


THEO. HOLM. 
(WITH FIVE FIGURES ) 


In a paper entitled ‘Structure de la racine et disposition des 
radicelles dans les Centrolepidées, Eriocaulées, Joncées, Maya- 
cées, et Xyridées,”* Van Tieghem arrives at the conclusion that 
these orders possess the same peculiarity in regard to the struc- 
ture of pericambium as do the Graminee and Cyperacee, and 
in this they appear to differ from the other orders of monocoty- 
ledons. His conclusions in regard to the Graminez and Cyper- 
acez, however, are mostly based on the observations of Johannes 
Klinge, which are recorded in his excellent work, ‘‘ Vergleichend 
histologische Untersuchung der Gramineen- und Cyperaceen- © 
Wurzeln.”’? It is the structure of the pericambium, Van Tieghem’s 
péricycle, which exhibits such marked variation as to seem charac- 
teristic of certain orders, at least in some of the species, and the 
variation consists in its continuity or interruption by the proto- 
hadrome vessels. Some very instructive tables are contained in 
Klinge’s paper, in which the position of these vesse!s has been 
given in relation to the pericambium of a number of species of 
Graminez and Cyperacee. From these tables it is seen that in 
some species all the proto-hadrome vessels are within the peri- 
cambium, in others only half of them or only a few, while in still 
others they are all in direct contact with the endodermis, having 
thus broken through the pericambium. Van Tieghem describes 
the same variation in Eriocaulacee, etc. Representatives of no 
less than seven orders of monocotyledons thus exhibit this pecu- 
liar structire. 

The continuity of the pericambial stratum is considered 

* Jour. de Botanique 1: 305. 1887. 

* Mém. de l’Acad. Imp. d. sc. d. St. Petersbourg VII. 26:12. 1879. 

Igor] 17 


18. BOTANICAL GAZETTE [JANUARY 


normal, its interruption abnormal. So far, botanists have not ven- 
tured to consider these two conditions as anything but specific 
characters, and if they were really constant they would certainly 
be of some importance; but my observations have lately shown 
that they are not to be depended upon as constant in all species. 
For example, in Carex jispidula3 the pericambium may be inter- 
rupted either by all the proto-hadrome vessels or by some of 
them, while in C. firma and C. supina this tissue is either con- 
tinuous or interrupted by the majority of these vessels. These 
variations were noticed in roots of a single individual, and it is 
very likely that such irregularities are much more common than 
is supposed, even in spite of an examination of abundant mate- 
rial. In regard to the Eriocaulacee, Van Tieghem describes the 
root-structure of a few species, and calls special attention to the 
very regular interruption of the pericambium by all the proto- 
hadrome vessels in #. decangulare, E. septangulare, E. Dregei, E. 
Sellowianum, and £. Paraguayense; while in £. atratum and E. 
Kunth only some of these vessels have broken through the 
pericambium. Moreover, this author attributes six or eight rays 
of hadrome to &. decangulare, and states that each ray is com- 
posed of two vessels, one bordering on endodermis, the other on 
the central vessel, but separated from it by a layer of conjunc- 
tive tissue. Besides Eriocaulon Van Tieghem has described 
Lachnocaulon,+* Paepalanthus, and Philodice. 

Having observed the irregular disposition of the proto- 
hadrome in some species of Carex, it seemed well to investigate 
the matter further, and I have examined the root structure of 
various other genera, and among them Eriocaulon; and since 
E. decangulare L.. exhibits a similar and even more striking varia- 
tion in its root structure, it was thought worth while to present 

3On a collection of Carices from Alaska, with remarks upon the affinities of 
-Carex circinata and C. leiocarpa C. A. Mey. Am. Jour. Sci. IV. 10:279. 1900. 


4 Judging from Van Tieghem’s description of the root structure in Lachnocaulon 


our North American Lachnocaulon Michauxii resembles that which Van ane has 
ascribed as sede to Paepalanthus, 


Igor] ERIOCAULON DECANGULARE 19 


the results. Furthermore, some notes upon the anatomy of the 
leaf and stem are appended, for it must be remembered that very 
little has been published heretofore upon the anatomy of Erio- 
caulon. As stated above, the root structure has been discussed 
by Van Tieghem, but only in reference to seven species of the 
genus ; and &. helichrysoides is the only one which ts treated from 
a general histological point of view by V. A. Poulsen in his ana- 
tomical study of the order. Besides these papers there are 
some few scattered remarks upon the structure of various organs 
of the genus in the works of DeBary, Russow, and Schwendener. 
There is thus only one paper in which a general anatomical 
treatment of the order has been presented, the work by Poulsen 
cited above. This author, who has studied fifteen species belong- 
ing to nine genera, especially Paepalanthus, all from Brazil, 
deserves credit for having detected several structural peculiari- 
ties; and it is a matter for regret that but one species of the 
large genus Eriocaulon has been examined. 

In the present paper some of the anatomical features of &. 
decangulare will be discussed, and attention will be called to such 
points as do not harmonize with the conclusions of Poulsen and 
Van Tieghem in reference to this species and others. The 
material was collected in sphagnum swamps in the vicinity of 
Washington, D. C. These swamps are wet during the entire 
season, so that the plants which were studied were not exposed 
to any drought during the summer months; furthermore, the 
plants were almost fully exposed to sunlight. Among the 
plants associated with Eriocaulon may be mentioned: Furena 
squarrosa, Scleria reticulata, Eleocharis tuberculosa, Rhynchospora alba, 
R. glomerata, R. cephalantha, Xyris flexuosa, Pogonia ophioglos- 
soides, Drosera rotundifolia, Utricularia subulata, Rhexia Mariana, 
and Asclepias rubra. 

THE ROOT. 

The roots of &. decangulare studied were all secondary, being 
developed from the rhizome. Two kinds may be distinguished : 
some that are quite thick, white, mostly unbranched, and with 


5 Anatomiske Studier over Eriocaulaceerne. Thesis, Copenhagen. 1888. 


20 BOTANICAL GAZETTE [JANUARY 


few root hairs; and others that are less white, somewhat thinner 
but decidedly longer, with many lateral rootlets, and amply pro- 
vided with root hairs. No special arrangement of these two 
forms was noticed, but the thick ones were perhaps the more 
abundant. Both forms appeared to have developed in the same 
year, and I was unable to detect any pronounced difference in 
their internal structure that might warrant any further distinc- 
tion than the one observed in their external appearance. 

The thick white roots show a very weak structure in general, 
which seems to imply that they are of short duration, and prob- 
ably do not last beyond one season. The epidermis is very thin 
walled, and root hairs are present, but rather scarce. The root 
hairs observed were all single, not in pairs, as Van Tieghem 
observed in species of Paepalanthus (Lachnocaulon?). Poulsen 
did not find any root hairs at all in E. helichrysoides, the speci- 
mens of which had perhaps not been lifted with sufficient care. 
Within the epidermis is a hypodermis of a single stratum, the 
cells of which are of the same size as those of the epidermis, and 
are also thin walled. The cortical parenchyma is quite broad, 
but almost totally collapsed radially, excepting the innermost 
stratum which borders on the endodermis, and which is often 
somewhat thick walled in contrast with the other portion of the 
cortex. There are thus several very wide lacunes in the cortex, 
and these contain horizontally placed diaphragms composed of 
star-shaped, thick walled cells, which contain chlorophyll. The 
lateral roots do not immediately break through the epidermis, 
but traverse the cortex for some distance downwards before they 
penetrate the hypodermis and epidermis. A similar course of 
the lateral roots was observed by Poulsen in Carpocephalus cau- 
lescens Kth. 

The endodermis (jig. z, Z) is usually thin walled, and the 
spots of Caspary are plainly visible. However, in some roots 
the cell walls of the endodermis are slightly thickened, especially 
in the basal portion of the root. The pericambium (figs. 7, 2, P) 
is very thin walled, and consists of a single layer, which in no case 
was observed to be continuous, although a very large number 


1901] ERIOCAULON DECANGULARE 21 


naira 


Ke hy 


CR 
ors. 


Figs. 1-5. ERIOCAULON DECANGULARE L. 


Fic. 1. Transverse sectionof a thick root: C, innermost stratum of cortex ; 
£, endodermis: P, pericambium; //, proto-hadrome vessel; the proto- 
leptome cells are drawn with heavy lines; the drawing shows five proto-had- 
tome vessels, all bordering on endodermis, and alternating with five groups 
of leptome; a large vessel occupies the center of the root. x 495. 

Fig. 2. Transverse section of a part of a thick root; letters as above; V, 
the two central vessels; one of the proto-hadrome vessels has not broken 
through the pericambium. X 495. 

Fig. 3. Transverse section of a lateral root; letters as above; there are 
only two proto-hadrome vessels, both of which border on endodermis. X 495. 

_ FIG. 4. Transverse section through a large mestome bundle from the 
leaf; 4, epidermis of upper face ; B, same of lower surface ; M7, the collenchy- 
matic tissue, which surrounds the mestome bundle completely and extends 
to the epidermis on both faces of the leaf; S, the inner sheath, which forms 
a closed ring around the leptome and hadrome; in the hadrome is a lacune 
with a ring vessel. x 320. 

FiG. 5. Transverse section of a smaller mestome bundle from the leaf; 
letters as above ; the inner sheath (.S) is interrupted by the vessels. X 495- 


22 BOTANICAL GAZETTE | JANUARY 


of sections were examined. To examine the position of the 
proto-hadrome in relation to the pericambium, not only were a 
number of sections taken from roots of various individuals, but 
also several roots of the same specimen in their entire length 
from apex to base. The latter method appears to give the most 
satisfactory results, and is sufficient to demonstrate that neither 
the position of the proto-hadrome vessels, nor the number of 
hadromatic rays or of the individual vessels is to be depended 
upon as a constant character. 

A few examples of these structural irregularities are as follows. 
One of the thick roots possessed at the base eight rays of proto- 
hadrome, each consisting of a single vessel, five of which had 
penetrated the pericambium, thus bordering immediately on 
endodermis. Nearer the apex of the same root the number of 
rays decreased from eight to seven and to six, five of which had 
broken through the pericambium, as in the first case. At the 
apex itself there were only five rays, all of which were bordering 
on the endodermis; thus the number of interruptions of the 
pericambium was five in the entire length of this root. In other 
roots only five rays were found at the base, four of which had 
broken through the pericambium ; and here also there was a 
similar decrease in number of rays when approaching the apex, 
where finally all the proto-hadrome vessels bordered on the 
endodermis. The number of interruptions may vary in the same 
root, instead of being five, for instance, throughout the entire 
root; or there may be roots in which all these vessels border on 
the endodermis, the most frequent condition (fg. 7). Van 
Tieghem ascribes six to eight hadrome rays to this species, each 
consisting of two vessels, but such regularity in number was not 
observed in my material. Not only did the number of rays vary, 
usually decreasing from base to apex, but five was the most com- 
mon number, each ray consisting of a single proto-hadrome 
vessel (fig. 1, 1) instead of two. The largest number of rays 
observed was eight, but this number, as well as seven, was rather 
rare, while six or four was not uncommon. While the proto- 
hadrome vessels were observed as being mostly one in each ray, 


1901] ERIOCAULON DECANGULARE 23 


sometimes two, three, or even four together were observed, 
either in the same radius or side by side. This multiplication 
of vessels was found in the same root in which the majority of 
sections showed only a single vessel in each ray. 

In considering the pericambium itself another irregularity in 
respect to the number of cells between each two proto-hadrome 
vessels was observed. Two may be found in many roots, and 
is no doubt the commonest number; three cells were found, 
however, in the same roots, but seldom four or five. The leptome 
forms groups which are roundish in transverse section, and the 
proto-leptome is plainly visible. The central portion of the 
root was constantly found occupied by one or two wide reticu- 
lated vessels, mostly surrounded by a thin walled conjunctive 
tissue. 

The thinner secondary roots show much the same structure 
as described above, but root hairs abound. The innermost 
stratum of the cortex is sometimes distinctly thick walled, and 
the endodermis often exhibits a similar thickening of its walls. 
These roots show the same irregularities in the structure of the 
pericambium and the hadrome as described above, and there is 
also a wide central vessel surrounded by a thin walled conjunc- 
tive tissue. 

The lateral roots exhibit a very simple structure (fig. 3). No 
hypoderm is developed, the cortex bordering immediately on 
epidermis, and composed of only two or three strata, the innermost 
of which persists (jig. 3, C), while the others sometimes collapse 
Tadially. The endodermis (fig. 3, Z) is very thin walled, the 
spots being very plainly visible; and the pericambium is inter- 
rupted by the two rays of proto-hadrome, which alternate with 
two minute groups of leptome. No central vessel was observed. 
The peculiar course of the lateral roots in the cortical parenchyma 
before they break through the hypoderm and epidermis has 
been described, and it might be added that root hairs are 
_ absent from the enclosed part of these roots, but are developed 
aS soon as the root became free. . 

The roots of Eriocaulon may well be classified as simply 


24 BOTANICAL GAZETTE [JANUARY 


“nutritive,” as has been described by Rimbach,° since they pos- 
sess no pronounced power of resistence, and are not contractile 
or especially adapted for storage. 

THE RHIZOME. 

According to Gray’s Manual (sixth edition) all our species 
should be ‘‘stemless,” but Morong’ has corrected this and 
describes for £. decangulare a short thick caudex one or two 
inches in length. This species has a nearly horizontal or ascend- 
ing rhizome, densely covered by remnants of old leaf bases, 
with no internodes. The numerous long hairs developed from 
the epidermis are very characteristic, being pluricellular, the 
cells in a single row, and the basal cell very short. The cortex 
is differentiated into two or three subepidermal strata, the cells 
of which are mostly pentagonal in tranverse section, with dis- 
tinct but narrow intercellular spaces, and an inner tissue of 
many layers of nearly roundish thin walled cells containing 
starch. This portion of the cortex is very open on account of 
large intercellular spaces, but no lacunes were observed. Within 
the cortex is an endodermis, the cell walls of which are not 
thickened and do not show the characteristic spots of Caspary. 
Nevertheless, the endodermis was readily visible by the peculiar 
clearness of its cell walls in contrast with the surrounding corti- 
cal parenchyma, and by the somewhat irregular shape of the 
cells. Within the endodermis is a large, solid, fundamental tissue, 
consisting of somewhat thick walled cells with distinct inter- 
cellular spaces, and containing starch. The mestome bundles 
occur in the cortex and in the fundamental tissue. Those in the 
latter tissue, being thus within the endodermis, are not arranged 
in any order, and are mostly bicollateral and perihadromatic, but 
not always completely so. They are not surrounded by any 
special sheath, and the vessels are either scalariform and quite 
narrow or reticulated and wider. 


‘ ®Beitrage zur Physiologie der Wurzeln. Ber. d. deutsch. Bot. Gesell. 17: 186 
1899. 


7 Notes on the North American species of Eriocaulez. Bull. Torr. Bot. Club 
18: 354. 1891. 


- eee ee ee i : 


1901] ERIOCAULON DECANGULARE 25 


When the mestome bundles break through the endodermis 
to the cortical parenchyma, they appear much smaller and are 
then surrounded by a thin-walled endodermis, showing the same 
power of resisting concentrated sulfuric acid as does the main 
endodermis. They are almost regularly arranged in two con- 
centric bands, and are either strictly collateral or more or less 
bicollateral, in the same manner as those of the fundamental 
tissue. The general structure of the rhizome, therefore, does 
not differ from that of other monocotyledonous plants, with the 
exception of the development of the epidermis into long hairs, 
which is known in a very few other orders. The mestome 
bundles of Eriocaulon being collateral or more or less com- 
pletely perihadromatic do not differ from those of rhizomes of 
most of the other monocotyledonous orders. Attention may be 
called, however, to a very peculiar structure which Poulsen dis- 
covered in the rhizome and stem of Actinocephalus polyanthus 
Kth. (Eriocaulacee), and which may be found in some of the 
other genera. The mestome bundles exhibit a form very 
unusual among the phanerogams, in being perileptomatic in 
the cortex and exohadromatic in the fundamental tissue; in the 
latter the perihadromatic bundles are surrounded by a tissue 
Which is suggestive of leptome, and around this again is a band 
of vessels, each mestome bundle consisting thus of a central 
Stroup of leptome surrounded by two bands of hadrome sepa- 
rated from each other by a band of apparently leptomatic tissue. 


THE SCAPE. 

The inflorescence, a capitulum, is borne at the apex of a long, 
slender, solid and twisted scape, which is distinctly furrowed 
and consists of only one internode with a single leaf, the long 
tubular sheath of which surrounds the scape to about its middle. 
In a transverse section of the free part of the scape, there is 
shown an epidermis of roundish cells, covered by a thin, smooth 
Cuticle. The size of the cells varies somewhat, and the largest 
are observable in the furrows, where they cover the green cor- 
tex. Stomata and hairs are present and exhibit the same 


26 BOTANICAL GAZETTE [JANUARY 


structure as in the leaves, under which they will be described. 
Underneath the epidermis is either a chlorophyll bearing corti- 
cal parenchyma (in the furrows) or a mechanical tissue (in the 
ridges). The former in our species occurs in nearly rectangu- 
lar groups in transverse section, separated from one another by 
the mechanical tissue. There is but one stratum of palisade 
cells, directly beneath the stomatiferous epidermis; while the 
other part of the green cortex consists of loosely connected 
cells of irregular shape and with very wide air spaces. No 
proper lacunes are developed in the cortex, nor are there any 
such diaphragms as in the leaves. 

The mechanical tissue constitutes quite a prominent portion 
of the stem section. It is this tissue which forms the eight 
ridges, and it extends inwards to the mestome bundles, which it 
surrounds as a closed ring, and also occurs as a few layers on 
the hadrome side of the mestome bundles. <A close examina- 
tion of this tissue, however, shows that even if it may be desig- 
nated as ‘“‘mechanical” throughout, it nevertheless represents 
two distinct kinds of tissues, collenchymatic and stereomatic, 
both somewhat modified in our species. The collenchymatic 
tissue reaches its highest development in the ridges of the scape, 
just beneath the epidermis. i1t appears there composed of dis- 
tinctly thickened cells, but roundish in cross section and with 
plainly visible intercellular spaces; viewed in longitudinal sec- 
tions these cells are rectangular and quite long. As mentioned 
above, this tissue extends inwards between the groups of green 
cortex to the mestome bundles, and occurs here as a closed ring 
of two or three strata. I have been unable to discover any dis- 
tinction between the strata which surround the band of mestome 
bundles and those which form the ridges. The thickening of 
the cell walls is of course most pronounced in the ridges, and 
especially near the epidermis, but if the tissue is followed 
inwards to the mestome bundles the thickness decreases very 
gradually, and the shape of the cells remains the same in both 
longitudinal and transverse sections. The term “ collenchy- 
matic,’ as suggested by Schwendener for this special tissue in 


1901] ERIOCAULON DECANGULARE 27 


Eriocaulon, seems more suitable than ‘‘stereomatic’’; moreover, 
it appears that it attains a still further development in other 
genera of the order, where it sometimes resembles true collen- 
chyma much more than in our species. 

The second form of mechanical tissue in the scape so much 
resembles stereome in its structure that it may be considered as 
stereomatic, but not as stereome proper. This tissue is but 
sparingly developed, and occurs only in one or two strata cover- 
ing the leptome of the bundles within the collenchymatic ribs, 
and as a few cells on the hadrome side of these same bundles. 
The lumen of these stereomatic cells is much narrower than in 
any portion of the collenchyma; moreover, the cells are longer 
and the cross walls are almost oblique. In no instance, however, 
did I observe this stereomatic tissue form a sheath around the 
mestome bundles; it seems to be restricted to a few strata on 
either face, separated from one another by the large vessels on 
the sides of the bundles. The small mestome bundles which 
are located within the chlorophyll bearing cortex are destitute 
of such stereomatic support. It would thus appear as if the 
mestome bundles are merely surrounded by continuous strata of 
collenchyma which only differs from that in the ridges in being 
less thick walled; and there is no indication of such structural 
peculiarities as might lead to a separation of it into two tissues, 
as a collenchymatic and an endodermal. The cells showed 
exactly the same shape and size all around the central cylinder, 
and in no place were the spots of Caspary, or any special 
arrangement usually noticeable in an endodermis, observed. 
However, when the sections were placed in concentrated sulfuric 
acid, a true endodermis became at once plainly visible. The 
collenchyma was rapidly destroyed, while a continuous ring of 
a single stratum of cells persisted for some time surrounding 
the mestome bundles and bordering on the leptome, but sepa- 
rated from it by the few strata of stereomatic tissue. The scape 
thus possesses an endodermis of the same structure as that 
observed in the rhizome. 

It has been stated already that the mestome bundles occur 


28 BOTANICAL, GAZETTE | JANUARY 


of two sizes, both constituting a single band, the larger situated 
in the same radius as the collenchymatic ridges, the smaller in 
alternation with them within the cortex. The leptome forms 
quite a large oval or roundish group in both kinds of bundles, 
all of which are collateral; and the hadrome is composed of 
very wide reticulated vessels which form a continuous arch in 
the smaller bundles, sometimes in the shape of a V. In the 
larger mestome bundles the arch is less complete, and here a 
small lacune with remains of an annular vessel was observed. 
This structure of the mestome bundles suggests what is known 
among other orders of monocotyledons, especially the larger 
ones. Finally, the pith, which occupies the inner portion of the 
central cyclinder, is solid, not hollow, but the cell walls are very 
thin, and no deposits were observed. 

The base and apex of the scape show the same arrangement 
and relative development of the various tissues, but no stomata 
or hairs were observed on the part enclosed by the tubular leaf 
sheath. It might be stated, also, that although the mestome 
bundles were noticed to be strictly collateral in a large number 
of sections, one instance was found in which a few (three) nar- 
row scalariform vessels were situated outside the leptome, this 
mestome bundle being thus approximately perihadromatic. It 
was only in one bundle, however, and very near the inflorescence. 
The structure of the bundles, therefore, does not seem to be invari- 
ably identical throughout the whole length of the stem. It must 
be remembered, also, that in the rhizome the bicollateral bundles 
within the endodermis pass gradually over into collateral when 
they enter the cortical parenchyma, whence they proceed to the 
leaves, in which they are constantly and strictly collateral. 


THE LEAF. 


The proper leaves are linear, nearly flat or with the margins : 
somewhat. involute, and have no sheath. They form a dense 
rosette at the apex of the rhizome, and in their axils the flower 
bearing scapes develop on very short one leaved shoots, while 
the terminal bud of the rhizome itself is purely vegetative. In 


190r] ERIOCAULON DECANGULARE 29 


contrast with the proper leaf, the leaf of the floral shoot has a 
long tubular sheath and a very short blade, and this leaf is by 
some authors considered as representing the prophyllum of the 
floral shoot. No other leaf is developed on this shoot, but its 
true position in relation to the mother axis cannot be ascertained, 
since the scape becomes twisted during its growth and forces 
the leaf away from its natural position. Thus the little blade 
may sometimes turn to the side or to the front, instead of turn- 
ing its dorsal face towards the main axis, as is the normal posi- 
tion of such a prophyllum. As will be shown later, the structure 
of this leaf is very different from that of prophylla as a rule, 
though some analogous cases do exist in other orders, where a 
similarly developed leaf unquestionably represents a true prophy]- 
lum. Attention might be called especially to the clado-prophyl- 
lum in the inflorescence of certain Cyperacee, as for example, 
Scirpus silvaticus, S. polyphyllus, Cyperus phymatodes, C. strigosus, 
and many others. So far, there is no very strong objection to 
considering the leaf of Eriocaulon as a prophyllum, even if its 
position cannot be determined accurately. There is no very 
Pronounced difference between these two kinds of leaves, but 
they will be treated separately in order that the view of their 
general structure may be made as distinct as possible. 

The proper leaf, in sections taken from the middle of the 
blade, shows a large celled epidermis on both surfaces, especially 
On the upper. There is a thin, but visible cuticle, which shows 
minute wrinklings on the lower surface, but is smooth on the 
upper. None of the epidermal cells are arranged so as to form 
longitudinal rows of bulliform cells, such as are so well known 
in many genera of Cyperacee and Graminez, where they espe- 
cially abound above the keel of the blade or between some of 
the larger mestome bundles. The cells in Eriocaulon are very 
large on the upper face, but decrease very gradually in size from 
the middle towards the margin, the outermost part of which is 
composed only of the two strata of epidermis corresponding to 
the two leaf surfaces. Viewed en face the radial cell walls are 
perfectly straight on both surfaces, and the outer cell wall of 


30 BOTANICAL GAZETTE [JANUARY 


the dorsal epidermis is somewhat thicker than the others. 
The inner wall may sometimes show a peculiar bend inwards, 
the lumen of the cell thus becoming diminished, while at the 
same time a wide air space is formed beneath the cell. This 
case was noticed only on the lower face of the blade, and Poul- 
sen found a similar structure, but much further developed, in 
the leaves of Dimeranthus and Actinocephalus. While hairs are 
present on both faces of the blade, stomata are restricted to 
the lower surface. The hairs are small and much shorter than 
those on the rhizome, consisting of but three cells, the terminal 
one being the longest. They are closely appressed to the leaf 
blade and point towards its apex. This form of hair was also 
observed by Poulsen in &. helichrysoides, and its early stage of 
development is represented in his f/. 5, figs. 2-4. 

The stomata occur only on the lower surface, outside the 
mesophyll, and possess very narrow guard cells and subsidiary 
cells, both of which, especially the former, are raised a little 
above the surrounding epidermis. 

The mesophyll in our species constitutes a very uniform struc- 
ture throughout, and the leaf in this respect may be considered 
isolateral. There is no continuous layer of palisade cells on the 
upper face, but the tissue consists of a single stratum, only some 
of the cells of which represent true palisade. Most of the other 
cells are roundish and very loosely connected with one another. 
On the lower face the cells of the mesophyll show a similar round- 
ish form, but the intercellular spaces are not so wide as on the 
upper face. A like structure is represented by the mesophyll 
_ around the mestome bundles, where it is also composed of a 
single stratum bordering on the mechanical tissue. The inner 
portion of the mesophyll is broken down into wide lacunes, one 
between each two mestome bundles, in which are found numer- 
ous diaphragms of star shaped, chlorophyll bearing cells, exhib- 
iting much the same structure as those in the root. 

Attention has been called, in connection with the stem, to 
the difficulty in designating the proper term for the mechanical 
tissue as it is developed in Eriocaulon; and the same is true 


1901] ERIOCAULON DECANGULARE 31 


in regard to it in the leaf, where there is to be observed 
collenchyma and another form of tissue which superficially 
resembles stereome, but which shows such strange occurrence 
and development that it is not even to be considered as a modi- 
fication of this tissue. The collenchyma in the leaf is thinner 
walled than in the stem, and accompanies the mestome bundles 
mostly as hypodermal groups on both faces. It appears to be 
more strongly developed on the lower face than on the upper, 
but each group is composed of only three or four layers of these 
cells (fig. g, M). Around each mestome bundle is a sheath, 
which at first glance was considered identical with the paren- 
chyma sheath (fig. 5, M), as seen in many genera of other 
orders. A longitudinal section, however, showed a structure 
very unlike that of a typical parenchyma sheath. The cells 
were relatively very long, and I was unable to separate them 
from the adjoining cells of the collenchyma, inasmuch as they 
both showed the same rapid dissolution when treated with con- 
centrated sulfuric acid. It is somewhat strange, also, that no 
trace of this outer sheath was found in the stem, a fact which 
seems to be in favor of the supposition that it is not identical 
with a true parenchyma sheath. When such sheaths are devel- 
oped, they seem to follow the mestome bundles not only in the 
leaves but also in the stem, at least in those which are in the 
cortex, but not in the pith. 

An examination of one of the large mestome bundles ( fig. 
4) reveals the fact that still another sheath (.S) occurs within 
the first. This is distinctly thicker walled than the outer, the 
cells are very much smaller, and the walls are very bright 
-yellow in contrast with the surrounding collenchyma. In longi- 
tudinal section the cells of this inner sheath are quite long, 
with mostly horizontal cross walls. A study of its occurrence 
throughout the leaf has assured me that it does not show the 
Same development in all the mestome bundles. In smaller 
bundles the sheath is a mere arch on the leptome side, while the 
vessels border immediately on the outer collenchymatic sheath. 
In the larger mestome bundles there is the same arch on the 


32 BOTANICAL GAZETTE [JANUARY 


leptome side, besides one (jig. 5, S), or sometimes two or three 
cells on the hadrome side, between the large vessels. Selecting 
one of the largest bundles in the leaf, the sheath appears as a 
closed continuous sheath (fig. 4, S) all around the leptome and 
hadrome. It seems characteristic of this inner sheath to show 
interruptions in the small mestome bundles, and to become con- 
fined in the leptome alone in the smaller of them. I regret that 
I was unable to decide whether the inner sheath is also devel- 
oped in the smallest of the mestome bundles, the anastomoses. 
The tissues are so little differentiated in these that it seemed 
impossible to decide whether the sheath was present or not; the 
outer one, however, was plainly visible. So far there would be 
no valid objection to considering the inner sheath as identical 
with a mestome sheath,® which, in several Graminez, exhibits 
the same kind of interruptions in the various forms of mestome 
bundles. However, there are two reasons for feeling uncertain 
about its identity. It is absent in the mestome bundles of the 
scape, where two or three strata of stereomatic tissue have taken 
its place within a true endodermis, and in longitudinal section 
the cells differ in a marked degree from those of a mestome 
sheath, being very long, with almost horizontal cross walls. 
Moreover, the thickening of the inner cell wall does not show 
the same structure as in typical mestome sheaths, where this 
wall almost constantly exhibits a conspicuous thickening in con- 
trast to the outer wall. When treated with concentrated sulfuric 
acid, however, the sheath in Eriocaulon showed the same resis- 
tance as in the endodermal sheaths in the mestome bundles in 
the cortex of the rhizome. If the outer sheath represents a 
parenchyma sheath, one might feel justified in considering the 
inner as a somewhat modified mestome sheath; but since it does 
not seem possible to separate the outer sheath from the collen- 
chyma, I can at present suggest no other term for the inner one 
than simply a protective sheath. 

Poulsen attributes two sheaths to the mestome bundles in the 


® SCHWENDENER, S.: Die Mestomscheiden der Gramineenblitter. Sitzungsber. 
K. Acad. Berlin 405. 1890. 


ee a ae ee ae nr oc en OES 


1901} ERIOCAULON DECANGULARE 33 


leaves of the genera which he examined, and, in accordance 
with his opinion, the outer sheath would represent true paren- 
chyma, the inner sclerenchyma. He saw, however, that the 
so-called sclerenchyma does not show the same development in 
all the bundles, but he makes no allusion to its possible identity 
with a mestome sheath. - The outer sheath is only described as 
being composed of thin walled cells ‘‘stretched longitudinally,” 
which does not seem to indicate that a true parenchyma sheath 
was observed. The testing of these sheaths with sulfuric acid 
seems to have been omitted. 

The mestome bundles of the leaf are almost parallel from 
the base to near the apex, where they unite and form an appar- 
ently single bundle. When viewed superficially the nerves 
might appear without anastomoses, since the numerous dia- 
phragms filled with chlorophyll make the inner portion of the 
leaf very indistinct. When the blade is divided into halves the 
anastomoses may be seen, not at all infrequent, connecting the 
larger nerves at more or less acute angles. In this way anasto- 
moses become plainly visible even without any further treatment 
with potassium hydrate, and they are often seen in transverse 
sections. It seems very surprising that Poulsen did not notice 
_ them in any of the fifteen species which he examined, and that 
he points out the absence of anastomoses as one of the principal 
characteristics of the order. It may be that they are not devel- 
oped in any of these fifteen species, but they are certainly very 
plainly visible in £. decangulare, and I have found them also in 
E. gnaphalodes and in Lachnocaulon Michauxii. 

The structure of the main nerves is very uniform. None of 
the mestome bundles near the middle of the blade are of any 
considerable size when compared with others, and it might be said 
that the leaf has no midrib. It has already been stated that 
some of the nerves are smaller than others, and that a corres- 
ponding variation in regard to the development of the inner 
sheath is noticeable. However, the hadrome and leptome are 
common to all the main nerves, but are naturally larger in the 
thicker mestome bundles than in the thinner. The leptome 


34 BOTANICAL GAZETTE [ JANUARY 


shows the same structure as in other orders, for example 
Cyperacee, Graminee, etc. The hadrome shows several rela- 
tively wide vessels (reticulated) besides a lacune, in which there 
occurs a ring vessel. The arrangement of the vessels is the 
same as in Graminee, etc., and in no instance was there seen the 
v-shaped hadrome as in the scape. Fhe very smallest of the 
mestome bundles exhibit both leptome and hadrome in two well 
differentiated groups, and several narrow vessels are developed 
in them, bordering immediately on the collenchyma, while the 
leptome has an arch-shaped support of an inner sheath, con- 
stantly incomplete in such small bundles. 

The base of the leaf differs but slightly in structure from 
the middle of the blade. The lacunes are wider, the mesophyll 
is more irregular, and no palisades were observed ; furthermore, 
the inner sheath of the mestome bundles is somewhat more 
thickened, but shows otherwise the same kind of interruptions 
in the small bundles. The epidermis does not seem to have 
become modified in any respect, and both hairs and stomata 
occur. The structure of the apex, however, is very different. 
The epidermis is unchanged, but the mesophyll presents a solid 
palisade tissue in three layers on the lower face, while the upper 
one consists of more roundish cells with narrow intercellular 
spaces; the mestome bundles are fused together into one oval 
group surrounded by a sheath of thin-walled cells, which evi- 
dently corresponds to the outer one, the collenchymatic ; and it 
may be added also that in the apex this sheath is not able to 
resist sulfuric acid. There is thus a single group of mestome in 
the apex, and the leptome seems much reduced, while the had- 
rome occupies a broad space and contains many very wide ves- 
sels, especially reticulated. This single mestome bundle is not 
supported by any hypodermal mechanical tissue or by an inner 
sheath, as farther down the blade. 


THE PROPHYLLUM. 


The prophyllum consists of a very long tubular sheath and a 
short free blade, in both of which there is much the saine 


1901} ERIOCAULON DECANGULARE 35 


structure as in the proper leaf. The tubular portion, however 
possesses an epidermis of several (three) layers on its morpho- 
logically ventral face, and this epidermal tissue is homogeneous 
in structure, the cells being of the same size and shape throughout. 
The dorsal epidermis consists of only one stratum, and the cells, 
are about the same size as those of the ventral epidermis. The 
mesophyll is very open on account of wide lacunes, as in the 
proper leaf. The mechanical tissue is developed further in the 
prophyllum than in the other leaves in having larger groups on 
the dorsal face and in being distinctly thick walled, almost like 
collenchyma. The mestome bundles are surrounded by this tis- 
sue, and also by the inner sheath, which is much less differen- 
tiated, however, in this leaf, being totally absent in the smallest 
bundles. Furthermore, a somewhat weaker development of the 
hadrome and leptome is to be seen in this portion of the pro- 
phyllum. It may be stated that the structure of the collen- 
chyma was plainly followed to several of the cells of the outer 
sheath ; thus there seems little doubt that this sheath actually 
represents a part of the mechanical tissue itself, instead of being 
a specially developed parenchyma sheath. 
The little blade resembles very much that of the proper leaf, 
in regard to the epidermis with its hairs and stomata, and the 
mesophyll with its lacunes; while the collenchyma is separated 
from the dorsal epidermis by strata of small palisade cells. 
There is the same distinct thickening of the collenchyma, and 
very plainly so where it surrounds the nerves; the inner sheath 
is but very incompletely developed in this portion of the leaf. 
The mestome bundles are almost orbicular in transverse section, 
and exhibit the same difference in relative size as described 
above; and finally anastomoses more frequently occur in the 
Prophyllum from apex to base. Thus the inner sheath is present 
also in the mestome bundles of the prophyllum, though less 
completely developed; and the same result was obtained as 
before when treating the sections with concentrated sulfuric acid, 
that is, the outer sheath dissolved at once, together with the 
adjoining cells of the collenchyma, from which it must be 


36 BOTANICAL GAZETTE [JANUARY 


considered as inseparable, while the inner sheath showed some 
resistance wherever it occurred. 


CONCLUSIONS. 

In comparing the general structure of £. decangulare with 
that of &. helichrysoides, as described by Poulsen, there seem to 
be certain peculiarities in common which may be characteristic 
of the order, inasmuch as they have also been observed in some 
of the other genera. The presence of collenchyma is one of 
these, although this tissue should not be considered so rare in 
monocotyledonous orders as Poulsen claims it to be. It is com- 
mon, for instance, among the Graminez, where it is well devel- 
oped in the nodes; it is also foundin the small “ pulvini’” which 
occur at the base of the lateral branches in paniculate inflores- 
cences of a number of genera. Moreover, in the Cyperacee 
collenchyma occurs in the basal portion of both rachis and pro- 
phyllum in certain species of Cyperus of the subgenera Eucype- 
rus, Diclidium, and Mariscus; besides in such species of Scirpus 
as possess a large open inflorescence (S. atrovirens, S. polyphyl- 
‘ dus, and others). A still more advanced development of this 
tissue may be observed in Commelinacee, Smilacez, and Dios- 
coreacez. It does not seem, therefore, as if this tissue exists 
only in ‘‘a very few monocotyledons.”’ But it seems charac- 
teristic of the Eriocaulacee examined that the collenchyma, or 
to be more exact ‘‘collenchymatic tissue,” occurs as prominent 
ridges in the stem, that it replaces the stereome in the leaves, 
and that it surrounds the mestome bundles as a closed sheath. 
The inner mechanical sheath, noted in the leaves, appears to have 
several points in common with the mestome sheath of the Gram- 
inez, yet it does not seem to be identical with this, and perhaps 
may be characteristic of Eriocaulon. The hairy rhizome consti- 
tutes an excellent character of the order, even if a few instances 


are known in other orders of hairs on underground stems, Viz 


Corallorhiza, Scheuchzeria, and Mercurialis. Otherwise the 
structure of Eriocaulon does not differ so very much from that 
of certain other orders, as Cyperacee and Juncacez, in which a 


naan sanescanenrmacaremnienmnanmasiaanasnitite is neal re | 


en 


1901] ERIOCAULON DECANGULARE 37 


similar leaf structure is provided with a uniform large celled 
epidermis (Kyllingia, Cyperus, Fimbristylis, etc. }. Also, the 
structure of the rhizome and roots is not to be separated from 
that of a number of Gramineze, Cyperacee, etc. The most 
peculiar structure is exhibited by the scape, not only in the pres- 
ence of collenchymatic ridges, but also in the mestome bundles 
surrounded by a common endodermis. 

But neither the order nor even the genus can be character- 
ized especially by absence of anastomoses in the leaves, since 
both occur in Eriocaulon and Lachnocaulon; and the continuity 
of interruption of the pericambium by the proto-hadrome ves- 
sels in the root is not of any importance as a family character, 
as it is not constant, at least in Z. decangulare. 

It appears, therefore, as if Eriocaulon and its allies possess 
some characters in common with other monocotyledonous orders, 
and a few which seem peculiar to themselves. The singular 
inflorescence which is unique among the monocotyledons, and 
the floral structure, together with the peculiar habit of these 
plants, make them very distinct among the other orders, to none 
of which they make any special approach, not even in respect to 
their morphological characters. 


BROOKLAND, D. C. 


PHYSIOLOGICAL STUDIES WITH REFERENCE TO THE 

GERMINATION OF CERTAIN FUNGOUS SPORES. 

B. M. DUGGAR. ; 
INTRODUCTORY AND HISTORICAL. 

Tus study was entered upon with the view of ascertaining 
somewhat more definitely than previous researches have indicated 
what may be some of the special factors which influence germina- 
tion. As particular lines of inquiry the following may be men- 
tioned as suggestive. In how far does there exist in the spores of 
fungi an essential physiological difference, whereby some may ger- 
minate by the mere absorption of water, while others may require 
for this germination a food supply from without? Where a food 
supply is required, does germination require a perfect food or a 
particular food? May a chemical irritant, or poison, which is 
not primarily a food substance, thus function as a stimulus? Is 
it possible by mechanical means, or by a change of conditions, 
to furnish the necessary stimulus for germination? Considering 
the broad field thus suggested, it has yet been possible to study 
but a very limited number of fungi. Moreover, there are 
numerous minor questions which must be considered in a sub- 
sidiary way. 

Using the term germination in its broadest sense, it may be 
well at the outset to notice some of the conditions of food sup- 
ply or stimulation characterizing germination in general. As a 
rule, the seed of the phanerogam requires for germination only 
water, along with suitable conditions of temperature and requisite 
oxygen supply. Not even does the force of imbibition of the 
seed coats have any special action in inciting to activity the 
dormant faculties. In about a dozen plants upon which I have 
experimented, the uninvested embryo, that is, entirely free from 
integuments or externally stored food material, is capable of 
manifesting the first stages of germination in distilled water. 


38 [JANUARY — 


Igor] GERMINATION OF SPORES 39 


A few phanerogams which have become adapted to a peculiar 
environment may require a stimulus of this environment. Koch’ 
and Heinricher? have shown that seeds of Orobanchaceze ger- 
minate only with the presence of a host plant. Heinricher found 
that the seed usually germinates in spring or autumn, in periods 
of greatest humidity. He states that the germination of the 
seed is indirectly a partial indicator of the health of the host — 
a substance excreted by the latter being evidently the source of 
the stimulus. What may be the nature of this attractive sub- 
stance has not been determined. On the other hand, seeds of 
Rhinanthus seem to germinate without such stimulus. 

According to Wiesner 3 the seeds of Viscum album are known 
to germinate only in light, and although apparently mature in 
autumn, they are not to be forced to germination until the fol- 
lowing spring. The same was found true of Loranthus Europaeus ; 
and, although light is required for germination, the most favor- 
able conditions of the tropical climate of Buitenzorg could not 
induce this activity without the intervention of the resting period. 
Wiesner then surmised that the factors concerned are the grad- 
ual availability of the reserve food, a phylogenetic light influ- 
ence, and the effect of the viscous substance of the integuments. 
If there are other phanerogams growing in peculiar situations 
for which a particular stimulus is necessary for germination, the 
matter seems to await study. We cannot appropriately include 
in this place seeds of some Rosaceae, for example, which germi- 
nate better after passing through the digestive tract of birds, or 
after artificial treatment with acids. 

The germination of pollen has been very much studied, and 
it is Surprising that this has yielded so little of special inter- 
est relative to particular or peculiar stimuli. It is of special 
es: Kocn, L.: Die Entwickelungsgeschichte der Orobanchen 119. 1887. Heidel- 

*HEINRICHER, E.: Die Keimung von Lathrea. Ber. d. deut. bot. Gesells. 
12: 117-132. 1894. : 

* Wiesner, J.: Vergleichende physiologische Studien iiber die Keimung europii- 
on und tropischer Arten von Viscum und Loranthus. Sitzb. d. kaisl. Akad. d. 

Ssenschaften zu Wien 103 : 423-437. 1894. 


40 BOTANICAL GAZETTE [JANUARY 


significance that the pollen of so many species of plants germi- 
nate either in sugar solution or in water. According to Molisch* 
many forms germinate best in about Io per cent. sugar solu- 
tion, and concentrations of from 15 to 40 per cent. are often 
required. This might suggest an osmotic stimulus. Neverthe- 
less, there are many forms germinating well in water, and for 
other forms perhaps the idea of a special stimulus should be 
further applied to a study of pollen. Molisch indeed found 
minute quantities of malic acid a stimulus to the germination of 
Ericaceae; but, in general, my results indicate that it is only a 
stronger stimulus than pure water, not equal to sugar, for 
example. 

Lidforss’ has recently made extensive studies with pollen 
germination in ‘‘pure water,” and among many plants whose 
pollen is so adapted may be mentioned Orchidaceae, Salicaceae, 
and many Liliaceae, Geraniaceae, etc. Following in the path of 
still earlier workers along that line, Borcodin® clearly indicated 
that light has a stimulating effect for the germination of fern 
spores. Kny’ and others further determined the necessity of 
this factor. Heald® has more recently shown that this stimulus 
may find a substitute in high temperatures; and apparently he 
has also cleared up some previous inconsistencies. In the same 
paper, Heald has demonstrated the necessity of light for the 
germination of moss spores. In a nutrient solution containing 
peptone or sugar, on the other hand, good germination resulted 
in yi and higher temperatures alone had no stimulating 


OLISCH, H.: Zur Physiologie des Pollens mit besonderer Riicksicht auf die 
p laeatiaree Bewegungen der Pollenschlauche. Sitzb. d. kaisl. Akad. d. Wissen- 
schaften zu Wien 102': 423-427. 1893. 
5 LipForss, B.: Weitere Beitrage zur Biologie des Pollens. Jahrb. £. wiss. Bot. 32; 
237-312. 1899. 
° Boropin, —: Bull. de l’Acad. imp. de St. es 433-440 1867, (ref. 
Heald, 1. c.). 


7 Kny, L.: Beitrage zur Entwickelunsgeschichte der Pasnereuet: Jahr. f. wiss- 
Bot. 8: 1-15. 1877. 


* HEALD, F, pE F.: Gametophytic regeneration exhibited by mosses and condi- 
somo for the germination of cryptogam spores. Inaugural-Dissertation. Leipzig, 
1897. ; 


ee ee a 


Igor | GERMINATION OF SPORES 4I 


influence, as likewise small quantities of poisonous substances. 
The further interesting fact was disclosed that the effect of 
light is not one of photosynthesis. Equisetum spores, moreover 
according to Sadebeck,? germinate well either in light or in 
darkness. 

Notwithstanding the excellent culture methods for the fungi, 
a study of germination in relation to the stimuli involved has 
been largely a matter of incidental consideration. A study of 
growth phenomena, thermal limitations of growth, toxic effect, 
etc., have furnished some data relative to the stimulus of partic- 
ular substances. It seems, however, that no summation of the 
results has been made since the work of DeBary.” 


METHODS. 

The most convenient method of observing spore germination 
is undoubtedly the hanging drop-culture. The principal points 
to be considered in properly handling the drop-culture, or Van 
Tieghem cell, where nutrient media are employed, have been 
recently set forth clearly by Clark. These notes bear repeti- 
tion to a certain extent. The employment of large rings is 
desirable, and they should be cemented to the glass slips by a 
mixture of refined beeswax and pure vaseline. The cover 
should be cemented to the ring with vaseline. The same char- 
acter of liquid should be used at the bottom of the cell as 
employed in the drop. 

While the form of cell culture above described is highly 
accurate for culturable forms in nutrient media, it is by no means 
@ccurate in all other cases. When a careful study is to be made 
of particular stimulants in water, or in a medium not ordinarily 
Causing abundant germination, and the like, recourse should be 
had to a different method. My experience has been that the 
“ saanaig Re Ueber die Entwickelungsgeschichte d. Le U, S. W der 

e. Sitzungsber. d. Versammlung deutscher Naturforscher u. Aerzte zur 
amburg, 1876. : 

* DeBary, A.: Morphologie und Biologie der Pilze, 376-377. 1884. 


™ CLARK, J. F.: Dissociation and toxic effect. Journal of Physical Chemistry 3: 
263-316. 1899, 


42 BOTANICAL GAZETTE [JANUARY 


results may be untrustworthy if any volatile or soluble substance 
besides the medium employed is used in connection with the 
cultures. As subsequently mentioned, even the purest vaseline 
may have an effect on sensitive forms. In all cell cultures in 
which full nutrient media were not employed, I have used a 
modified method. The cells were used in small Petri dishes. 
On the bottom of the Petri dish was placed filter paper with 
holes made for the insertion of the cells, thus securing them 
against movement. The covers were laid on without vaseline, 
and only in a few cases with volatile substances was any vase- 
line placed on the outer rim of the Petri dish. Dishes with 
ground-glass tops are preferable. All cultures were kept in 
moist chambers. The one difficulty remains of opening the cul- 
tures for examination, but as it is done only once, or at most 
twice during the continuance of the experiment, it is perhaps a 
matter of small significance. 

All cells, dishes, flasks, etc., used in these experiments were 
first cleaned with an alkali, then an acid, and finally, after 
thorough washing in distilled water, steamed for an hour or two 
before use. Particular care was taken with the covers, which 
were also boiled in the cleaning materials, and kept soaking 
either in distilled water, or in some cleaning agent when not 
in use. Cultures were kept in the warm room at a tempera- 
ture of about 25° C., this temperature being especially 
commendable on account of ease of examination at the same 
temperature. 

After considerable experience with vegetable decoctions for 
the growth of fungi, I] have generally adopted a decoction of 
green string beans or of sugar beets as the best culture medium 
for most readily culturable fungi. Any absolute standard of 
strength is impossible, but as a working basis, fifty grams of dry 
matter for each liter of water has been found convenient; thus 
for green beans, from an average of analyses, three hundred and 
ninety-two grams would be required per liter. 

The chemicals used have been the purest obtainable; and the 
sugar was recrystallized by the alcohol method, and subsequently 


1901 | GERMINATION OF SPORES 43 


washed with ether. As a standard nutrient-salt solution the 
following well-known formula was adopted : 


Ammonium nitrate - - : - - 1.0 gram 
Acid potassic phosphate - - - Ob 2 
Magnesium sulfate - . - - Ak RG ee 
Iron sulfate - - - - - - Trace 
Cane sugar - - - - - - 3-5 grams, 
Water - - - -* too" 


As a standard salt solution the above formula has been used 
without the sugar, the osmotic influence being neglected as of 
little consequence in comparison with the desirability of having 
equivalent salt constituents. The experiments with the so-called 
paraffin water resulted from a test of the value of a paraffin 
lining for flasks in which distilled water was to be used or kept 
on hand. 

It is evident that so far as the fungi are culturable only 
pure cultures should be used for inoculation purposes. It is 
desirable, moreover, to avoid old cultures, and cultures which 
have been exposed to direct sunlight. When possible, I have 
used spores from cultures five to ten days old. 


EXPERIMENTAL. 

Table I will serve as the basis of some general comparisons 
with regard to the amount of germination on various nutrient 
media. In general a perfect food is the best stimulus for the 
germination of saprophytic forms, but in particular cases special 
stimuli are necessary. The standard organic solution has for 
germination purposes less strength than decoctions of plants. 
Generally speaking, the standard inorganic solution has about the 
value of sugar, except for the Mucoraceae. There is considerable 
difference, however, even with related species of fungi. 

Of the purely saprophytic fungi studied, Oedocephalum albidum 
is the only one capable of germinating to considerable extent on 
pure water. This one exception is suggestive in that it is not 
necessarily a characteristic of saprophytic fungi that the spores 
do not contain within themselves the nourishment required for 
8ermination. Botrytis, though parasitic at times, would further 


44 BOTANICAL GAZETTE ' [JANUARY 


TABLE I. 
PERCENTAGE OF GERMINATION. 
In all cases the cultures were examined 15 hours after the spores were sown, them 
in most cases 24 and 48 hours later. When germination was not complete at the end 


of 15 hours there was very my germination. The results as given, however, are for 
the second or third observation 


Culture media 
~— Wares Bean Standard | Standard Sug: 
decoction jnutr.saltsol.| inorg. sol, | solution 
Aspergillus flavus . .. . ot 100 100 75 75 
Sterigmatocystis nigra . fe) 100 100 3-75 20+ 
Penic ucu 100 100 20-50 Sie 
(Edocephalum albidum toot 100 100 50-75 100 
Botrytis vulgaris ot 100 100 100 100 100 
Mo ena. . 75 100 100 100 100 
Circinella umbellata o 100 fe) 5 i 
Mucor s as te ? 100 ie ahs 25 
cor racemosus : ?§ 100 100— ?§ 80 
Mucor spino fo) 100 oo* fe} oe 
Phycomyces of 100 50-100 fC) 2-10 
Chaetocladium Jonesii o 100 ) 80 
Coprinus fimetarius . oO 5-I0 o) ° ° 
Coprinus co ) ) fc) fo) oO 
oprinus micaceus fo) 100 oO fe) 0 
Boletus sp. nee ae fc) ) ) fe) sd 
summer. I os 
Ustilago perennans ans { pikes 50-70 100 ‘ 100f 
summer . 2-10 an Ais: ee 9 
Ustilago avenae { autumn .:| 50 root 
Ustilago striiformis . . . . ) as mre ae 25 
ocystis anemones : ) On oO - 
Uredo (P. graminis on wheat) 20 ee aay fs) 
Uredo (P. graminis on rye a) 2 deos 20 iy wis 50-100 
esac caryophillinus . . toot 7S Pen fel roof 
Exoascus s ; os *e ii et ac 
Galera primulina ‘ nae Ber 75 din xs 75+ 
* Abnormal. t Nearly. ** Budding. 
t Usually. § Small. . *** Some budding. 


confirm this view. With the exception of Coprinus micaceus, - 
most of the Hymenomycetes employed fail to germinate readily 
on bean decoction. Brefeld and others have determined that a 
large number of Coprini, Clavariae, and Tremellineae germinate 
well upon dung decoctions. 
Glycerin, although slow in action, often gives somewhat 
** Ductavx,—: Annales de l'Institut Pasteur 3: 112. —. 


TABLE II. 
PERCENTAGE OF GERMINATION, 
: Strength Strength Strength 
Spores of of solu- Peptone o Cane sugar} KH,PO, | NH,NO, MgSO, of KNO, 
tion % solution solution 
a 2H 
I 100 . 20 et 0 15-30 * 5-10 
nu n 
O.1 100 ~ - 20-75 at i 20 10-20 = 5-10 
Aspergillus flavus 0.01 50+ =~ 75-100 7 70-100 5-10 = 5-10 
0.001 10-25 = 50-75 >t 5-30 aes 
L — 30-50 : a 
ploy 100* . | 10-50 ?+ c ?t 
O.1 tot = 50-100 0 o ) 
Sterigmatocystis a ox & na 20-50 a . fi 
Ta 100 
- 0.001 ?f- — 5 0) re) 
\ ine 5 : 


* Nearly, when spores are single. 


t+ Very sma 


{At edges of drop. 


| 1065 


SHAOdS FO NOILVNIWLDZD 


46 BOTANICAL GAZETTE [JANUARY 


TABLE Il. 


PERCENTAGE OF GERMINATION IN GLYCERIN. 


: | 
ae Aspergillus ition Penicillium seaue ti pr ae Ustilago | Mucor 
tation flavus nigra glaucum alka vulga avenae_ |spinosus 

| 
fo | | 
= 75 fa) cae 100 75 50-90 fe) 
nu 
. 100* fe) ney 100 10-25 100* re) 
n 
oe 75-100 fe) Are 100* 25 100* fe) 
a 
- 20T 
* Nearly. + After two days. 


more germination than sugar, yet with Sterigmatocystis the case 
is reversed. In general, the difference in action has seemed to 
bear no particular relation to the physical properties of the 
substance, as for example its power of penetrating membranes. 

Most of the smut forms gave good germination on bean 
decoction, although less on pure distilled water than has been 
reported by observers using ordinary tap water. Uvomyces 
caryophillinus not only germinated less efficiently in bean decoc- 
tion than in distilled water, but in beet decoction it failed entirely 
to germinate. 

Of the three salts generally used in the standard adtriant 
salt solution, ammonium nitrate at a particular concentration 
gives abundant germination with Aspergillus flavus, but has no 
effect upon Sterigmatocystis. In general, the neutral salts give 
a greater stimulation than the one with acid properties. In this 
connection, reference should be made to the interesting results 
of Benecke.** He considers the presence of potassium absolutely 
necessary, and that without this metal no germination, or only 
traces of germination, can occur. Since his cultures were made 
in flasks, it is perhaps to be asked if he refers to germination 
(growth) visible to the unaided eye. Not only do some of the 
mold fungi germinate to considerable extent in solutions of 


BeNeECcKE, W.: Die zur Ernahrung der Schimmelpilze notwendigen Metalle. 


Jahrb. f. wiss. Bot. 28: 487-530. 1895. 


a 


Igor] GERMINATION OF SPORES 47 


simple substances, but the form of Botrytis which I have used 
germinates within a few hours on large surfaces of pure distilled 
water. Moreover, the potassium compounds alone are only very 
slightly stimulating for germination, and I have found no marked 
increase in germination of Phycomyces and Penicillium by the 
addition of potassium nitrate to a solutionof sugar. The above 
remarks are also partially applicable to the conclusions which 
Molisch** has drawn from his most interesting experiments with . 
Aspergillus and Penicillium. He found no germination without 
magnesium, saying: ‘‘da ohne Magnesium nicht einmal ein 
Auskeimen der Pilzsporen stattfindet, und dieses Element weder 
durch die Elemente der alkalischen Erden (Ca, Str, Bar) noch 
durch die der Zinkgruppe vertreten werden kann.” He also 
applied these results to all “lower” fungi, that is, apparently, to 
all culturable forms. 

Pepsin and asparagin gave almost no germination with 
Sterigmatocystis. The latter substance had also very little effect 
on Aspergillus flavus, while the percentage of germination with the 
former substance was as high as ninety. 

EFFECT OF SPECIAL STIMULI ON GERMINATION. 

Under the head of special stimuli, or substances which are 
not normal sources of food supply, are also included, for con- 
venience, certain carbon compounds, as well as the metallic salts 
and mineral acids. 

In this connection an unexpected result was obtained with 
cultures of Aspergillus flavus on distilled water which had been 
Standing in paraffin-lined flasks. Nearly all of the spores on 
the edge of the drop, or where single, germinated; and a large 
per cent. of germination occurred throughout the drop. In similar 
cultures Sterigmatocystis gave usually only 10 to 20 per cent. 
of germination. Penicillium and Phycomyces were not stimulated. 

Ethyl alcohol affords a marked stimulus for the germination 
of aspergillus; germination being more nearly perfect on the 
edge of the hanging drop, but occurring markedly throughout. 


ae: “ MOLIscH, H.: Die mineralische Nahrung der; niederen Pilze. oo mmsaaagie ss 
Kaisl. Akad. d, Wiss. zu Wien 103: 554-574. 


TABLE IV. 


PERCENTAGE OF GERMINATION. ~ 
d tren’th Stren’th : cl 
Phenol | Strychnin of, Ether of Camphor | Petroleum Vaseline re 
olution solution ae 
n ae 4 
aa ° 3-5 6 1-2 cone. 50 1-3 - 50-75 || ) 
Bee a z 
peo 80 1-3 on 2-5 = 10 2 50-75 || I 
n n I 
pron Og 24 = I is 10-25 “oh 50-75 || I 
u% So n I 
josoos | ae t Sa I pau 5-10 bees 50-75 || 0 
ie ee % 10-2 
e pee ea = ie . vee Se ee eee e 
ate ies ae ae ee conc. | 30-70 oe! dee ) 
nn nn Tr 
= fc) te) 4 oO. = ) t) ta) t) 
an : nm x 
ae ° sas e ng 0 8 5 0 
ah. - 8: oie - 
gree 1-5 0 on Co) — 0 BEE 25-50 || , 
an Co) co) ee oO soles ee rire casts sek 
TOO00O = 3 70000 ; 
eee eeee wee Bis: ie) ee ee wees eee ee ne oe 
760000 
{Small. _ - ${ Considerable at edges. § Irregular. || When single. 


ALLAZVI TVIINVLIOF 


AUvaNv!] 


oo a a ae 


Ce eee ae 


i901] GERMINATION OF SPORES 49 


Methyl] alcohol is slower in action, and eventually less effective. 
Immersion of spores for short periods of time in injurious con- 
centrations of ethyl alcohol, and subsequently sowing these 
spores in water cultures, afford no stimulus for germination. 

The results with phenol, here only partially given, were 
unusually variable. More than with any other substance used, 
difficulty was here experienced from the evaporation and changes 
in form,of the drop within the culture cell, which may partially 
account for the dissimilarity of results. 

Experiments with chloroform were not made in drop culture, 
but an exposure of half an hour in a saturated atmosphere was 
found fatal to Aspergillus and Phycomyces, and without bene- 
ficial effect upon Sterigmatocystis. 

Ether has proved of little consequence as a stimulus, except 
with Aspergillus at the lowest concentrations used. This was 
hardly to be expected in consideration of its action on the cell 
activities. It is to be noted that all of the above mentioned 
substances penetrate membranes immediately.*5 

At the end of fifteen hours there is an almost inappreciable 
stimulus with camphor, but its effect gradually increases to the 
third day. 

With Aspergillus flavus germination in pure water was 
increased 10 to 20 per cent. when vaseline was used for cement- 
ing the covers to the rings. 

The above experiments with Aspergillus on alcohol, camphor, 
strychnine, and vaseline were repeated in flask cultures. Again 
2 concentration of — alcohol gave more than 50 per cent. ger- 
mination; but under these conditions camphor, strychnine, and 
vaseline gave uniformly little or no germination. This, as wel 
as other experiments, suggested conditions in the hanging drop 
more favorable for germination than in flask culture. 

esides the results presented in Table V, an interesting fact 
is noted in connection with hydrochloric acid, as well as later 


“Overton, E.: Ueber die osmotischen Eigenschaften der lebenden Pflanzen- 
und Tierzelle. Vierteljahrsch. d. Naturf. Ges. in Zurich go: 1-43. 1895. 


Mo,Bot. Garden, 


TABLE: VY: 
PERCENTAGE OF GERMINATION AFTER THREE DAYS. 
Strength T Strength inl 
of HCl HNO, | Aceticacid artaric | Oxalic acid | Cu(NO of CuSO MaCiy | ZnSO 
solution : acid - solution % . FeSO, : 
atl re) fe) C) 
10 eeee I . 0 e] 
bd 
oe te) oO 15 5 te) fe) Ra § fe) 3 : 
” 
Aspergillus ae I 5-25 20 10 I-5 ft) 0.01 0-40 10-20 I-5 
avus n' 207% s o : ? 
a 20-3 I-2 15 25-50 5 4: 0.001 3-5 10-20 0 
x 
: C) 2-4 : 5-10 0.0001 25 30 
aes eres 0.00001| 3-10 
n 
= fo) 5 fe) 1.0 fe) 
— I- ) 10 I 
= 3 25f ) O.1 fC) ) Ce) 
Sterigma- cS = = ss a 
ferbitia —— 3 I-3 10-20 I-5 2-10 Co) 0.01 fC) ft) ) 
nigra n 
8 or 5-10 0 20 2-3 3-5 oO 0.001 15-30 >t 0 
ante 0 2-4 0 0.0001 0 ?+ 
es 0.00001 ft) 
* On edges. t Very small. { Throughout; 100 per cent. on the edges. 


of 


ALLAZVI TVIINVLIOF 


AUVANV!] 


leat. 


1901 | GERMINATION OF SPORES 51 


for some other substances. The spores of Aspergillus flavus 
readily fly off from the surface of the drop, and such spores, 
falling on the cover glass beyond the limits of the medium, but 
of course moistened by the slight water of evaporation, gave at 
nt n 
van and at ans 
The salts of the heavy metals have generally caused a slight 
increase in the germination of Aspergillus, but in no case have 
they acted very strongly. 
A repetition of the experiments with nitric acid, CuSO,, 


Cu(NO,), and FeSO, were also made in flask cultures at con- 
nw 


a germination of 50 to 70 per cent. 


centrations ranging from ae ee Nitric acid of ee 
> 


and below has in every case given a small percentage of germina- 
tion. The spores thus germinated show a considerable length of 
germ tube. Iron has had a similar but weaker effect. So far 
as could be ascertained from flask cultures, the copper com- 
pounds have at most caused a swelling of the spore, and in 
isolated cases the very slight protusion of a tube. 

Although the stimulating action of the organic acids here 
included, as also the action of alcohol, may be a very different 
one from that of the inorganic acids, yet this stimulus of the 
Organic acids is in no case a very remarkable one. As to 
whether or not these organic substances act as peculiar stimuli, 
Or as food substances direct, there is evidently no good clue, for 
it is not yet possible to draw the line between those concentra- 
tions which should be considered stimulating or poisonous and 
those at which there may be an action merely as food. Clark’s 


nm, 
results previously mentioned have shown that alcohol at — is the 


lowest concentration at which any inhibition of the germination 
of Aspergillus in nutrient media occurred. 

In Seneral, the action of the organic acids as food sub- 
Stances has been determined by means of the corresponding 
salts, and further than this we know very little about them. 

72 


The stimulus given by Joo OF less of acetic acid to both fungi 


52 BOTANICAL GAZETTE [JANUARY 


deserves mention, although the percentage of germination is not 
great. 

Oxalic acid stands quite alone as a substance stimulating 
Sterigmatocystis more than Aspergillus, in fact causing its 
maximum stimulus at a concentration which affords no germina- 
tion with Aspergillus. This effect on Sterigmatocystis was so 
variant and marked that this acid is to be regarded as a peculiar 
stimulant for that plant. 

: THE INFLUENCE OF CERTAIN PHYSICAL STIMULI. 

From some variations in results obtained it became evident 
that by the form of the drop and the amount of evaporation 
therefrom, or perhaps by some other physical force closely con- 
nected with these conditions, a considerable stimulus was given 
to germination. When the cultures of Aspergillus flavus were 
prepared at the same temperature at which they were to be 
incubated, with all possible precautions being observed as to 
purity of water and cleanliness of cover glasses, there was seldom 
more than a fraction of I per cent. of germination, provided the 
culture drop retained its original form and dimensions. In 
cases of some evident change of conditions within the culture, 
however, whether by slight dispersal of the drop or by a certain 
amount of evaporation, the percentage of germination was often 
greatly increased. In special cases the percentage of germina- 
tion was as high as go after an incubation of fifteen hours, this 
maximum being reached particularly when the spores were free 
from each other and collected on the periphery of the drop next 
to the glass. 

Many experiments were introduced with the hope of eliminat- 
ing the single factors which might be involved and of accurately 
determining the cause of the stimulus. Unfortunately the matter 
is as yet very inconclusive. 

If even very small amounts of some salts were present in the 
drop, these during evaporation would be concentrated at the 
periphery, especially at the edge in contact with the glass. 
This would hardly be sufficient, however, to account for the 


1901] : GERMINATION OF SPORES 53 


germination observed, and such an occurrence of salts could 
only come from the well-cleaned glass surface. 

As a check on the carefully prepared distilled water, cover 
glasses were moistened over a steam jet, and upon this condensa- 
tion the spores were sown as before, the results also being 
parallel to those previously obtained. 

The possible stimulus of rapid evaporation gave only nega- 
tive results in the following experiment: Properly cleaned 
ground-glass slips were placed on benches (glass rings) in Petri. 
dishes of distilled water. Strips of filter paper which had been 
soaked for days in acidulated water and then in distilled water 
were passed over the edges of the slips with the ends reaching 
into the water. A clear ground-glass surface of nearly an inch 
was left between the strips, and on this the spores were sown. 
This arrangement sufficed to keep a constant thin film of moist- 
ure over the glass from which evaporation might readily ensue. 
The covers of the Petri dishes were slightly raised, and these 
cultures were placed in a fairly dry atmosphere, in ordinary 
laboratory atmosphere, and in a moist chamber. After twenty- 
four hours there was from 10 to 20 per cent. of germination, 
and those cultures in the dry atmosphere gave perhaps less than 
the other two. Surface tension could hardly be considered a 
factor, for these fungi remained practically unchanged after a 
month on the surface of water in flask cultures. 

To test the effect of contact and surface tension with the 
Evaporation factor eliminated, well cleaned glass tubes were 
drawn out into a capillary end to be used as a culture cell. The 
tube was partially filled with water and the spores inserted. 
The water was then forced out until the spores reached capillary 
parts, when the larger end was also closed with water. Such 
cultures gave very slight germination after two days. 

Attempts to increase the surface tension by means of small 
quantities of oil in the water gave only negative results. Massart*° 
found surface tension productive of contact phenomena in 
. “Cte J.: La sensibilité tactile chez les organismes inférieures. Journ. 

- méd, et nat. de Bruxelles, December 1890 


54 BOTANICAL GAZETTE [JANUARY 


bacteria, amoebae, flagellates,etc. Busgen*? found that Botrytis 
reacts to surface tension by the formation of little bundles of 
branches perpendicular to the touched surface. 

Besides the previously mentioned contact experiments, 
recourse was also had to the clinostat. By shaking, spores 
were submerged in flasks containing a small amount of distilled 
water. These flasks were then rotated horizontally so that there 
was constant movement of the spores and considerable contact 
with the glass surface. The results were negative. Likewise 
experiments made with a shock-imparting clinostat, and also with 
a combination of the rotation and shock, failed to give any posi- 
tive results. 

A small number of experiments was made to test the effect 
of evaporation caused by a lowering of the vapor tension. To 
effect this the cells were arranged in the Petri dishes as before. 
Water was used in the hanging drop above, except in control 
experiments, and below was placed the salt solution of various 
strengths. Over solutions of MgSO,, varying in strength from 
$n to 3n, by far the best germination occurred over $x. where 
drying out was quite gradual. 

According to Lesage,** who made a number of experiments 
to determine the dampness of the air in which spores would 
germinate best, the spores of Penicillium germinate well at all 
of the higher densities, but reached the lower limit between 82 and 
84 per cent. humidity. Aspergillus germinates so well ordinarily 
at the higher humidities that the factor of evaporation seems of 
more significance. 

The action of a change of concentration of the medium was 
tested in a small way. Spores of Aspergillus flavus were sown 
in Erlenmeyer flasks containing respectively 20, 10, and 5 per 
cent. KNO, in one series, and in another the same strengths of 
MgSO,. After twenty-four hours about 5 to 10 per cent. of 
germination had occurred in the KNO, cultures and about 10 

‘7 BUSGEN, M.: Ueber einige Eigenschaften der Keimlinge parasitischer Pilze. 
Bot. Zeit. 51 : 53-72. 1893. 


**LesaGeE, P.: Recherches expérimentales sur la germination des spores du 
Penicillium glaucum. Ann. d. sci. nat. Bot. 8: 309-322. 1895. 


ew 


1901 | GERMINATION OF SPORES 55 


per cent. in the MgSO,. Water was then added to these cultures 
until the liquid was reduced tenfold in concentration. A second 
examination after the lapse of two days showed little or no 
increase of germination in the original 10 and 20 per cent. 
KNO,, but an increase to about 25 per cent. in the 5 per cent. 
solution, and a similar increase in the 5 and 10 per cent. MgSO,,. 
In the culture containing 20 per cent. MgSO,, the increased 
_ germination and the amount of growth was greater than in any 
other. 
EFFECTS OF TEMPERATURE AND OXYGEN SUPPLY. 


As previously mentioned, Heald found that fern spores kept 
at high temperature were incited to germination as by light, but 
a longer period of time was required and the response less uni- 
form. In general I have found very little difference between 
the germination in water of fungi at 25° C. and at temperatures 
nearer the maximum. At 32° C. there is some increase in the 
germination of Aspergillus flavus in hanging drop, but not in 
flask cultures. The former might well be due to other conditions 
than to any augmentation from the higher temperature. On 
nutrient solution Coprinus fimetarius was slightly benefited by the 
Same temperature. Such forms as Coprinus comatus, C. micaceus, 
Boletus sp., and Urocystis anemones could not be incited to germi- 
nation at higher temperatures when no germination occurred at 
25°. Botrytis and Phycomyces were both injured at the tem- 
perature of 32° C., Botrytis failing to germinate on water, and 
Phycomyces giving a slight growth at the bottom of the. liquid. 
Changes of temperature from 28° to 32° C. and vice versa did not 
materially affect the germination of Aspergillus and Sterig- 
Matocystis. After experiments of various kinds with the 
aecidiospores of Puccinia graminis, Eriksson and Henning’ found 
that the best results were secured by placing the spores for 
a time on melting ice, and then sowing them on water. The 
next best results were obtained when the spores were soaked in 
water at 3° C. for three hours, and then sown at room temperature. 


*9ERIKSSON and HENNING: Die Getreideroste, 71. 1896. Stockholm. 


56 BOTANICAL GAZETTE [JANUARY 


The poorest results were secured when the fresh spores were 
directly sown at the room temperature. Cooling also had a 
favorable influence upon the uredospores. This excessive cool- 
ing is hardly a natural stimulus. It may be regarded perhaps 
as a substitution stimulus, able effectually to. replace some other 
incitation of the natural environment. 

A few experiments were made with reduced oxygen supply, 
mainly to see if slight variations in this regard would at all 
vary the results. With cultures at room temperature with an 
air pressure of 60™™ there was no noticeable effect on germina- 
tion, either in water or in nutrient solution. At below 40™™ of 
air pressure there was marked retardation, but since such lower 
pressures were of little concern in these results, the matter was 
not carried farther. 


INHIBITION OF GERMINATION BY NUTRIENT SOLUTIONS. 


Various authors have made casual reference to the fact that 
ordinary nutrient solutions may injure the germination of certain 
fungi normally germinating in water alone. A thorough study 
of this matter should throw some light upon the conditions 
necessary for the penetration of the host plant by the parasite. 
As yet I have had opportunity to make but few experiments in 
this direction, but an accidental attempt to make a substitute 
for bean decoction by adding peptone to the standard nutrient 
salt solution gave some results of interest with certain smuts 
used. Ustilago Avenae and U. perennans gave but a small per 
cent. of germination on any solution containing 1 per cent. of 
a German preparation of peptone; and the pure peptone solu- 
tion gave only 1 or 2 per cent. of germination. An American 
manufacture of peptone did not act as an inhibiting agent; but 
the pure peptone solution afforded no better germination than 
distilled water. On the other hand, Ustilago was not inhibited 
by either preparation of peptone. 

Certain rust fungi also comport themselves sornewbial pecul- 
iarly towards nutrient solutions, as seen in Table VI. 


“~ 


i a 


LLL AAI OEE = 


1901 | GERMINATION OF SPORES 57 


TABLE VI. 


PERCENTAGE OF GERMINATION. 


PHO. | Pertone | gecrction | decoction | 22 tims | oom 
2 Solution | 5 ze 
Puccinia Helianth 
uredos spores) 1oo— 20 25 * 50 10 * " 
Uromyces caryophi 
linus (uredospores)| 100— 100— 0 g5° 1: 200 160 ne? 


* Germ tubes very short and often ill-formed. 


DeBary and others have noted an injurious effect of nutrient 
media upon the formation of zoospores in certain Peronosporeae. 
The effect is to suppress the amount of zoosporic germination, 
and to develop germ tubes instead. Wuthrich* found the same 
phenomenon characterizing the germination of Phytophthora 
infestans under the influence of small amounts of poisons. From 
experiments with Plasmopara viticola, | have found no germ tube 
development; but many nutrient media inhibit the germination 
of the species. Winogradski and Omeliansky** have determined 
that a number of organic compounds act even at considerable 
dilutions to hinder or prevent the normal action of the nitrite 
and nitrate bacteria, and often even to sterilize the solution in 
which it was attempted to grow these organisms. Among these 
inhibiting substances are peptone and other albuminoids, 
glycerin, salts of organic acids, and also ammonia. 

We are at this time far from a rational conception of the 
most important problems concerning the relation between host 
and parasite. The resistance of species and varieties cannot be 
viewed merely from a histological standpoint, and so far as the 
problem is capable of solution, a complicated set of factors is 
to be expected. If peptone and other nutrient media may be 
injurious to the a of certain fungi, not only the 

Woruricn, E.: Ueber die Einwirkung von Metallsalzen auf die Keimfahigkeit 
der Sporen Pune ais ae Pilze. Inaugural-Dissertation, Berne, 1592. 

WinocRaDski und OMELIANSKY: Ueber den Einfluss der organischen Sub- 
Stanzen auf die Arbeit der nitrifizierenden Mikroben. Centrlb. f. Backt. u. Parasitenk. 
5°:319-343, 371-387, 425-440. 18 


58 BOTANICAL GAZETTE | JANUARY 


poisonous excretions of plants, but all excretions may have their 
role to play regarding infection. At any rate, from the point 
of view of the fungus, a further study of chemotropism, of stimu- 
lants to germination, and of the inhibition of germination and 
growth by injurious substances cannot fail to lead us somewhat 
farther toward a knowledge of parasitic attack. 


RESTING STAGES AND DRYING-OUT OF SPORES. 

It is well known that seeds of certain phanerogams do not 
readily germinate on reaching maturity. From the researches 
of Weisner and others with rather peculiar phanerogams, it is 
seen that this resting period is not merely due to an absence of 
the best conditions, but that the element of time, as far as we 
know, is absolutely essential. Whether we may in some cases 
substitute for this element of time artificial changes of condition 
is mostly a matter of conjecture. 

The same remarks will hold in a general way pit the so-called 
resting stages of fungi. For the maturity of the spores of the 
Peronosporaceae and many of tlie teleutospores of the Uredineae, 
as examples, a certain resting period is indispensable. From 
the cytological studies of Wager,”? it would appear probable 
that in the case of Cystopus candidus the maturity of the oospores, 
so far as the external appearances are concerned, does not 
denote the maturity of the zoosporangium with reference to the 
full quota of nuclei as a basis for the formation of zoospores. 
It appears that a resting stage must intervene before the final 
divisions of the nuclei. 

Teleutospores of Puccinia graminis germinate best when they 
have been subjected to all the changes of the winter months. 
Observations on the presence of a certain coloring matter in the 
walls of such spores led Dietel?3 to attribute to this certain prop- 
erties for the prevention of the germination, and for protection 
against unfavorable conditions. DeBary** has found that the 
teleutospores of the previous harvest cannot be brought to germi- 

7? WaGER, H.: Reproduction of Cystopus candidus. Ann. Bot. 10: 245-339: 

"3 DIETEL, P.: Flora 74: 151. 1891. 24 DEBaryY, A.: lL. c. 


Re ee ee, er 


Igor] GERMINATION OF SPORES 59 


nation later than August of the following year, and the optimum 
germination occurs during the spring of the latter year. Eriksson 
and Henning’ are of the opinion that a passage through the 
animal body will not act as a substitute for the dormant period, 
Such lines of work have not been systematically followed out, and 
even for the Uredineae, which are very variable in their disposi- 
tion toward the resting period, the limitations are not sufficiently 
known. According to Kihn, Brefeld, and others, as previously 
mentioned, germination of certain smuts in water can only be 
induced after a period of rest, while immediate germination may 
result from the addition of food material. My own results also 
show that the per cent. of germination with Ustilago Avenae and 
U. perennans increases considerably from summer to autumn, 
even though the material is kept in dry condition. 

In other experiments, Sterigmatocystis nigra kept dry in the 
laboratory for five years gave no germination, while material one 
and two years old gave good germination in nutrient media. 
This age of the material did not act asa stimulus to germina- 
tion, however, since only a fraction of 1 per cent. germinated 
when sown on distilled water. After drying out for four days 
on slide at 25° C. neither Aspergillus flavus, Penicillium glaucum, 
nor Sterigmatocystis nigra. showed any germination as tested by 
distilled water. 

SUBMERGENCE OF SPORES. 

Ordinarily the spores of such molds as Aspergillus flavus and 
Sterigmatocystis nigra float on the surface of solutions; and, 
depending upon the solution, they may or may not tend to collect 
at the line of attachment to the glass in drop cultures. The 
spores of Phycomyces and other Phycomycetes, as well as many 
Hymenomycetes, however, very readily sink beneath the surface. 
To test the capacity of Aspergillus for submerged germination, 
Spores were sown in bean infusion between the parts of a strip of 


mica lightly separated. The mica was then dropped into a flask 


of bean decoction. After two days the removal and examina- 
tion of the mica showed that all spores were germinated, those 
*S ERIKSSON and HENNING: Op. c. 54. 


60 BOTANICAL GAZETTE [JANUARY 


in the middle, however, having developed a germ tube only 
about ten times the diameter of the spore, while those on the 
margin were growing luxuriantly. Under pressure submerged 
spores do not germinate. 

Spores of Aspergillus flavus were also sown on a layer of agar 
beneath a considerable layer of the same material, the upper 
layer being poured on while the first was still soft. Germina- 
tion readily resulted. Also spores in flask cultures of ¢%, alcohol, 
submerged by shaking, germinated readily. 


SOME PECULIARITIES OF GERMINATION VERSUS GROWTH. 


The Hymenomycetes will doubtless form an interesting field 
for the study of germination relative to special stimuli. With 
studies which are yet merely preliminary I have secured only a 
single positive result of interest, but in many cases failures are 
likewise suggestive. 

Brefeld, in his Untersuchungen iiber Schimmelpilze (part III), 
records that spores of Gasteromycetes and of Phallus (p. 174) 
particularly, as well as other members of the fleshy fungi, do 
not germinate under any conditions tried. On the other hand, 
Coprinus stercorarius (p. 14), C. lagopus (p. 99), and C. ephemer- 
oides (p. 117) germinate well on any plant decoction, as likewise 
spores of Clavariae and Tremellinae(p. 181). Coprinus ephemerus 
(p. 109) is said to germinate once in perhaps ten trials. While 
I have not been able to germinate Coprinus comatus and Boletus 
sp., these forms have been studied only from fresh spores and from 
spores keptin the laboratory about two months. Coprinus fimeta- 
rius has given various small percentages of germination in different 
vegetable decoctions, but otherwise no germination. A species, 
which unfortunately was not determined while fresh, but later 
identified as Coprinus micaceus, has given little or no germination 
in all solutions containing no plant decoction. Inbeanand dung 
decoction the same material has furnished perfect germination. 
The question then of interest was to determine if there might 
exist in the bean decoction a substance stimulating germination 
but unnecessary for growth; in other words, if we may here 


a ell 


SRE 


1901] GERMINATION OF SPORES 61 


distinguish between a medium for germination and a growth 
medium. Spores of this fungus, caught with all possible sterili- 
zation precautions, were germinated on bean decoctions in flask 
cultures. About eight hours after the sowing, the liquid was 
filtered off in a sterile filter, and the collected mass of germi- 
nated spores was removed by a needle to a second filter. Here 
the spores were washed, and finally transferred to a flask of 
sterile water. In the latter they remained two days, the water 
being then poured off and the standard nutrient salt solution 
added. Growth proceeded gradually, and at the end of one 
month there was a thick mat covering the bottom of the flask, 
as if with a circular piece of canton flannel. With all of the pre- 
cautions observed, and by a comparison of the mycelium, this must 
be taken to justify the belief that we may here deal with a case in 
which a medium failing to stimulate to germination may yet 
afford growth. Bean decoction, moreover, is a better growth 
medium, and it would seem that the stimulus to germination 
would be a food stimulus. Nevertheless, the addition of pep- 
tone to the standard nutrient salt solution also gave no germina- 
tion, and if the stimulus is that of a food, it must be considered 
in the class of peculiar foods. 

Some other results, scarcely comparable to the above, may, 
however, be mentioned at this place. Janczewski* has deter- 
mined that Ascobolus furfuraceus, a plant growing normally on the 
dung of herbivorous animals, could only be prepared for germi- 
nation by being passed through the digestive tract of such ani- 
mals. White rabbits were the animals used in his experiments. 
We have here evidently a case in which the spore is immediately 
capable of germination provided it may be first acted upon chemi- 
cally or otherwise, so that it is rendered capable of using the 
stimulus of the medium on which it normally grows. 

DeBary” also found that Onygena corvina, growing on the 
feathers of birds of prey, seemed to require a particular stimulus 

*6 JANCZEWSKI : Morphologische Untersuchungen iiber Ascobolus furfuraceus. 
Bot. Zeit. 29: 257-262, 1870. 

” DEBary, A.: op. c. pp. 376-377. 


62 BOTANICAL GAZETTE [JANUARY 


of its normal environment in order that germination might be 
effected. 

Coprinus comatus, Boletus sp., and a few other forms failed 
also to germinate on filtrates or decoctions of the soil in which 
the plants grew. The soil filtrate cultures were of course swarm- 
ing with the bacteria which would thrive under such conditions. 
Equally futile have been the attempts to germinate these spores 
in the presence of alkaline substances, in a slightly acid medium, 
or in the presence of a reducing agent. Hartig* mentions the 
germination of Merulius lacrimans in the presence of ammonium 
and other alkaline compounds, after the failure of many other 
substances. 

Brefeld® found that Zil/etia caries fails to germinate in 
nutrient solution. If already germinated in water and then 
transferred to nutrient solution the death and bursting of the 
promycelium soon occurs. 


DILUTION OF FOOD MATERIALS. 


Concerning the minimum food supply necessary for more or 
less perfect germination there seem to be almost no references 
in the literature. The concentration, however, at which some 
substances begin to attract chemotropically, or practically this 
lower concentration, has been determined by Miyoshi.*° For 
example, cane sugar at 0.01 per cent. attracted the hyphae of Mucor 
stolonifer, and ammonium nitrate attracted the same fungus at 
0.05 per cent.; while meat extract of 0.005 per cent. was attractive 
for Saprolegnia. On the other hand, Eschenhagen* and others 
have found that germination and growth of the mold fungi may 
occur at very high concentrations. 

Examining horizontal lines in table VII, these experiments 
are more or less comparable from the point of view of the con- 
centrations of the medium. The standard nutrient salt solution 

7 HARTIG, R.: Der echte Hausschwamm. 

*° BREFELD, O.: Vgl. Unters. a. d. Gesammtgebeit der Mykologie. Part V, p. 152- 

»® MIYOSHE, M.: Ueber Chemotropismus der Pilze. Bot. Zeit. 52: 1-28. 1894- 

% ESCHENHAGEN, F.: Ueber den Einfluss von Lésungen verschiedener Concen- 
tration auf das Wachstum von Schimmelpilzen. Inaug.-Dissertation, Leipzig. 1889- 


1901] GERMINATION OF SPORES 63 


TABLE VII. 


Bean decoction Standard nutr.-salt solution Sugar solution 
Strength | Per cent. Per cent. Strength) Per cent, | 
of of germi- Remarks of germi- Remarks oO of germi- Remarks 
solution nation nation solution | nation 
. 
( | = 20 From 10-30% 
Standard | 100 100 | lad 20 FON GET 
Io 
n 
a 100 100 aw 75 Drops _ poor, 
- spreading 
n 
ian 100* 65+ Jooo | 10-50 | Varying from 
middle to 
Asper- bins 
gillus ¢ ‘ : : od 
fax fae 60+ | From 40-90% ?{ (3% in middle}, 5 5-20 “6 
ay in different of drop to 
s cultures 20% on edges 
10000 20+ | From 5-40% ?f{ + |1% in middle 
in different of drop to 
s cultures 20%0n edges 
Aa >t ?7~ (Slight germi- 
nation on 
L edges only 
eee bs cee: | 
( Sd 5 | Very few in 
" middle 
Standard | 100 100 oad 20-75 | Varied from 
center to 
fs S 
s 
Sterig- - 100 100 == | 10-50 ‘ 
mato- ee n 
cystis a re 20+ | Varied from pena 5-10 Sythe 
nigra 10-50% 
about 60% 
itil % 5-10 “or ‘ce ee 
1000 rca 225 I 
; s 
. 10000 0|| te) 
S 
L | To0000 ° fo) 
a 
* ‘ 
Nearly, + Very small. t Small. || Practically. 


contains 5 per cent. of sugar, the beet decoction about 3 per cent., 
nn * 

and the Fo Sugar solution 3.4 per cent. It is noticeable that 

with Aspergillus on bean — practically normal germina- 


| tion takes place as low as —— (standard solution diluted one 


64 BOTANICAL GAZETTE [JANUARY 


hundred times), and otherwise the coefficient of maximum ger- 
S 

mination is constantly above sca Parallel with the results 

cited above, flask cultures were made with the more variable 

Aspergillus. These cultures for the first two media showed at 


s he s : 
s and — complete germination, at Fen VOY good germi- 
Io 


: Ss naa 

nation, at ety small per cent., and at further dilutions 
1°) 

none. Equivalent cultures on sugar solution are interesting, 


ee mre n 5 
> Siving almost no germination; co nae Per cent. ; ae 


nn 


and perhaps about Io per cent.; and a very small 


I9000 To000 
percentage. From this it will be seen that a tenfold dilution of 
the ordinary culture media affords perfect germination, and a 
dilution below one thousand times gives practically no germina- 
tion except with very sensitive fungi. 


INDIVIDUAL VARIATION OF SPORES. 

No studies of importance seem to have been made upon the 
variation in capacity for germination of individual spores pro- 
duced under similar conditions, or of spores from the same 
conidiophore or sporangium. Nevertheless, great individual 
differences exist, and in any medium which is not a strong 
stimulus for germination, varying percentages of perfect germi- 
nation will invariably occur, whatever precautions of method 
may be observed. 


CAPACITY FOR GERMINATION OF SPORES LONG INCUBATED ON 
WATER SURFACES. 

Spores of Aspergillus flavus and Sterigmatocystis nigra were 
sown on distilled water in Erlenmeyer flasks. At the end of 10, 
30, and go days some of these were transferred to bean decoc- 
tion. Practically no germination had occurred on the water; 
and in nutrient solution of the new cultures these spores gave 
perfect germination. After 125 days, some germination had 
occurred among spores in masses, but this germination was by 


1901] GERMINATION OF SPORES 65 


no means general. The ungerminated spores of these fungi 
were entirely uninjured as to their capacity for germination after 
this period of incubation. 


LENGTH OF LIFE OF GERMINATED SPORES IN WATER. 


Spores of Aspergillus flavus and Botrytis vulgaris were sown in 
weak bean decoction, and after about eight hours, or when all 
had germinated, they were filtered and thoroughly washed. The 
masses of spores were then transferred to fresh distilled water 
and so preserved. The last sowing from these germinated 
spores was made after eighty days with Aspergillus, and at this 
time all were yet alive. With Botrytis, the last inoculation was 
made after forty days, with the same result. In the cultures in 
which these spores were tested, general growth alone was not 
depended upon, but individual germ tubes were located and 
growth from these directly observed. 


LENGTH OF LIFE OF SPORES DRIED OUT AFTER GERMINATION. 


Spores of Aspergillus and Botrytis were germinated as in 
the above experiments, and then dried on filter paper. With 
Botrytis no further growth could be secured from spores thus 
dried after twenty-four hours, thus in perfect agreement with 
the results of Nordhausen.32. On the other hand, inoculations 
from the Aspergillus material and careful marking and observa- 
tion of individual germinated spores gave a very general new 
Srowth after being dried out twenty days. After sixty-five 
days there was new growth from about half of those transferred, 
and after one hundred days there was no sign of growth from 
germinated spores. It would be interesting to compare further 
the behavior of parasitic and saprophytic forms in this respect. 
We know in a general way that the germ tubes of parasitic forms 
die quickly when dried. Indeed in an early paper by Hoff- 
mann it is stated that. “ Austrocknen im gekeimten Zustande, 


* NORDHAUSEN, M.: Beitrage zur parisitirer Pilze. Jahrb. f. wiss. Bot. 33 : 1-46. 


‘“SHoFrMANN, H.: Untersuchungen iiber die Keimung der Pilzsporen. Jahrb. 
f. wiss. Bot. 2: 267-337. 1860. 


66 BOTANICAL GAZETTE [JANUARY 


also Austrocknung ‘des Keimfadens, fiir das Weiterwachsen 
absolut tédlich ist.” 

I am pleased to express my thanks to Professor Pfeffer for 
his kind advice and encouragement in the progress of this work 
completed in his laboratory at Leipsic. 


CoRNELL UNIVERSITY, 
Ithaca, N. Y. 


CURRENT LITERAL Ue: 
BOOK REVIEWS. 
Agricultural botany." 


THOSE practically interested in the cultivation of plants have felt the 
need of a text-book of botany adapted to their wants. They are interested 
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of the practical cultivator. The appearance of a text-book definitely designed 
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hat is equally to the point, he is evidently in touch with what is best 
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This book is divided into eight parts. The first deals with the gross 
Structures of seed plants, the topics being seeds, the root, the vegetative shoot, 
the leaf, the flower, the inflorescence, and the fruit. The account is clear and 
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: The third part is devoted to plant physiology, and it is a pleasure to meet 

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a CIVAL, JOHN: Agricultural Botany, theoretical and practical. 8vo. pp. 

xi + 798, illustrated. New York: Henry Holt & Co. 1900. $2.50. 

Igor] 67 


68 BOTANICAL GAZETTE [JANUARY 


worth mentioning is that little idea is given of the plant as an irritable 
organism, irritability being scarcely alluded to except in connection with 
movements. e subjects treated are the chemical composition of plants, 
osmosis, absorption of water, transpiration, absorption of food materials, 
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the author in the main steers clear of the usual misleading terminology, but 
evidently finds it impossible to make clear the real situation without more 
training than the book demands. 

The fourth part has to do with the “classification and special botany of 
farm crops.’ After a general discussion of the classification of plants, in 
which the emphasis is naturally laid upon seed plants, the author selects for 
special presentation the following families: Cannabaceae, Chenopodiaceae, 
Cruciferae, Rosaceae, Leguminosae, Umbelliferae, Solanaceae, Compositae, 
and Gramineae, with special chapters on cultivated and wild oats, cultivated 
barleys, rye, wheats, grasses, and clovers. The fifth part discusses the weeds 
of the farm. A general discussion of the injurious effects of weeds, their 
duration, their habit of growth, how they spread, and their extermination, is 
followed by an account of special weeds. 

The sixth part presents the subject of farm seeds, the purity, germination 
capacity, germination energy, weight, form, color, etc., of seeds being discussed. 
The seventh part considers fungi chiefly in relation to some common diseases 
of plants. After a presentation of the structure and habits of fungi in gen- 
eral, the different diseases which they produce in farm crops are discussed. 
The last part is devoted toa consideration of bacteria. The morphology 
and reproduction of bacteria are first presented, and afterwards their work 
in lactic, butyric, and acetic fermentations, and in putrefaction, nitrification, 
denitrification, fixation of free nitrogen, and diseases of animals. 

It will be observed that the scope of the book is broad, and that it includes 
the subjects of special interest to cultivators of plants. It certainly deserves 
to be received Bide by agricultural schools and those engaged in agricul- 
tural pursuits.— J. M. C. 

A botanical dictionary. 

THE making of a good dictionary, even though limited to the special 
vocabulary of a single science, is not the easy task a novice might imagine. 
It is fortunate, therefore, that the task was undertaken by so competent 2 
person as Mr. B. Daydon Jackson, who, if we may judge by his works, delights 
in painstaking labor of a kind that is intolerable drudgery to most men. e 
fully realizes the difficulties of the present task and the meager appreciation 
it is likely to receive, for he aptly quotes Dr. Johnson: ‘Every other author 
may aspire to praise, the lexicographer can only hope to escape reproach.” 


—_— 


1901] CURRENT LITERATURE 69 


It will be only just, therefore, to express at once our hearty commenda- 
tion of the work which Mr. Jackson has done, and to say that the glossary, as 
he modestly calls it,? is not only by far the largest and most comprehensive 
botanical dictionary in the English language, but by far the best. Concise 
definitions, brief derivations, and the accents are given for almost 15,000 
words, which is about three times the number in Crozier’s hastily compiled 
dictionary, issued a few years ago in this country. The only English botani- 
cal dictionaries are long since out of date and practically useless. 

Mr. Jackson has succeeded remarkably well in traversing the whole 
range of our terminology. Even very new words have not escaped him, for 
he includes such terms as edaphic, tropophyte, geophyte, coenocentrum, com- 
pound oosphere, etc. Most of the definitions are concise an good ; some, how- 
ever, are incomplete, e. g., (ree, xerophyte , some are careless or ambiguous, 
é. £., coenocentrum, mycorhiza (misspelled mycorrhiza), geophyte ; and some 
are antiquated or erroneous, ¢. g., archesforial cells, chlorovaporization, 
oogenesis, fertilization, sperm cell, etc. The accent given does not always 
coincide with lexicons, ¢. g., medillary and eléter. In the former case it 
does not coincide with usage in this country though the author says 
medullary is the usage in England. And if usage even permits eldter, it 
violates all rules of quantity. 

Part, perhaps a large part, of the faults are due to the extreme conden- 
sation of the definitions. This might have been avoided, without making the 
volume of inconvenient size, by dispensing with a number of words derived 
from Crozier’s dictionary, which, as inquiry and search indicate, neither have 
nor have had botanical use in literature, ¢. ia gusset, ensate, hydroid, polydel- 
Phous, secondine, etc. Greater fullness of the definitions might also have 
been compensated for by using thinner paper, instead of the thick and stiff 
Stock, which, together with the stiff binding (entirely unfit for such a refer- 
ence book), prevents the book from opening comfortably or lying open. 
But blemishes suchas these, the more noticeable because so easily avoidable, 
may well be overlooked in view of the good qualities, too many to enumerate, 
which distinguish this book from its predecessors. Every laboratory needs 
a Copy on its shelves.—C. R. B. 


MINOR NOTICES. 

THE FouRTH PART of the “ Catalogue of Welwitsch’s African plants,” 
by W. P. Hiern, and published by the British Museum, has just appeared, 
including Lentibulariacee to Ceratophyllee. The occasion of the publica- 
tion was stated in the review of the first part, published in the GAZETTE (23: 
210. 1897). The present part contains some important families, as Acantha- 
ceae, Verbenaceae, Labiatae, Euphorbiaceae, etc. Approximately ninety 
~ *JACKsoN, BENJAMIN Daypon: A glossary of botanic terms, with their deriva- 
LB. accent. 12mo. pp. xii-+ 327. London: Duckworth & Co. Philadelphia : 
. ippincott Co. 1900. 


7° ; BOTANICAL GAZETTE [JANUARY 


new species are described, about one third of which are Labiatae; and to 
the same family the single new genus (Sy#phostemon) belongs.—J. M. C 

THE SERIES OF PUBLICATIONS by de Wildeman and Durand, setting forth 
the flora of Congo, has now reached the completion of the second fas- 
cicle.3 In this fascicle it is stated that the number of spermatophytes, which 
a year ago was estimated to reach 1500, has now reached 2000. The fascicle 
deals with collections made by Belgian explorers, officers, and missionaries, 
and contains the descriptions of about twenty-five new species.—J. M. C 

THE FIRST VOLUME of Wiesner’s Die Rohstoffe des Pflanzenreiches \s 
now completed by the publication of the fifth part. The subject of yeasts is 
completed; products of alge and lichens are elaborated by Dr. F. Krasser 
(31 pp.), of which the most important is agar, and carrageen or “ Irish moss," 
dye-yielding lichens, “ Iceland moss,” and “reindeer moss.’ Dr. W. Figdor 
treats the galls (27 pp.), and Dr. F. von Héhnel the barks (96 pp.), with which 
important section the volume concludes.—C. R. B. 

Dr. H. A. HARDING, of the N. Y. Agricultural Experiment Station at 
Geneva, has found that the black rot of cabbage and allied plants is a wide- 
spread disease in Europe,’ which has been overlooked by continental pathol- 
ogists. He observed it in the vicinity of Versailles, Bern, Ziirich, Karlsruhe, 
Bonn, Harlem, Fulda, Halle a. S., Berlin, Kiel and Slagelse in the course of 
some months of travel in France, Germany, Switzerland, Belgium, Holland, 
and Denmark in the autumnof 1898. Only in Switzerland and Denmark did 
the disease seem to be of economic importanc B. 

THE Proceedings of the Indiana pre of Science for 1889, just 
received, en the following botanical papers : “Contributions to the flora 
of Indiana,” and “Some unrecognized forms of native trees’ (Asimina tri- 
loba, Jet nigra, Liriodendron Tulipifera, and Disopyros Virginiana), by 
STANLEY COULTER; “A list of plants collected at Cedar, Shriner, and 
Round Lakes,” by C. C. DeaM; “The resin ducts and strengthening cells of 
Abies and Picea (illustrated), by HERMAN B, Dorner; “A proteolytic 
enzyme of yeast,” and ‘Saccharomyces anomalus,” by KATHERINE E. 
GOLDEN; “Some problems in Corallorhiza,” and “The disappearance of 
Sedum ternatum,” by M. B. THomas.—J. M. C 


3 WILDEMAN, Em. DE ET DurRAND, Tu.: Annales du Musée du Congo. Bota- 
nique, série II. Contributions a la flore du Congo. Tome I. Fasc. 2. (2™* Partie.) 
4to. pp. ete Bruxelles: Charles Vande Weghe. July 1900. 

4 WIESNER, JULIUS: Die Rohstoffe des Pflanzenreiches. Versuch einer tech- 
nischen westatcllian tea 4s Picuscaae oak Ed. 2. Lief. 5. pp. 641-795, figs. 123-153: 
Leipzig: Wm. Engelmann. 1900. 

5 Die schwarze Faulniss des Kohls und verwandter Pflanzen, eine in Europa weit 
verbreitete bakterielle Pflanzenkrankheit. Centralbl. f. Bakt. II. 6: 305-313- P/- 2 
and map. 1900. 


1901] CURRENT LITERATURE 71 


NOTES FOR STUDENTS. 


IN THE Gardener's Chronicle for November to last, Mr. W. G. Smith 
figures a curious malformation on the pileus of Agaricus albus, recently col- 
lected and now in the British Museum. On the lamellar surface of the 
obconical pileus there are five smaller pilei, each raised on a short stipe, 
arising almost in the same plane, midway from stipe to margin.—C. R. B 

CZAPEK’s recent paper ® upon the sensitiveness of the root tip to geotropic 
influence will be read with exceptional interest by all who have attempted to 
repeat, for class demonstration or otherwise, his ingenious experiments with 
bent glass caps. There appeared last year, in Russian, a paper by Wachtel? 
containing a mass of experimental evidence which the writer believed to dis- 
prove Czapek’s thesis entirely. Wachtel failed to obtain the curvatures 
described by Czapek for capped roots, and he obtained other curvatures 
which seemed to prove the absence of any localization of the sensitive region 
in such organs. Czapek has retraversed the whole disputed ground in his 
usual careful manner. He was able to reproduce all of Wachtel’s results 
and shows that they were due to imperfect manipulation in the preparation 
of the glass caps. The walls of Wachtel’s capillary tubes were too thin, 
thus bringing about either a narrowing of the bore at the angle, or the pro- 
duction of an angle too blunt for the desired purpose. Either of these con- 
ditions prevents the success of the experiment. The present article contains 
a translation into German of a good part of Wachtel’s paper, with critical 
remarks, and a discussion of the difficulties of manipulation, including a very 
detailed account of Czapek’s own methods. 

There is also added a new experimental proof of the fact that only the 
formative region of the root tip is sensitive to the earth’s gravitation. 
Briefly, this is as follows: Vertically placed roots are allowed to grow into 
right- -angled caps in the usual way; then the caps are removed, the seedlings 
placed upon the revolving klinostat, and their behavior is observed. Within 
a few hours a bending becomes evident, the after effect of the stimulus 
received by the stationary horizontal tip while still within the cap. Various 
Positions of the roots were tried and all point to the same conclusion. The 
new method is even more elegant than the older one, and in itself amounts to 
a proof of Darwin’s hypothesis of the localization of the sensitive region .,— 
Burton Epwarb LIvINGsTON. 


°CZAPEK, FRIEDRICH : Ueber den Nachweis mee oe Sensibilitat der 
Wurzelspitze. Jahrbiicher fiir wiss. Bot. 35: 313. 19 


WacutTeL, M.: Zur Frage iiber den Geotropismus der Wurzeln. Berichte der 
neurussischen iS leach der Naturforscher in Odessa 23 : 48. 1599. 


NEWS. 


Dr. A. NESTLER has been appointed assistant professor in the German 
University at Prag. 

Dr. R. HEGLER, frivat-docent in the University of Rostock, died at 
Stuttgart on September 2 

Sir Jos—EPH HooKER has been elected one of the foreign associates of 
the French Academy of Sciences, Institute of France.—Gard. Chron. 

THE SECOND MEETING of the naturalists of the central states was held 
at Chicago, on December 27 and 28, in the biological buildings of the Uni- 
versity of Chicago. There was a good attendance of both zoologists and 


and Holferty of Illinois, Mottier and King of Indiana, Grover of Ohio, Cope- 
land of West Virginia, and Jeffrey of Toronto. The visitors, together with 
the botanical staff of the University of Chicago, and the ten or twelve graduate 
students who were still at the University during the holidays, made upa satis- 
factory botanical attendance. The papers were numerous enough to demand 
the division on Friday afternoon into botanical and zoological sections. The 
botanical papers were as follows: H. C. CowLes, H. N. WHITFORD, and C. C. 
ApAms: The relation of base-leveling to specific differentiation. A symp0- 
sium, treated from the standpoint of both plants and animals. B.M. DAvIs: 
Coenogametes. H. G. TIMBERLAKE: Swarmspore formation in Hydro- 
dictyon. . H. SHULL: Variation in the florets and bracts of Aster Puniceus 
Linn. H.N. WuitrorD: The genetic development of the forests of nor- 
thern Michigan and a consideration of the factors controlling the distribution 
of ‘coniferous forests. E.C. JEFFREY: An anatomical classification of vas- 
cular plants. H.C. CowLes: Application of the quantitative method to the 
dynamical study of plant societies. R. B. THomson: North American 
Chalazogams. O. W. CALDWELL: The use of color photography in demon- 
stration work, J. H. FAULL: Some interesting features in the anatomy of 
the Osmundaceae. E. B. CopELAND: Stomata of some Liliaceae. The 
success of this meeting justifies the expectation that a society of the naturalists 
of the upper Mississippi valley meeting at this time of year, can be made 
successful. A third meeting, at which a formal organization may be effected, 
will be held in Chicago in December rgor. 

72 [JANUARY, 190! 


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Vol. XXXI IPEBRUARY, 1901 | No. 2 


THE 


BOTANICAL GAZETTE 


EDITORS 
JOHN M. COULTER anp CHARLES R. BARNES, 


WITH OTHER MEMBERS OF THE BOTANICAL STAFF 
OF THE UNIVERSITY OF CHICAGO 


ASSOCIATE EDITORS 


J. C. ARTHUR 
Purdue University 
CASIMIR DeCANDOLLE 
Geneva 


J. B. DeTONI 
University of Padua 
ADOLF ENGLER 
University of Berlin 
LEON oe 
L’ Ecole de Pharmacie, Paris 
ROBERT A. HARP 
Oniversity of Wisconsin 
JINZG MATSUMURA 
Lmperial Oniversity, Tokyo 


FRITZ NOLL” 
University we Bonn 
VOLNEY M. scene 
University - Wished 
ROLAND THAXTER 
flarvard pws 
WILLIAM TRELEA 
Missouri font Garden 
H. MARSHALL W 
Un cide =r Cambridge 
EUGEN. WARMING 
University of Copenhagen 
VEIT wap do eae 
oval aor as Setences 
Stock 


CHICAGO, ILLINOIS ; 
ecaksme by the Giniversity of Pe. 


ete @niversity of fhicage Press 


corvRiGHt Pe BY THE A oem vein aatade! oF CHICAGO See 4: 


ii aaa 


e 
ee 


Botanical Gazette 


A Montbly Fournal Embracing all Departments of Botanical Science 
Subscription per year, $4.00 Single Numbers, 40 Cents 
The subscription price must be paid in advance. No numbers are sent after the expiration 
e time paid for. No reduction is made to dealers or agents 


FOREIGN AGENTS: 


Great Britain—-W™M. WestEY & Son, 28 Essex Germany — GEBRUDER BORNTRAEGER, Berlin 
St., Strand, London. 18 Shillings. SW. 46, Schonebergerstr. 17a. 18 Marks 
Vol. XXXI, No. 2 Issued February 23, 1901 


CONTENTS 


THE Bestia aanrni ECOLOGY OF CHICAGO AND VICINITY ; A STUDY OF THE 
ORIGIN, DEVELOPMENT, AND CLASSIFICATION OF PLANT SOCIETIES. Con- 
TRIBUTIONS FROM THE Hutt BoTANICAL LABORATORY. XXIV on THIRTY-FIVE 


FIGURES). Henry Chandler Cowles - 73 
ea hare PLANTS FROM GUATEMALA AND OTHER CENTRAL AMERICAN 
BLICS. XXII (witH pLaTE 1). John Donnell Smith 109 
BRIEFER ARTICLES. 
ITRATES AS A SOURCE OF NITROGEN FOR SAPROPHYTIC FunGcI. Mary H. Smith - 126 
NON-SEXUAL PROPAGATION IN OpuNTIA. II... Carleton E. Preston - - oon ZF 
CURRENT LITERATURE. 
BOOK REVIEWS - . - - - - : i - 129 
A NEw ScuHoo. Botany. THE UMBELLIFERAE. 
MINOR NOTICES - - - - - + - “ ‘ ESR 
NOTES FOR STUDENTS - - 4 . “ “ - - eae, 1) 
NEWS = 5 7 i i Z a : : a = oi sg hQe 


Separates, if d desired, must be pet sper a i advance of publication. Not less than = und meouend vad ge 
ing oe will sed printed, of which 25 thout covers) will be furnished gva/zs, th ual ¢ f the 


1 remainder (and covers, if desired) to be paid for by the author. Separates of “briefer “aeticles sw "(with or 
without covers) will also be supplied at cost he table below shows the approximate cost of sep 
consisting of plain text or text with line engravings. The actual cost may vary from the figures given, 


“ vings. e ua. BIN 
and will depend upon the amount nf a < in re-making the : pages into forms, press work, paper, binding, 
etc. Separates containing half-tones may be expec ero to somewhat more than Aas erates given, the 
increase depending upon the neal of cuts and the a $e work fess upon th 


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Works of General Literature 
OF VALUE TO STUDENTS 


McLOUGHLIN AND OLD OREGON 


By Eva Emery Dye. 12mo, gilt top, with frontispiece, $1. sy 


‘« The vital history ofa State . . . told with art and power.’’— The Argonaut, San Francisco. 
‘ Mrs. Dye’s narrativ: . . reads likea rom . some of the chapters are more interesting 
ae most of the recent powalas’ —New York Times’ Sitarday Review 
‘* From beginning to end the story is lively, brilliant, phsharencaks” he Independent, New Y 


ork. 
‘Mrs s. Dye has the ser iornenrd s _— for bringing out scuieai ey events, the novelist’s gift for vivifying 
isiaaaes.”” — The Buffalo Expre. 


THE LAST YEARS OF THE 190TH HUDSON’S WORK 
CENTURY THE piv E PEDIGREE OF MAN 
OR, THE TESTIMONY OF EVOLUTION AND 
By ELIzABETH WORMELEY LATIMER, author of PSYCHOLOGY TO THE FATHERHOOD OF GOD. 
France in the 19th Century, etc. LUlustrated. By THomSON Jay Jupson, LL.D. Third edi- 
Crown 8vo, $2.50. tion. I2mo, $1. eee 
t t in fav 
rs imer’s esl —— of the 19th Century, Ch Pebiiady ‘Theism, | ful and oe congaed ace of Evolution 
namely. Spain, Ttaly, Eu se cali eH » England, Russia, and Psychology. The work presents an original interpre 
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s0 as to include all ner events oh note up to the very c tation of key phenomena of dev velopment, and advances 
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the series. The is a model of elegant, fa miliar, Thirty-fifth thousand, 0, $1.5 
epistolary a, clear insight, and judicious selection of A Scientific Demonstration, ot “the Future Life 
interesting ma oo 
THE PRIVATE MEMOIRS OF MEMOIRS OF ALEXANDER I. AND 
MADAME ROLAND ge ee sie RUSSIA 
ure ; ‘ By Mm Co DE CH pene -GOUF- 
Edited, with an Introduction, by Epwarp GIL- FIER. ‘Translated pote the French by Mary 
PIN JOHNSON. Illustrated. 12mo, gilt top, Berenice Patterson. With per poet: 
deckle edges, $1.50. eit mi “decks edges, $1.50. 
Only two copies of the original of this work me eee 
e, Roland is botha heroine of romance and an to exist, from one of which the present translatio is made. 
historic ‘sane whom no one could think of overlook- The account of Napoleon is of exceptional int terest, ‘ 
ing.”—The Nation, New York. h h __New York ie. 
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.UNC. AN ONIZED A.comance oF By MARGARET Horton Potter. 
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life story of Anthony Fitz-Hubert, son of the Archbishop of Can icine, who abandoned the brilliant 
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Every pte eat isa ig ade every reer jos court pb pom eots caneholy bi ee os Se utile Comstex Jouraal 


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A GENERAL SURVEY OF AMERI- | OPPORTUNITY, AND OTHER 
CAN LITERATURE ESSAYS AND ADDRESSES ais 
* eorla, 
By MARY FISHER, author of A Group of French pe Pah aah nts arc ag? her Life, 
Pyines te i kee: deckie ot author of Education and t.: z 
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names in our literature. well-balanced, thoughtful, oo es fg in wholesome thought, full of a fine, strong ¥: 
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WALKER 
PRIZES IN NATURAL HISTORY 


B* the provisions of the will of the late Dr. William Johnson Walker two prizes 
are annually offered by the Boston Society oF NATURAL History for the 
best memoirs written in the English language on subjects proposed by a com- 

mittee appointed by the Council. 

For the best memoir presented a prize of sixty dollars may be awarded ; if, how- 
ever, the memoir be one of marked merit, the amount may be increased to one hundred 
dollars, at the discretion of the committee. 

For the next best memoir, a prize not exceeding fifty dollars may be awarded. 

Prizes will not be awarded unless the memoirs presented are of adequate merit. 

The competition for these prizes is not restricted, but is open to all. 

Attention is especially called to the following points : 


In all cases the memoirs are to be based on a considerable body of original and 
unpublished work, accompanied by a general review of the literature of the subject. 


- Anything in the memoir which shall furnish proof of the identity of the author 
shall ot considered as debarring the essay from competition. 


3. Preference will be given to memoirs showing intrinsic evidence of being based 
upon researches made directly in competition for the prize. 


4. Each memoir must be accompanied by a sealed envelope enclosing the author's 
name and superscribed with a motto corresponding to one borne by the manuscript, 
and must be in the hands of the Secretary on or before April 1st of the year for which 
the prize is offered. 


Subject for ro0r: 
Monograph on any problem connected with or any group belonging to the North 
American fauna or flora. 


Subjects for 1902: 
- Nuclear fusions in plants. 
- The fate of specific areas of the germ of chordates, as determined by local 
Mec on. 
- The reactions of organisms to solutions, considered from the standpoint of the 
chemical theory of dissociation. 


C. F,. BaTCHELDER, 
Secretary. 


BOSTON SOCIETY OF sii HISTORY, 
BOSTON, MASS., U.S 


The Forester 


URING the satin year, THE FORESTER, bask eiatiney ascii aso of the 
se rican Forestry ne ation, will be more ing and v fhe 
ev fore. No one who cares for “st 5 OF life i sea woods, or sc 

ested in the a to encourage the pr eservation ae care of the see pat 
be withou 3 rae the contributors gi Nad ag aie HOT, Chief o 
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Forestry; HENRY fetes "Geographer of the i. S. Geo foaieal ee ; Dr. JOHN 
GiIFFORD; Pror. Henry S. GRAVES, of the Yale Forest School; Dr. C. A. SCHENCK, 

i . JAMES acer ager ae ah pane ay Pror. Wo. R. 
DUDLEY, of Sta my oa d pecan: Cal.; FN, SeSH rR, of Harvard University ; 
and many others of note and author nity 0 on S ehete poceiuisias. :: :: Besides a number of 
contributed n tienes pia ye issue of the magazine will contain a record of legislation 
touching the nterests of the etapa S oese (of which there Tae probably be a great 
deal during the coming year), with beitemetes comment, and reviews of recent oe 
tions sae - Lsanot ee 0p etent exper Each number is handsomely illust rated, 

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THE FORESTER is sent to members of the American Forestry Association free of charge. 
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Wm. Wesley & Son’s Botanical Catalogue, 190! 


CONTENTS: 
0. 137 138 of Transactions of Scientific Societies Periodicals 
THE Bibliography History Biographies and Portraits 
Herbals i i Li US 
TURAL Early Botanical Science Linnae 
Handbooks 


Se a Se SS 


Microscopy Morphology and Physiology 
Encyclopaedic works Classification Nomenclature 


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Fossil Plants 
; ptogams Phanerogams 
Natural a of Plants (Floras) 
BOTANY Agricul culture tothe end ofthe S8th centr 


Gesleniey.  Lendease Ganley 

The Flower and Ornamental Garden 

headings Husbandry Tropical Agriculture Commercial Plants 
304 pages Medical Botany Forestry Diseases of Plants 


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A NEW BOOK FOR BIBLE STUDENTS 
CONSTRUCTIVE STUDIES IN THE 
FE OF sitet 


By ERNEST D. BURTON and SHAILER ets 
Professors in the Univ lag of Chic 


THE ORDER OF TREATMEN 
: t for bead the Stevens & Burton * 7 “Harmony of the rag. ge 
a n the 


radi as thus arran 


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THE METHOD OF T 


'estament times are desert 
€ is made to or es events of the Gos; 
‘story in a true, historical persp. 


pn caren in its title; ide th cor 
NStruction ~ himsel a a a Eile. of | Christ, derived ai 

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from the sourc s they exist in the New Tes f 

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contai n: (x). an ypenciat A of the Gos a narra- 
€ notes of information on matters about 
is not easily cheniiee “G) brief 
ifficu cult passages; (4) geogra bhi cal and 
explanations ; (5) ers directions for study, 
student himscit- 6) ort Life of Christ by the 
pig understanding ot ie Gone w algae the student into 
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THE PROSPECTS 
OF THE SMALL 
COLLEGE 


By WILLIAM R. HARPER 


EC AST from an address delivered before 
sso 


The substance o as also given as a 
course of lectures at The University of Chicago, 
during the past summer. 


Now Ready, t2mo, paper, 25¢ ( pos: stpaid), sent 
on veeaise of price by the publishers 


The University of Chicago Press 
CHICAGO, ILLINOIS 


A complete Catalogue of Publications sent on request 


Every Botanist 


Should be familiar with 
the prominent works of 


GEBRUDER BORNTRAEGER 
Publishers. 


Die Glykoside. Chemische Monographie der Pflanzengly- 
koside nebst syst tischer Darstellung der kiinstlichen 
Glykoside von Dr. I. I. L. von Run, Director der Reichsver- 
suchsstation in-Mastricht. 8vo. Cloth, $2.50. 


Das Werk giebt—wie es bisher noch nirgends geschehen — eine eingehende 
chemische Behandlung der Glykoside—nicht nur eine kurzgefasste Zusammen- 
stellung der chemischen Eigenschaften dieser Korperkilasse, sondern die Darstel- 
lungsmethode, die Griinde, welche zur Aufstellung der Constitutionsformeln 
gefithrt haben etc., so dass das Buch in chemisch-pharmaceutischen wie pharma- 
hologischen Kreisen sowie unter den Studirenden und sonstigen Freunden der 
phytochemischen Forschung sicher mit grosser Freude begriisst werden wird. 


Die Harze und die Harzbehalter. Historisch 
kritische und experimentelle, in Gemeinschaft mit zahlreichen 
Mitarbeitern ausgefiihrte Untersuchungen von Proressor DR. 
A. Tscuircn, Director des pharmaceutischen Institutes der Uni 
versitat Bern. Mit 6 Tafeln. 8vo. Half calf, $5.00. 

Das Werk stellt zum ersten Mal das gesammte Material dieser wichtigen 
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Write for free Catalogue; postpaid. Address: 
Gebriider Borntraeger, Publishers, 
BERLIN, S. W. 46: 


SCHONEBERGERSTRASSE 17a 


2s SS ui ee 


F | 
*rench, and Latin, as well as in English. 


The Journals of the University of Chicago 
BEING THE DESCRIPTIVE LIST OF ONE WEEKLY, FOUR 
MONTHLY, ONE BI-MONTHLY, THREE QUARTERLY, 
AND ONE SEMI-QUARTERLY PUBLICATIONS & #& 


THE BIBLICAL WORLD 


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onthl = Siemens oh Subicrption price in the bijou States, 

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8 


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VOLUME XXxXI1 NUMBER 2 


DOTANICAL (Gee 1 e 
FEBRUARY, 1901 


THE PHYSIOGRAPHIC ECOLOGY OF CHICAGO AND 
VICINITY; A STUDY OF THE ORIGIN, DEVELOP- 
MENT, AND CLASSIFICATION OF PLANT SOCIE- 
gy 


CONTRIBUTIONS FROM THE HULL BOTANICAL LABORATORY. 


HENRY CHANDLER COWLES. 
(WITH THIRTY-FIVE FIGURES.) 
I. The content and scope of physiographic ecology. 


Wirun the last few years the subject of ecology has come to 
find a place of more or less importance wherever botany is 
Studied in its general aspects. The limits of the subject, how- 
ever, have not yet been defined, nor have many attempts been 
made to bring order out of the chaos which exists with regard 
to the arrangement of the subject-matter. Thé main purpose 
of the present paper is to suggest a classification of a portion of 
the ecological field. 

Whatever its limits may be, ecology i is essentially a study of 
origins and life histories, having two well-marked phases; one 
phase is concerned with the. origin and development of plant 
structures, the other with the origin and development of plant 
Societies or formations. The plant structure side again has two 
aspects, one viewing organs or plant forms as a whole, the other 
viewing the tissues which make up the organs; the former might 

73 


74 BOTANICAL GAZETTE [FEBRUARY 


be called organographic ecology or even organography, while 
the latter may be called ecological anatomy. 

It may be well to speak more in detail concerning the dis- 
tributional phase of ecology. There are two distinct aspects 
here also, the one local, the other regional. Climatic factors, 
particularly temperature and atmospheric moisture, permit the 
subdivision of the earth into great zones or regions with charac- 
teristic plant formations which extend over wide areas. Exam- 
ples of this type are tropical evergreen forests, deserts in 
continental interiors, prairies, deciduous forests, arctic tundras. 
These formations are widespread because the factors that pro- 
duce them are widespread. We might call these formations cli- 
matic formations (following Schimper’) and the subject that 
deals with them geographic ecology or ecological plant geogra- 
phy. In contrast with the above there are the local or edaphic 
factors, such as soil (including its moisture, air, and temperature 
relations), slope, light; in other words, factors that are largely 
due to the physiographic nature of the district. Where the cli- 
mate is the same these factors produce marked changes locally, 
and there results a variety of plant societies, such as swamp, 
dune, bottom forest, river bluff, etc. These correspond to 
Schimper’s' edaphic formations or Warming’s plant societies, and 
the subject that deals with them may be called physiographic 
ecology. 

In order to justify the terminology here given it will be 
desirable to trace briefly the history of the study of plant socie- 
ties and then to depict the intimate relations which exist between 
the physiography of a region and its flora.’ Before the appear- 
ance of Warming’s ecological plant geography” there had been 
no attempt to classify the plant formations of the globe in a sys- 
tematic manner. Warming introduced the term plant society in 
place of plant formation, because of the varied use of the latter, 

*SCHIMPER, A. F. W.: Pflanzengeographie auf physiologischer Grundlage 
173-176. Jena, 1898. 


* WARMING, E.: Plantesamfund. Copenhagen, 1895. German edition, trans- 
lated by Knoblauch. Berlin, 1896. 


| 
f 


1gor | PHYSIOGRAPHIC ECOLOGY OF CHICAGO 75 


and made variations in the water content of the soil a basis of 
classification. Plant societies were divided into hydrophytes, 
mesophytes, and xerophytes; further than this, however, little 
attempt was made by Warming to subdivide the plant socie- 
ties, except in the case of swamp and dune plants. In these two 
instances, as will appear later, Warming adopted the order of 
succession in his method of treatment. 

Since Warming’s great work appeared, ecologists have in gen- 
eral followed his ideas and have attempted to work them out. 
Noteworthy contributions have also been made which make a 
comprehensive view of the subject more possible. Schimper? 
has analyzed in a most thorough manner the conditions which 
determine the distribution of plants in the large, though he has 
discussed but briefly the purely local or habitat factors. We owe 
to him, however, the first clear statement of the distinction 
between edaphic and climatic factors and formations. The 
minuter treatment of the edaphic formations did not lie within 
his field, and he has attempted in no sense to give a classifica- 
tion, except in the case of climatic formations. 

_ Graebner*5 has published a classification of some of the Ger- 
man vegetation formations, which has not received the attention 
it deserves. This classification is based in the main on the 
chemical and physical characteristics of the soil. The primary 
divisions are chemical, depending on the richness or poverty of 
the soil in plant foods. The secondary divisions are based 
chiefly: on soil moisture.. 

N. H. Nilsson® in some preliminary notes on Swedish 
Swamps and their vegetation called attention to the striking 
difference between hydrophytic and xerophytic same ae and 

3 Op. cit. 

*GRAEBNER, P.: Gliederung der westpreussischen Vegetationsformationen. 
Schrift. Natale Ges. Danzig 9 : 43-74. 1898. See Bot. Centralb. 75: 277-279. 1898. 

EBNER, P.: Ueber die Bildung natiirlichen Vegetationsformationen im 
norddeutschen Flachlande. Archiv der Brandenburgia 4: 137-161. 1898. See Bot 
Centralb, 77 :212-214. 1899. 


° NItsson N. H.: Einiges tiber die Biologie der scicecotirer Sumpfpflanzen. 
Bot. Centralb. 76:9-14. 1898. 


76 BOTANICAL GAZETTE © [FEBRUARY 


gave expression to the view that differences in food supply may 
account for the facts observed. Schimper’? has a somewhat 
similar view, but explains differences in the vegetation of various 
swamps more along the line of relative ease or difficulty of 
absorption in the swampy soils. Both Nilsson and Schimper 
regard peat bogs as essentially xerophytic. 

In this historical sketch mention should be made of the 
work of Flahault*9, who has projected and begun to execute a 
detailed map of the plant societies of France. Conventional 
color tones are chosen for the various plant societies, and they 
are plotted on topographic contour maps. Robert. Smith*”* 
had just entered upon a similar work in Scotland when death put 
a stop to his labors. So far as the author knows, Flahault has 
not concerned himself particularly as yet with the matter of 
classification. Alb. Nilsson* has recently published some inter- 
esting studies of Swedish plant societies, tracing the order of 
succession of vegetation on cliffs and moors. Still more recently 
Meigen*3 has published a series of short articles, tracing the 
order of succession in a number of plant societies. Besides the 
authors named thus far, Drude™*, MacMillan*s, and Pound and 
Clements have given excellent treatments of the plant forma- 
_ tions in their respective regions. 

7 Op. cit. 18. 

SFLAHAULT, Ch.: Projet de carte botanique forestitre et agricole de la France. 
Bull, Soc. Bot. France 41: 56-94. 1894. Ann. de Géographie 5 : 449-457. 1896. 

9FLAHAULT, Ch.: Essai d’une pes botanique et forestiére de col France. 
Ann. de Gésammpble 6: 289-312. 1897, etc. 

7°SMITH, ROBERT: On the study a plant associations. Nat. Sci. 14: 109-120. 

9. 
H, ROBERT: Botanical survey Spa ae I. Edi ee district. HU. 
North Perthshire district. Scot. Geog. Mag. 16: 385~416; 441-467. 

12 NILSSON, ALB.: Nagra drag ur de Pens vaxtsamhallenas ia ada ea 
Bot. Not. ee enc 123-135. 

MEI PP aniprena tea ek Formationsfolge im Kaiserstuhl. _ Deutsch. 
Bot. hone ia 145~147, e 
™ DRuDE, O.: Deutschlands ees I. Teil. Stuttgart. 1896. 

5 MACMILLAN, C.: Observations on the couligeian of oe along shore at 
Lake of the Woods. Minn. Bot. Stud. 1: 949-10 

76PouUND and CLEMENTS: The shee cae at seit I. General Survey- 
Lincoln. 1898. 


Igor] PHYSIOGRAPHIC ECOLOGY OF CHICAGO 77 


During 1896 and 1897 the author of this paper, in company 
with his students, endeavored to classify the vegetation about 
Chicago in accord with Warming’s principles. In 1898 a similar 
and more careful study of this kind was made in northern 
Michigan. It was of course found to be possible to classify the 
plant societies by the amount of water in the soil, but it was 
found that such a classification put together plant societies 
radically different in their character, and separated plant societies 
that were obviously closely related. The best instances of these 
difficulties were seen in the case of heaths and moors. Not 
only were heaths and moors found to have closely similar species 
and vegetative adaptations, but these plant societies were often 
found grading into each other. In water content these societies 
were very different, the peat moor or bog being hydrophytic and 
the heath xerophytic. Thus some factor other than water con- 
tent is responsible for both. In that same year (1898) Nilsson 
and Schimper published their views on the causes of the xero- 
phytic character of moor vegetation, as outlined above. Further- 
more the vegetation of peat bogs is radically different from that 
of river swamps which have the same water content. 

Further field study but added to the difficulties of the situa- 
tion, and the need of another classification was keenly felt. It 
Was seen at once that no one factor could take the place of the 
water content of the soil, since that is obviously the most impor- 
tant of all direct factors in distribution, as Warming so ably 
shows. An attempt was therefore made to relate the facts of 
distribution to combinations of factors, with the following 
results. The classification which is about to follow is based in 
the main on two ideas, viz., that a classification to be true must 
be genetic and dynamic. In other words, an attempt is made to 
Sroup plant societies according to their relationship and their 
evolution. 

The influences which govern the distribution of plants reside 
in the air or soil (regarding water as soil, for the sake of con- 
venience). The atmospheric influences (light, heat, air) operate 
Over wide areas and have subordinate edaphic importance, 


78 BOTANICAL GAZETTE [FEBRUARY 


whereas the soil influences (soil heat, soil air, soil water, soil 
chemistry and physics) are of predominant edaphic importance, 
though of little account when distribution over wide areas is 
considered. We may say then that atmospheric or climatic 
factors determine distribution in the large, while local differences 
are produced by changes in the edaphic or soil factors. 

The soil conditions are chiefly determined by the surface 
geology and the topography. The original character of the soil, 
whether rock, sand, clay, or marl, depends upon the geological 
relations. From the vegetation standpoint the topographic rela- 
tions are commonly much more important, since they condition 
the presence or absence of drainage, and hence cause striking 
variations in air content and humus. Doubtless the charac- 
teristic features of peat bog vegetation are due to the absence 
of drainage and consequent poor aeration and accumulation 
of organic products. Moreover, in so far as the atmospheric 
factors have an influence on distribution locally, it is largely 
due to topographic diversities, such as angle and direction of 
slope. 

Having related the vegetation largely to topography, we must 
recognize that topography changes, not in a haphazard manner, 
but according to well-defined laws. The processes of erosion 
ultimately cause the wearing down of the hills and the filling up 
of the hollows. These two processes, denudation and deposi- 
tion, working in harmony produce planation; the inequalities are 
brought down to a base level. The chief agent in all these 
activities is water, and no fact is better established than the 
gradual eating back of the rivers into the land and the wearing 
away of coast lines; the material thus gathered fills up lakes, 
forms the alluvium of flood plains, or is taken to the sea. Veg- 
etation plays a part in all these processes, the peat deposits add- 
ing greatly to the rapidity with which lakes and swamps are 
filled, while the plant covering of the hills, on the contrary, 
greatly retards the erosive processes. Thus the hollows are 
filled more rapidly than the hills are worn away. As a conse- 
quence of all these changes, the slopes and soils must change ; 


Igor | PHYSIOGRAPHIC ECOLOGY OF CHICAGU 79 


So, too, the plant societies, which are replaced in turn by others 
that are adapted to the new conditions. 

There must be, then, an order of succession of plant societies, 
just as there is an order of succession of topographic forms in 
the changing landscape. As the years pass by, one plant society 
must necessarily be supplanted by another, though the one 
passes into the other by imperceptible gradations. Here then 
is a classification both genetic and dynamic, a classification 
which has a place for all possible ecological factors. It is based 
on the normal physiographic changes of a region and hence 
should be called a physiographic classification. One thing 
more must be recognized, and that is that environmental influ- 
ences are normally cumulative. A plant society is not a prod- 
uct of present conditions alone, but the past is involved as 
well. For example, a hydrophytic plant society may be seen 
growing in a mesophytic soil ; the author has seen a mesophytic 
tamarack swamp which can be explained only in this way. We 
have in this phenomenon a lagging of effects behind their 
cumulative causes, just as the climax of the heat in summer 
comes long after the solstice. 

In a classification like this great emphasis is placed on bor- 
der lines or zones of tension, for here, rather than at the center 
of the society, one can best interpret the changes that are tak- 
ing place. Of course the order of succession referred to above 
is a vertical or historical one. One plant society is said to fol- 
low another if it is actually superimposed upon the one preced- 
ing. In many cases, if not in most, there is a horizontal order 
of succession at the present time that resembles the vertical 
Succession of which we now have only the topmost member. 
Instances of similarity between vertical and horizontal orders of 
Succession are well shown in peat swamps and along shores and 
flood plains. Along a sandy shore it is only by studying the 
horizontal succession that one can get any idea of the vertical, 
Since all fossil traces of preceding plant societies have passed 
away. In peat swamps one can sometimes verify the results of a 
horizontal zonal study by investigating the fossil remains beneath 


80 BOTANICAL GAZETTE [FEBRUARY 


We may now outline the main features of a physiographic 
classification of plant societies. Speaking in the large, the tend- 
ency of the erosive processes is to reduce the inequalities of the 
topography and produce a base level. This base level may not 
soon be reached, though geological history furnishes instances 
of extensive base leveling. Crustal movements interfere with 
the erosive agencies and a mature base level topography may 
become rejuvenated by a great uplift of the land, or sinking on 
the other hand may check the rapid action of erosion. Yet 
even with the crustal movements there go these topographic 
changes and with them the plant societies must change. Put- 
ting the facts of physiography in the terms of ecology, the con- 
ditions become more and more mesophytic as the centuries pass. 
In a young topography, such as the recently glaciated areas of 
Michigan, Wisconsin, and Minnesota, there is a great variety of 
topographic conditions and of plant societies. Among these are 
many hydrophytic lakes and swamps and many xerophytic hills. 
The hills are being denuded and the swamps and lakes are being 
filled, so that the hydrophytic and xerophytic areas are becoming 
more and more restricted, while the mesophytic areas are becom- 
ing more and more enlarged. In passing from youth to old age 
then, a region gradually loses its hydrophytic areas and also its 
xerophytic areas, though in the latter case there is usually at first 
an increase in the xerophytic areas which is due to the working 
back of the young streams into the hills. The latter conditions 
are well shown in Iowa; in the comparatively recent Wisconsin 
drift of north-central Iowa the topography is much less diversi- 
fied and there are fewer xerophytic areas than in the older Iowan 
drift farther south, which has been greatly dissected by stream 
erosion. Later, however, the inequalities are removed, and we 
find great mesophytic flood plain areas, such as are seen along 
the lower Mississippi. 

_ From what has been stated it will be seen that the ultimate 
stage of a region is mesophytic. The various plant societies 
pass in a series of successive types from their original condition 
to the mesophytic forest, which may be regarded as the climax 


Igor | PHYVSIOGRAPHIC ECOLOGY OF CHICAGO 8 


or culminating type. These stages may be slow or rapid; some 
habitats may be mesophytic from the start; undrained lakes and 
swamps fill up and become mesophytic with great rapidity, 
whereas granite hills might take many centuries or even geological 
epochs in being reduced to the mesophytic level. Again the 
stages may be direct or tortuous; we have already seen how the 
first consequences of stream erosion may be to make mesophytic 
areas xerophytic. So, too, in flood plains, the meanderings of 
the river may cause retrogressions to the hydrophytic condition 
such as are seen in oxbow lakes, or the river may lower its bed 
and the mesophytic flood plain become a xerophytic terrace. 
But through all these changes and counterchanges the great 
mesophytic tendency is clearly seen; mesophytic areas may be 
lost here and there but many more are gained, so that the 
approach to the mesophytic base level is unmistakable. More- 
over, the retrogressive phases are relatively ephemeral, while the 
progressive phases often take long periods of time for their full 
development, especially in their later stages. 

The above phenomena postulate congenial climates and more 
or less static crustal conditions. It is obvious, however, that 
€roSive processes in a desert region do not result in a mesophytic 
flora; the same is true of alpine and arctic climates. Again, the 
climate of all regions is doubtless changing, as it has changed 
in past ages. So, too, there are crustal movements up and down. 
In other words the condition of equilibrium is never reached, 
and when we say that there is an approach to the mesophytic 
forest, we speak only roughly and approximately. As a matter 
of fact we have a variable approaching a variable rather than a 
constant. These conditions do not destroy the validity ofa 
physiographic classification, but rather they require an enlarge- 
ment of conception. Retrogressive phases, 2. ¢., away from the 
mesophytic and toward the hydrophytic or xerophytic, must be 
_ included, as well as progressive phases away from the hydrophytic 
oF xerophytic and toward the mesophytic. In this way all pos- 
 Sible conditions are accounted for. For example, upward crustal 
movements make hills more xerophytic and swamps more 


o 


82 BOTANICAL GAZETTE [ FEBRUARY 


mesophytic, whereas downward movements make hills more 
mesophytic and swamps more hydrophytic. Thus in the upward 
movement of hills and the downward movement of swamps, 
physiographic processes are more or less neutralized and we may 
speak of retrogressive tendencies ; in the other two cases physio- 
graphic processes are accelerated and we have more rapid prog- 
ress toward the mesophytic climax. !faclimate grows colder or 
more arid, we find retrogressive tendencies toward the xerophytic 
condition, while in a climate that is getting more moist or more 
genial the mesophytic tendencies of the erosive processes are 
accelerated. Furthermore, climatic and crustal changes are 
commonly so slow in comparison with physiographic changes, 
that it is usually difficult to decipher their tendencies. We can be 
far more sure, in other words, with relation to the past and future 
of a topographic form and its plant societies, so far as erosion is 
concerned, than we can as to the actual effect that changing 
climatic and crustal conditions are making. 

One other modification of the physiographic theory is 
necessary, as has been clearly shown by recent field studies. 
While changes in plant societies are certain to follow changes 
in topography, it does not necessarily follow that plant societies 
remain the same if topographic conditions remain unchanged. 
In other words, changes may take place in plant societies more 
rapidly than in the topography. A cycle of vegetation may be 
much shorter than a cycle of erosion. One of the most interest- 
ing cases of this is seen in a growing river system. In the 
ravine stage there may be a rapid change from the xerophytic 
to the mesophytic plant societies on the slopes. As the valley 
widens xerophytic conditions appear on the slopes once more. 
This first and relatively short-lived mesophytic condition may 
be called a temporary climax, in distinction to the more perma- 
nent climax of the base level. 

In a study of plant societies such as this, it must be recog- 
nized that orders of succession are not the same in various 
regions. There is probably a close analogy between the various 
society life histories where climatic conditions are the same, but 


ee 


1901} PHYSIOGRAPHIC ECOLOGY OF CHICAGO 83 


it remains true, nevertheless, that each region must be worked 
out by itself. The general principles that are involved in the 
dynamics of plant societies, however, ought to be essentially the 
same everywhere. Some instances will be given which will 
show the necessity for working out the life history of the plant 
societies in all regions. While the culminating type throughout 
the northern states east of the Mississippi river is probably a 
deciduous mesophytic forest, yet the elements of this forest dif- 
fer greatly in different localities. In central Michigan the 
maple, beech, and the evergreen hemlock appear to be the lead- 
ing character trees of the mesophytic forest. In Indiana and 
Illinois the hemlock is not one of the dominant trees of this 
forest. In the Alleghanies of Tennessee a large number of tree 
Species assume a place of almost equal importance in the meso- 
phytic forest. Again, in the Chicago region the tulip tree and 
buckeye are rare and confined to the flood plain forests, while 
in Tennessee these trees are found in many other plant societies. 
In the Chicago region the arbor vitae is confined to undrained 
swamps and xerophytic cliffs, while in northern Michigan it is 
found in many other habitats. We may perhaps summarize 
these data by saying that each species varies in habitat in dif- 
ferent regions, and that in general a species can grow in the 
largest number of plant societies at its center of distribution, since 
there the climatic conditions favor it most highly. In other 
regions, especially near its areal limits, it can grow only in those 
plant societies which resemble most closely in an edaphic way 
the climatic features at the distribution center. Thus the tulip 
and buckeye, which flourish best in the mesophytic forest climate 
of the Alleghanies, are found near Chicago only in the most pro-. 
nounced of our mesophytic societies, those of the flood plain. 
Again, the arbor vitae, and with it many conifers and heaths, 
Stow near Chicago only on the cliffs and dunes or in the 
undrained swamps, since these are the most pronounced of our 
xerophytic habitats and most closely resemble the xerophytic 
northern climates. 

A few words should be said’in the way of indicating the 


\ 


84 BOTANICAL GAZETTE | FEBRUARY 


relationship between this and other classifications. Warming’s 
classification, based on the water relations, at two points agrees 
with the physiographic theory, viz., in the treatment of swamps 
and dunes. Each of these is treated from the standpoint of the 
order of succession as revealed by zonal distribution, though in the 
case of the dunes this order is not one of decreasing or increas- 
ing water content. Alb. Nilsson and Meigen (see above), and 
for that matter many other authors, have studied various plant 
societies from the standpoint of their order of succession, but so 
far as the author is aware no previous attempt has been made to 
establish a comprehensive theory on this basis. Graebner’s 
classification (see above) has several points in common with 
the physiographic theory, especially as it relates heaths with 
moors. In this connection it will be of interest to refer toa 
paper by J. B. Woodworth"? which indicates a fertile line of 
research that is but now being taken up by biologists. He 
shows how the base-leveling processes must influence the evolu- 
tion of species, since these processes constantly erect new and 
destroy old barriers, and hence cause isolation in the one case 
and intermingling of species in the other. Woodworth gives a 
number of instances of the influence of base leveling upon ani- 
mal life, and he refers, although but slightly, to the changes 
which must take place in the plant life as regions are uplifted or 
approach base level. It seems surprising that such a great field 
of study has been neglected until now. C.C. Adams, in a paper 
as yet unpublished, and C. T. Simpson * have recently given spe- 
cial cases of the interrelations between physiographic changes 
and animal distribution. 

The general principles of the physiographic theory have 
been developed as a result of studies in various sections of the 
country. Since 1898, when the author first began to work along 
these physiographic lines, the main thought has been to subject 

*7 WoopworTH, J. B.: ss Song between base-leveling and organic evolu- 
tion. Am. Geol. 14 : 209-235. 

%8StmpPson, C. T.: On the cee of the Unionidae regarding the former 
course of the Tennessee and other Southern rivers. Science N. S. 12:133-136- 
1900. 


rw” A 


Igor] PHYSIOGRAPHIC ECOLOGY OF CHICAGO 85 


the theory to the most rigid test possible. In connection with 
a number of students, investigations have been carried on about 
Chicago, in northern Michigan, in Tennessee, and along the 
Atlantic coast. In all cases it’ has been possible to find a 
general consonance between the facts of distribution. and the 
principles as stated above. The theory has suffered many modi- 
fications since its first conception, and doubtless it will suffer 
more. Indeed, it may be discarded altogether for.some other 
better theory. Nevertheless, publication at this time seems to 
be justified, and it is hoped that this paper may aid in solving 
some of the riddles of ecology. 

The author especially wishes to mention in this place the 
work of his student and associate, Mr. H. N. Whitford, who has 
in preparation a physiographic study of the forests of north- 
ern Michigan. The author is likewise especially indebted to 
another of his students, Mr. W. B. McCallum, who has taken all 
of the photographs with which this paper is illustrated, with the 
exception of figs. 7 and 73, which were contributed by Professor 
J. J. Allison, of Joliet. Acknowledgment should also be made 
of the help given by three excellent papers which deal with 
the general physiographic and geographic features of the Chi- 
cCagoarea. To these works by Leverett, Blatchley,” and Salis- 
bury and Alden* the author has made constant reference. The 
author has likewise freely used the work of Higley and Raddin.” 

In the following pages the various series of the Chicago area 
are discussed in some detail. Two general groups are made; 
the inland and the coastal. The inland group is subdivided into 
three series, river, swamp, and upland. The coastal group is 
eccanione into two series, lake bluff and dune. The river series 


ERETT, F.: The Pleistocene features and eo of the Chicago area, 
Chica or 
7 BLAT LEY, W. S.: The geology of Lake and Porter counties, Indiana. 
tae fron ie Twenty-second Annual Report of the Department of Geology and 
Natural Resources of Indiana. Indianapolis, 1 1897. 
* SALIsBuRY, R. D., and ALDEN, W. C.: The geography of Chicago and its 
environs. Chicago, 1899. 
*“ HIGLEY, W. K., and RAppIN, C. S.: The flora of Cook county, Illinois, and a 
part of Lake county, judas Chicago; 1891. 


86 BOTANICAL GAZETTE [FEBRUARY 


is remarkably tortuous, involving constructive and destructive, 
progressive and retrogressive phases. The treatment begins 
with an erosion gully; then there follow in order the ravine, 
both in clay and in rock, the xerophytic bluff, and the mesophytic 
forest. The depositional phases of the river begin with the 
appearance of a permanent stream; then follow the various 
stages of the flood plain culminating in the mesophytic forest. 
The swamp series begins with the pond, treats next the various 
types of swamps and ends with a brief discussion of the prairie. 
In the upland series the various stages of the rock hills and 
then of the clay hills are taken up in turn, culminating in the 
mesophytic forest. The coastal group is next discussed, begin- 
ning with the lake bluff. Finally, there is a brief treatment of 
the dune series from the beach on through the embryonic and 
active dune to the established dune on which there finally appears 
the mesophytic forest. 


II. The plant societies. 
A. The inland group. 
I, THE RIVER SERIES. 

A. The ravine-—No topographic forms lend themselves so 
well toa physiographic sketch of the vegetation as do those 
that are connected with the life history of a river. Beginning 
with the ravines, which are deep and narrow, because of the 
dominance of vertical cutting, we pass to the broader valleys, 
where lateral cutting becomes more pronounced. From this 
stage on we have to deal with two phases of river action, the 
destructive, which is concerned with the life history of the bluff, 
and the constructive, which has to do with the development of 
the flood plain. 

Wherever there is an elevated stretch of land adjoining a 
body of water, such as a lake bluff, one is apt to.find excellent 
illustrations of the beginning of a ravine. Fig. z shows an 
embryonic ravine of a type that may be seen frequently along 
the clay bluffs between Evanston and Waukegan. A ravine of 
this type is essentially a desert, so far as plant life is concerned. 


1901 | PHYSIOGRAPHIC ECOLOGY OF CHICAGO 87 


The exposure to wind and to alternations of temperature and 
moisture is excessive. The lack of vegetation, however, is due 
chiefly to the instability of the soil; this instability is particularly 
great in the case of clay bluffs such as these, where the seepage 
of water causes extensive landslide action. No plants can get 
a foothold in such a place, unless it be a few species that may be 
able to make their appearance between periods of landslide 
action; among these plants annuals particularly predominate. 
The perennials that may be found in such places are almost 
entirely plants which have slid down the bank. Near the center 
of fig. zis a clump of shrubs that has slid down in this way. 
Ravines of a similar type may also be seen at many places 
inland, and wherever found the poverty of vegetation on the 
slopes is the most striking character. 

As a ravine extends itself inland the conditions outlined 
above may be always seen about its head, but toward the mouth 
of the ravine the slopes are less precipitous. Torrents cut down 
the bed of the ravine until a depth is reached approaching the 
water level at its mouth. From this time on the slopes become 
reduced and the ravine widens more than it deepens, by reason 
of lateral cutting, landslide action, and side gullies. After a 
time a sufficient stability is reached to permit a considerable 
growth of vegetation. If the erosion is slight enough to allowa 
vegetation carpet to develop, a high degree of luxuriance may 
be attained. In fact ravine conditions are usually extremely favor- 
able for plants, after the initial stages have passed. In a com- 
paratively few years the vegetation leaps as it were by bounds 
through the herbaceous and shrubby stages into a mesophytic 
forest, and that, too, a maple forest, the highest type found in 
our region. Nothing shows so well as this the brief period 
necessary for a vegetation cycle in a favored situation when 
compared with an erosion cycle. 

Of such interest are the facts just noted that it is worth while 
to mention some of the characteristic ravine plants. Perhaps the 
most characteristic trees of the Glencoe ravines are the bass- 
wood (Tilia Americana) and the sugar maple (Acer saccharinum), 


88 BOTANICAL GAZETTE [FEBRUARY 


though the ash, elm, and other trees are frequent. The most 
characteristic undershrub is the witch hazel (Hamamels Vzr- 
giniana). The herbaceous plants are notoriously vernal forms, 
such as Hepatica, Thalictrum, Trillium, Mitella, Dicentra, 
Sanguinaria; mosses abound and liverworts are frequent. A 
ravine with the above vegetation is shown in jig. 2. We can 
explain this flora only by regarding it as having reached a tem- 
porary climax. Ravine conditions are more favorable for plants 
than those that precede or follow. The instability and exposure 
of the gully have gone; in their place there is protection from 
wind and exposure. The shade and topography favor the collec- 
tion and conservation of moisture, and as a result there is.a rapid 
development intoa high-grade forest, as outlined above. 
Rock ravines are much less common in the Chicago area than 
are those of clay, since the underlying limestone rarely comes 
near the surface. Excellent illustrations of stream gorges are 
to be seen at Lockport, and also inivarious tributaries of the IIli- 
nois river near: Starved rock, A striking difference between 
these rock’ gorges or cafions and the clay ravines is in the slope 
of the sides.; The physical nature of the rock excludes landslide 
action, hence the sides are often nearly vertical for a long time. 
Lateral cutting is also relatively slow as compared with clay. 
Thus the conditions for vegetation at the outset are much more 
favorable than in a clay ravine. Rock-bound gorges are very 
shady and often dripping with moisture, hence liverworts and 
many mosses find here a habitat even more congenial than in 
the clay. Among the higher forms are found the most extreme 
shade plants that we have, such as Impatiens, Pilea, and shade- 
loving ferns, plants whose leaves are broad and remarkably thin. 
figs. 3 and 4 represent cafions of the above description, whose 
rocks drip with moisture. 
_ The stages of development pass much more slowly in caiions 
than in clay ravines, largely because the primitive conditions of 
shade and moisture remain for a long period of time. Nor do 
the steep slopes permit the development of a wealth of trees and 
shrubs, since a secure foothold is not easily found. However, 


1901 | PHYSIOGRAPHIC ECOLOGY OF CHICAGO 89 


as the cafion broadens out and the slopes become less steep, 
shrubs and trees come in, though a typical mesophytic forest is 
rarely seen.. The Starved rock ravines are cut in St. Peters 
sandstone, those at Lockport in the Niagara limestone, yet the 
vegetation in the two places is essentially alike; at any rate the 
resemblances are greater than the differences. Much has been 


IG. I.—Embryonic ravine in the lake bluff at Glencoe. Entire absence of 
eee on the unstable ae slopes with the exception of shrubs and grasses that 
have slid down from the t 


written on the physical and chemical influences of rocks upon 
the vegetation. The facts seen here seem to show that the 
phy siographic stage of a region is more important than either. 
The flora of a youthful topography in limestone, so far as the 
author has observed, more closely resembles the flora of a similar 
Stage in sandstone than a young limestone topography resembles 
an old limestone topography. A limestone ravine resembles a 
Sandstone ravine far more than a limestone ravine resembles an 
€xposed limestone bluff or a sandstone ravine resembles an 


go BOTANICAL GAZETTE [FEBRUARY 


exposed sandstone bluff. We may make the above statements 

in another form. Rock as such, or even the soil which comes 

from it, is of less importance in determining vegetation than are 

the aerial conditions, especially exposure. And it is the stage 

in the topography which determines the exposure. | 
All of the preceding statements as to topographic stages, | 


TT 


ee es 


FIG. 2.—Ravine at Glencoe with a mesophytic forest vegetation on the slopes 
(temporary climax). Presence of erosive forces indicated by leaning trees. Water 


in the stream bed only after rains. ( 


whether young or old, refer not to times but to constructional 
forms. Two ravines, equally youthful from the topographic 
standpoint, may differ widely as to actual age in years or centu- 
ries, Since erosion is more rapid in one rock than in another. 1 
In our region, however, elements of actual time are not very 
important, except as between rock and clay, since the limestone 
is less soluble and the sandstone is more easily eroded than is 
often the case. 

| B. The river bluff—As a valley deepens and widens, the 


Igor] PHYSIOGRAPHIC ECOLOGY OF CHICAGO gi 


conditions outlined above undergo radical changes. From this 
point it will be necessary to discuss two phases in the growing 
river, the bluff phase and the bottom phase. We have left the 
clay ravine bluffs in a state of temporary climax, clothed with 
luxuriant mesophytic forest trees and with a rich undergrowth 
of vernal herbs. More and more the erosive processes are 
conspicuous laterally, and 
widening processes prevail 
over the more primitive 
deepening. As a result, 
the exposure to wind, sun- 
light, and changes of tem- 
perature increases; thé 
moisture content of the 
slopes becomes less and 
less. The rich mesophytic 
herbs, including the liver- 
worts and mosses, dry up 
and die. The humus oxi- 
dizes_ more rapidly, and 
a xerophytic undergrowth 
comes in. In place of 
Hepatica and its associ- 
ates, we find Antennaria, 
Poa compressa, Equisetum 
hyemale, and other xero- eo ce eae ae 
phytic herbs ; Polytrichum i ce bagi soc opiate 
also replaces the mesophy- forces prominent, and vegetation slight on 
tic mosses. The first signs the dripping slopes. 

of the new xerophytic flora ae 
are seen at the top of the ravine slope; indeed the original 
x€rophytic plants may never have been displaced here by the 
ravine mesophytes. As the ravine widens, the xerophytic plants 
creep down the slope, often almost to the water’s edge. Some 
of the young ravines between Evanston and Waukegan show 
xerophytes at the summits of the slopes. Fig. 5 shows a 


g2 BOTANICAL GAZETTE [ FEBRUARY 


widening ravine at Beverly hills; the vegetation is much less 
luxuriant than that shown in the young ravine of fig. 2. 

After a few years have passed, xerophytic shrubs appear on 
the bluff in place of the witch hazel and its associates. And it 
is not long until xerophytic or semi-xerophytic thickets prevail, 


Fic. 4.—Side of a cafion in the St. Peters sandstone at Starved rock. Herbaceous 
shade vegetation on the precipitous slopes. 


in place of the former mesophytic undershrubs. Among the 
more characteristic of these shrubs are the hop tree (Prelea trifo- 
liata), bittersweet ( Celastrus scandens), sumachs (Rhus typhina 
and R. glabra), choke cherry (Prunus Virginiana), nine-bark 
(Physocarpus opulifolius), wild crab (Pyrus coronaria). Two 
small trees are common on stream bluffs, the service berry 
(Amelanchier Canadensis) and the hop hornbeam ( Osétrya Virgin- 
ica); this last species is perhaps the chief character tree of river 


inten ee: 


a 


— £r 


Igor] PHYSIOGRAPHIC ECOLOGY OF CHICAGO 93 


bluffs and is rarely absent. Perhaps the best examples of xero- 
phytic stream bluffs near Chicago are along Thorn creek. One 
of the most interesting things about these bluff societies is the 
frequent presence of basswoods and sugar maples. Doubtless 
these trees look back to the mesophytic associations that have 
otherwise disappeared. As would be expected, the last of the 


Fic. 5.—Open ravine at Beverly ~~ gsi gentle slopes covered with a less 
mesophytic vegetation than is shown n fig. 2. Dominance of oaks in place of 
maples and basswoods. 


mesophytes to die are trees, because they are longer lived than 
herbs and shrubs, and also because their roots reach down to the 
moisture. But they cannot be succeeded by their own kind, 
inasmuch as the critical seedling stages cannot be passed success- 
full lly. 

The life history of the rock ravines or cafions is somewhat 
different. When the ravine vegetation is at its height, the mois- 
ture and shade are greater here than in the clay, hence the high 
development of liverworts and their associates. As the ravine 


94 BOTANICAL GAZETTE [ FEBRUARY 


widens these extreme shade forms are doubtless driven out 
almost immediately by xerophytes, since intermediate or meso- 
phytic conditions are seldom seen where the soil is rock. Fur- 
thermore, the xerophytic conditions become much more extreme 
on rock bluffs than on clay bluffs. This is well illustrated at 
Starved rock (fig. 6), where the dominant tree vegetation is 


Fic. 6.—Xerophytic bluff of St. Peters sandstone at Starved rock, on the Illinois 
river, Showing conifers and other plants of dry rocks. Influence of erosive forces 
seen at the base. 


coniferous, consisting especially of the white pine (Pinus Stro- 
bus) and the arbor vitae ( Thuya occidentalis). The herbs and 
undershrubs here are also pronouncedly xerophilous, resembling 
the vegetation of the sand dunes, e. g., Selaginella rupestris, Cam- 
panula rotundifolia, Pellaea atropurpurea, Talinum teretifolium, 
Opuntia Rafinesquit, etc. The entire bluff flora down to the 
river's edge is xerophytic, except in shaded situations. 

When a stream in its meanderings ceases to erode at the base 
of a bluff, increased opportunity is given for plant life. Through 


—— 


ee eee 


Igor] PHYSIOGRAPHIC ECOLOGY OF CHICAGO 95 


surface wash the slopes become more and more gentle. Meso- 
phytic vegetation comes in at the foot of the bluff and creeps 
up as the slopes decrease. Finally the xerophytes are driven 
from their last stronghold, the top of the slope, and the meso- 
phytes have come to stay, at least until the river returns and 


7-—Ravine in the Niagara limestone at Lockport, showing the beginnings 


FIc, 
of a flood plain. 


enters upon another stage of cliff erosion. The growth of a 
ravine intoa valley with xerophytic bluffs is rapid, when expressed 
in terms of geology, but far less rapid when expressed in terms 
of vegetation. A ravine in the vigor of youth may develop so 
slowly that forest trees may grow to a considerable size with- 
out any perceptible change in the erectness of their trunks. 
Thus in figs. 2 and 5 it will be seen that most of the trees stand 
approximately vertical. But the activity of the erosive forces, 


96 BOTANICAL GAZETTE [FEBRUARY 


slow as it may be, is nevertheless revealed by occasional leaning 
or even falling trees. From the above it is easy to understand 
that cycles of vegetation often pass much more rapidly than 
cycles of erosion, but never more slowly. During one erosion 
cycle the mesophytic forest develops at least twice, once on 
the ravine slopes and then finally on the gentler slopes that 
betoken approach toward base level. 


Fic. 8.—General view of the Illinois valley near Starved rock, showing islands 
and an extensive flood plain with trees along the margin. Young islands in the fore- 
ground, older islands in the background, 


C. The flood plain— We may now follow the successive stages 
in the development of the flood plain vegetation. While the 
ravine is still young, as in fig. 2, there is no permanent stream, 
but merely torrents which remain but a short time. As the 
ravine deepens, widens, and lengthens, thus approaching the 
underground water level and increasing the drainage area, the 
water remains for a longer and longer time after each rainfall. 
As the ravine conditions thus become more and more hydro- 
phytic, the original flora, perhaps of shade mesophytes (as 
Impatiens), becomes replaced by amphibious shade plants, such 


lente eee 


1901 | PHYSIOGRAPHIC ECOLOGY OF CHICAGO 97 


as the common buttercup (Ranunculus septentrionalis\, Plantago 
cordata, various mosses, etc. Together with these forms algae of 
short vegetative period may be found in the wet seasons. When 
the ravine at last is sufficiently developed to have a permanent 
stream, a definite hydrophytic flora appears, consisting largely 
of algae (¢. g., Batrachospermum), aquatic mosses, and seed 


1G. 9.—Young island in the Illinois river at Starved reek (close view of island 
in foreground of fg. 8), seen from above, and showing the destructive action of tke 
iver, 


plants with finely dissected leaves and strong holdfast roots 
(such as Myriophyllum), though these latter plants are more 
characteristic of ponds. In the early phases of a stream, the 
currents are rapid and the vegetation (apart from lower forms) 
is sparse, by reason of the difficulty which plants have in secur- 
ing and retaining a foothold on the stream bed. This difficulty 
is due to the rapid erosion and consequent instability of the sub- 
stratum, as well as to the direct destructive action of the currents 
/ig. 7 shows one of these young streams, whose flora is sparse 


98 BOTANICAL GAZETTE [FEBRUARY 


Springs and spring brooks may be classed with ravine streams, 
but differ from them in the relative absence of erosion phenomena. 
This type of stream is uncommon in the Chicago area, though 
there are a few spring brooks near Chesterton. The water supply 
is much more constant than in ravine streams, and the shade of 
the ravines is often lacking. Besides the aquatics there may be 
mentioned a characteristic brookside flora, including such plants 
as Symplocarpus foetidus, Asclepias incarnata, Chelone glabra, Poly- 


Fic, 10.—Same island as shown in fg. 9, but seen from below, and showing the 
eee action of the river. Naked sand bar recently formed at the lower end of 
the island (left hand), Ambrosia farther Gite the right, willows on the older part of 
the island (extreme right). 


gonum sagittatum,and two or more species each of Eupatorium, 
Lobelia, Mentha, Lycopus, and Bidens. The most characteristic 
spring brook shrub is the alder (A/nus incana), though the exten- 
sive northern development of alder thickets has no parallel here. 

As the energy of the developing stream is checked, the con- 
ditions for plant life become more favorable. Inthe quiet pond- 
like waters of an older stream there may be found many of the 
aquatics that frequent the ponds and lakes. In fact the flora 
that is given later as characteristic of half-drained ponds and 
lakes (such as Calumet lake) may be transferred almost bodily 
to sluggish streams, such as the Calumet and Desplaines rivers. 

When streams are old enough and therefore slow enough to 


i a agency eas po tl gg: aaa 


| 
) 
| 


Igor] PHYSIOGRAPHIC ECOLOGY OF CHICAGO 99 


support a pond vegetation, they have become essentially deposit- 
ing rather than eroding streams, and we find there the develop- 
ment of a flood plain. While the river is still confined within 
narrow walls and may thus be called young, there may be 
embryonic patches of flood plain, representing alternations of 
erosion and deposition in the stream. ig. 7 shows such a con- 


IG. 11.—Flood plain of the Desplaines river at Glendon park, showing 
€ncroachment on the river. Willows in the foreground, cottonwoods farther back. 


dition of affairs ; though the stream is young and more destruc- 
tive than constructive at that point, there are to be seen small 
flood plain areas with their typical tree inhabitants. 

There is no place where flood plain development can be bet- 
ter studied than on growing islands in relatively rapid and yet 
essentially depositing streams, such as the Illinois river at 
Starved rock. Fig. 8 gives a general view of the Illinois islands 
and flood plain. In figs. g and zo the lower island (foreground 
of fig. 8) is seen close at hand. Any obstacle, such as a par- 
tially submerged tree trunk, serves to check the river current 
and cause a deposition of sand or silt, and before long a sand bar 


100 BOTANICAL GAZETTE [ FEBRUARY 
originates. As in the case of a sand dune, the bar itself becomes 
an obstacle to the currents and hence continually grows larger. 
The first vegetation, as on the lake beach, consists largely of 
annuals, especially the giant ragweed (Ambrosia trifida); rushes 
and sedges, some annual and some not, are also present but are 


less conspicuous. The perennials that manage to survive one 


aS" 


all 


ef 
4 
e 


Fic. 12.—Mesophytic flood plain forest in the bottoms of the Desplaines river at 
Riverside. Elms and basswoods. Rich herbaceous vegetation, consisting largely of 
Phi 


season are largely washed away in the winter and spring, so that 
in reality the vegetation is almost exclusively annual. The first 
woody plants to get a more or less permanent foothold here are 
willows (Salix nigra and S. longifolia). 

While islands of the above type gain more soil than they 
lose, a comparison of figs. 9 and zo shows that the river erodes 
above and deposits below. As a consequence these islands 
migrate down the river, as well as grow in area year by year. 


Igo! | PHYSIOGRAPHIC ECOLOGY OF CHICAGO 1o1 


Hence the upper part of the island is the oldest, as the vegeta- 
tion well shows. igs. 8 and zo show at the lower end the sand 
bar, which comes to a point and is so young or so exposed to 
submergence as to be barren of vegetation. Next comes the 
Ambrosia, then the willows, and finally a characteristic flood plain 


4 a 
— ae line 


Fic. 13.—Flood plain forest along Fraction run at Lockport, showing a rather 
striking collection of southern trees (see text). Coffee tree in the foreground. 


forest (background of fig. 8). The asymmetry of the river 
island vegetation is in striking contrast with the zonal symmetry 
of pond islands, as will be shown later (fig. 79). The cause is 
evident, viz., the relative lack of symmetry in river currents as 
compared with pond currents. 

The gradual encroachment of the land upon a stream through 
continuous deposition is well shown along the Desplaines river, 
and to a less complete degree along the Chicago river and Thorn 


102 BOTANICAL GAZETTE [ FEBRUARY 


>. creek. In the Desplaines bottoms the sand bar and island 
s { formations of the Illinois are largely absent, the currents being 


much less rapid. In the shallow water near the margin of the 
river are various hydrophytes, such as Sagittaria, Rumex verticil- 
latus, etc. The outermost fringe of land at ordinary low water 
is often almost as barren of vegetation as are the islands, but the 


Fic. 14.—Flood plain of the Calumet river near Chesterton, showing the begin- 
nings of terrace formation, indicated more by the falling elm than by the topography. 


soil is fine and hence makes a mud flat instead of a sand bar. 
Immediately after the spring freshets have gone, an alga vege- 
tation is frequently found on these flats, consisting especially 
of Botrydium and Vaucheria. Later in the season annuals, or 
even scattered perennials, may occur here, though the winter 
and spring floods uproot or bury most of this vegetation. The 
Ambrosia and willow vegetation soon appear as described 
above. The river maple (Acer dasycarpum) usually appears with 
or soon after the willows. After the willows the cottonwood 
(Populus monilifera) and the ash (Fraxinus Americana) soon 
come in. fig. rz shows an advancing flood plain of this type; 
willows are seen on the margin and cottonwoods farther back. 


Igor | PHYSIOGRAPHIC ECOLOGY OF CHICAGO 103 


Gradually the growing flood plain becomes dry enough to 
permit the germination and development of a true mesophytic 
flora. The trees named above, especially the willows, are 
largely replaced by others that seem better adapted to the 
changed conditions; among these are the elms ( Ulmus Americana 
and U. fulva), the basswood (Zilia Americana), the walnut and 


Fic. 15.—Terrace in the flood plain of the Desplaines river at Glendon park 
showing how a mesophytic flood plain may become xerophytic. The opposite bank 
shows deposition and flood plain enlargement (fg. 7/). 


butternut (Juglans nigra and J. cinerea), the pig-nut (Carya por- 
cima). In this rich flood plain forest there are many lianas 
climbing over the trees, e. g., greenbrier (Slax hispida), grape 
(Vitis spp.) , Virginia creeper (Ampelopsis quinguefolia),and poison 
ivy (Rhus T. oxicodendron). 

The undergrowth in these river woods is very dense and 
luxuriant, the alluvial character of the soil making it very fertile. 
Among the shrubs are the thorns (various species of Crataegus), 
the gooseberry (Ribes Cynosbati), and many others. ate Bee: 
baceous vegetation is dominantly vernal, the shade being too 
dense for a typical estival flora. Prominent among the spring 


104 BOTANICAL GAZETTE [ FEBRUARY 


flowering herbs are Trillium recurvatum, Phlox divaricata, Polemo- 
nium reptans, Hydrophyllum Virginicum, Mertensia Virginica, Col- 
linsia verna, Claytonia Virginica, Erythronium albidum, Arisaema 
triphyllum and A. Dracontium, Nepeta Glechoma, Isopyrum biterna- 
tum, Caulophyllum thalictroides, Viola cucullata, Galium Aparine. 
Other characteristic herbs are the nettles (Urtica gracilis, Lapor- 


Fic. 16.—An oxbow lake in the flood plain of Thorn creek. The willows are 
subsequent, dating back to a stream margin, while the shrub (Cephalanthus) and herb 
vegetation is associated with the present undrained condition. 


tea Canadensis), various umbellifers (Heracleum, Cryptotaenia, 
Sanicula, Osmorrhiza), and the parasitic dodder ( Cuscuta Gro- 
novit). fig. 12 shows a characteristic mesophytic flood plain 
forest along the Desplaines river; underneath the elms and bass- 
woods is seen a rich herbaceous flora, consisting largely of Phlox, 
which the picture shows in full bloom. 

In some of the bottom lands there is a rather striking collec- 
tion of trees, whose chief range is mainly southward. Fig. 13 
shows a flood plain tree group near Lockport, most of whose 
members are largely southern, viz., the coffee tree ( Gymnocladus 


1901] PHYSIOGRAPHIC ECOLOGY OF CHICAGO 105 


Canadensis), seen in the foreground, the papaw (Asimina triloba), 
the sycamore (Platanus occidentalis), and the hackberry (Celtis 
occidentalis). In other flood plains there may be found the 
mulberry (Morus rubra), the red bud (Cercis Canadensis), the 
buckeye (Aesculus glabra), and the tulip (Liriodendron Tulipifera). 
None of these trees are common in our district, and only Celtis 


Fic. 17.—A dead oxbow lake in the flood plain of Thorn creek. A willow still 
remains at the right, while the shrubs (Cephalanthus) have closed in upon the lake. 


may be regarded as frequent. These relatively southern trees 
are found not only along the Desplaines and its tributaries, 
where there is supplied a continuous habitat along the river 
southward, but also along the Calumet and its tributary, Thorn 
creek. The occurrence of the tulip tree is full of interest, since it 
has been found thus far chiefly (perhaps only) in the vicinity of 
the dunes. Its occurrence has been noted especially at Chester- 
ton along a small stream which empties into Lake Michigan at 
that point; the tulip tree has also been found away from present 
Streams, but apparently in old valleys whose streams have been 
diverted by dune activity. The confinement of these southern 


106 BOTANICAL GAZETTE | FEBRUARY 


trees to flood plains is not strange, since in such habitats are given 
the most congenial conditions that can be found in our area. 
The vegetation on flood plains is not always as described 
above. Sometimes meadows are found instead of forests; this 
condition is particularly well shown along Thorn creek. Fig. 18 
shows a stretch of meadow of this type. Besides various grasses 


Fic. 18.—Flood plain of Thorn creek near Glenwood, showing a meadow instead _ 
of aforest. At the center is an uneroded island, detached from the morainic main- 
land, seen at the left. The vegetation of the island is similar to that of the morainic 

plands. 


(such as Poa pratensis and Agrostis alba vulgaris), there are often 
other plants in abundance, ¢. g. Thalictrum purpurascens, Fragaria 
Virginiana, Anemone Pennsylvanica. The ecological meaning of the 
meadow is not clear. Probably mowing or grazing is responsible 
for the failure of a mesophytic forest to develop. Extensive 
thorn (Crataegus) thickets sometimes occur in these meadows 
and probably betoken the beginning of a mesophytic forest. 
Extensive and apparently natural meadows are found in the 
Calumet valley. 

As we have seen, the climax type of vegetation on the flood 


Igor] . PHYSIOGRAPHIC ECOLOGY OF CHICAGO 107 


plain is the mesophytic forest, but here, as well as on the river 
bluffs, the climax may be but temporary. Retrogression is 
almost sure to come in connection with terrace formation. 
While it is true that deposition is the main feature of flood plains, 
it is also true that erosion has not ceased; the downward cutting 
of the river once more causes vertical banks, though this time 
in its own flood plain. This action is seen in fig. rg which 
shows the beginning of the new erosive phase, and its indication 
in the falling elm. There has doubtless been lateral erosion here 
also, since elms are not usually marginal trees. /%g. 75 shows 
the erosion of the flood plain still farther advanced; this bank is 
just opposite the willow vegetation shown in fig. zz, hence there 
is deposition on one side and cutting on the other. A river may 


thus swing quite across its flood plain, destroying all that it has 


built, including the mesophytic forest. Not only is the vegeta- 


tion destroyed directly, as shown in fig. 14, but also indirectly, 


Since the lowering of the river causes the banks to become more 
xerophytic. In place of the herbaceous mesophytes, Equisetum 
and other relatively xerophytic forms may appear, though the 
trees usually live until directly overthrown by the river. _ 

One more phase of river activity may be briefly sketched. 
In meandering over a flood plain, serpentine curves or oxbows 
are frequently formed. In time the river breaks across the 
peninsula and the oxbow remains as a crescentic lake. The 
conditions radically change almost immediately, and the river 
life is replaced by pond life. The change is even more striking 
on the margins, where the old plants pass away and the forms of 
undrained Swamps come in. fig. 76 shows the remnant of one of 
these oxbows; on the farther side are old and dying willows, 
trees that look back to the well drained river margin. On either 
Side of the pond are seen clumps of the button bush ( Cepha/lan- 
thus occidentalis), one of the most characteristic plants of 
undrained swamps. Thus the willows are antecedent and the 
button bush subsequent to the formation of the cut-off. Fig. 77 
shows a portion of the same, in which the willows and even the 
pond itself have gone, and only the marginal button bush is left, 


108 BOTANICAL GAZETTE [FEBRUARY 


though in this case, the margin occupies the center of the original 
pond. Near Starved rock an extinct oxbow lake on the flood 
plain of the Illinois river contains an extensive patch of Sphag- 
num and Osmunda, among the most characteristic plants of 
undrained swamps. There are many undrained swamps, some 
with tamaracks, in the Calumet valley. The future of these 
swamps is like that of other swamps, and will be described in 
the next section. fig. 78 shows a morainic island in the Thorn 
creek flood plain; the stream has meandered but has thus far left 
this detached fragment of the morainic mainland with a large 
part of its original flora. 

In closing the section on rivers, all that is needed is to 
emphasize again the idea that the life history of a river shows 
retrogression at many points, but that the progressions outnum- 
ber the retrogressions. Not only this, but retrogressive phases 
are relatively ephemeral. Thus a river system, viewed asa whole, 
is progressive, and through all its vicissitudes there is an ever 
increasing area of mesophytic forest. When the theoretical 
base level is reached there seems to be no apparent reason why 
mesophytic forests should not be developed throughout most of 

‘ss the great plain. 
[Zo be concluded.) 


eo: eee iar 


" I90r) 


UNDESCRIBED PLANTS FROM GUATEMALA AND 
OTHER CENTRAL AMERICAN REPUBLICS. XXII.: 
JOHN DONNELL SMITH. 

(WITH PLATE 1) 

Xylosma Turrialbanum Donn. Sm.— Folia magna oblongo- 
elliptica longe obtuseque acuminata in basin obtusiusculam aut 
retusam angustata membranacea integerrima glabra minute pel- 
lucido-punctata venis venulisque transversis subparallelis. Flo- 


rum masculinorum pedicelli fasciculato-congesti floribus 4-5-plo 


longiores ad tres partes longitudinis articulati. Sepala5. Sta- 
mina I5 sepalis paulo breviora. Discus subinteger, glandulis 
nullis. 

Ramis spiniformibus 1-7 longis armatum, his florum fasciculis saepe 
ornatis, Folia g—13°™ longa 3~5™ lata pellucida basi nonnunquam minute 
bilobata, nervis lateralibus utrinque 6-8 subtus prominulis, petiolis 5-10™ 
longis. Pedicelli indefiniti g-11™™ longi pubescentes, bracteis pilosis, brac- 
teolis rudimentariis. Sepala oblongo-ovata 3™ longa parce pubescentia. 
Filamenta 2™™ longa infra medium pubescentia, antheris elliptico-globosis 
dorso affixis. Discus annularis carnosus 2™ diametralis ciliolatus ceterum 
glaber. Flores feminini fructusque ignoti.— Inter species Americanas foliis 
integerrimis insigne. 

“In fundo Aragon vocato in declivibus Turrialbanis sito, Costa Rica, alt. 
630", Jan. 1899, Pittier, no. 7518 Pl. Guat. &c., qu. ed. Donn, Sm. (n. 13217 
herb. nat. Cost,). 


Monnina saprogena Donn. Sm. (§ Hepeanpra Chodat, Bull. 
Herb. Boiss. 4: 243.)—Folia estipulata crassiuscula glabra lan- 
ceolata-elliptica acuminata in petiolum attenuata, nervis lat- 
eralibus utrinque 3~4 erecto-patentibus distinctis ante marginem 
€vanidis supra subimmersis, venis obsoletis. Racemi paniculati, 
floribus pedicellos paulo superantibus. Sepala interiora per duas 
partes connata. Carina subintegra. Discus conspicue unilat- 
€ralis, ovario dimidiato-elliptico glabro uniloculari. Fructus 

*Continued from Bor. GAZ., 27: 443. 1899. 

100 


110 BOTANICAL GAZETTE [FEBRUARY 


inaequilateralis oblongo-ovatus acutiusculus bicarinatus et bico- 
status indistincte areolatus. 

Arbuscula (Tonduz in schedula), caulibus teretibus glabris. Folia 6-g™ 
longa 2-3 lata, petiolis 6-7™™ longis. Paniculae terminales et subterminales 
puberulae laxiflorae 10-16 longae, bracteis ovatis 4™™ longis ciliatis cito 
deciduis, pedicellis 3-4™" longis, floribus 6™™ longis. Sepala pubescentia 
ciliataque, exteriora inferiora ovalia, superius majus ovatum 2.5™" longum, 
alae orbiculares 5™" diametrales extus pube punctatae e basi distincte 
nervosae. Carina ad apicem biplicata, petalis interioribus intus cum vagina 
staminea cano-pilosis, limbo lingulato 3™" longo. Stylus complanatus ovario 
bis longior. Fructus glaber sublaevis 6™™ longus 2.5™" latus, disco gibboso 
glanduliformi. 

vIn truncis putridis, La Palma, Costa Rica, alt. 1542™, Aug. 1898, Tonduz, 
no, 7406 Pl. Guat. &c. qu. ed. Donn. Sm. (n. 12501 herb, nat. Cost.). 


MyropiA GuATEMALTECA Donn. Sm., Bort. Gaz. 16: 2. 1891.— 
Hujus speciei e floribus nondum plane evolutis descriptae speci- 
mina typica cum Quararibea Guyanensi Aubl. satis congruunt. 
Ejusdem plantae exempla Guatemalensia nuper a Barone de 
Tuerckheim transmissa et flores evolutos et fructus praebent. 


Heliocarpus Donnellsmithii Rose.— Mature leaves nearly 
orbicular, not lobed, rounded at base, shortly acuminate (tips 
mostly broken), glabrous and shining ‘above, nearly glabrous 
below, 3 to 5-palmately veined from the base, somewhat glandu- 
lar-toothed especially at base; inflorescence a large spreading 
panicle; sepals not appendaged; stipe slender, 5™™ long; body 
of fruit oblong, 5™" long, somewhat hairy, becoming glabrate 
and rugose in age. 

Near Arenal, Department Alta Verapaz, Guatemala, alt. 1500™, April, 
1889, John Donnell Smith, n. 1722 (type) ; Coatzacoalcos, Isthmus of 
Tehauntepec, State of Vera Cruz, Mexico, 1895, Charles L. Smith, n. 1002. 

This species is very different from the numerous Mexican species, and is 
near H. Americanus Schumanni Baker, but has more glabrous leaves and an 


absence of hairs both on leaves and inflorescence. Mr. Baker’s variety 
seems to me to deserve specific rank. 


EXPLANATION OF PLATE I.— Branch two thirds natural size; fruit X 3- 


Villaresia Costaricensis Donn. Sm.— Folia oblongo-elliptica 
utrinque praesertim apice acuminata submembranacea diaphana 


BOTANICAL GAZETTE, XXX/. PLATE I, 


) f 


: 


B. Meisel. lith. Boston 


HELIOCARPUS DONNELLSMITHIHT Rose, n.sp. 


1901] UNDESCRIBED PLANTS FROM CENTRAL AMERICA rit 


pellucido-punctulata integra, costa cum nervis lateralibus utrinque 
4-5 venisque subtus conspicua. Racemi extra-axillares termi- 
nalesque foliis breviores spiciformes, glomerulis subsessilibus 
inferne dissitis. Sepala ovata pilosa. Petala glabra oblongo- 
elliptica sepalis vix bis longiora. Stamina petala paene aequan- 
tia. Ovarium pilosum ovoideum t-loculare. Drupa magna 
olivaeformis. 

Ramuli petiolique recentiores puberuli. Folia glaberrima aetate provec- 
tiore subcoriacea g-12.5°" longa 4.5-5°™ lata, petiolis crassis canaliculatis 
5-7™ longis. Racemi pubescentes singuli, floriferi 3-5°™ longi, fructiferi 
incrassati usque ad 8 longi, floribus 3-8 in pedicello crasso 1-2™™ longo ses- 
silibus. Sepala 1.5™™ longa. Petala quincuncialia apice cuspidata inflexo- 


aequans, stigmate obliquo nudo. Drupa pedicello vix ullo insidens 2.5 
longa 1.5°™ lata atque crassa apiculata in sicco nigrescens et sete taee: semi- 
septo 5™™ lato, seminis testa pallida et atro-venosa.— Genus in America cis 
circulum aequinoctialem adhuc non obvium. 

“In silvis prope Copey, Prov. Cartago, Costa Rica, alt. 1800”, Febr. et 
Mart. 1898, Tonduz, nn. 11664, 11995, 11791 herb. nat. Cost. 


Blakea tuberculata Donn. Sm. (§ Eusiakea Triana.)— Rami 
crassi teretes cum petiolis pedunculis bracteis et foliorum 
tergo ferrugineo-furfuracei et -strigillosi. Folia ovato-orbicularia 
abrupte caudato-acuminata basi rotundata supra glabra. Flores 
fasciculati brevissime pedunculati inter maximos. Bracteae 
exteriores usque ad mediam connatae calycis tubum paulo super- 
antes, interiores breviores omnino connatae. Calycis limbus 
ovali-cylindricus tubum aequans intus coloratus et tuberculatus, 
lobi retroflexi limbo bis longiores. Petala utrinque tuberculata 
cum staminibus basi limbi inserta. Genitalia limbo inclusa. 


Folia uniuscujusque paris aequalia 13-21™ longa 10-15% lata coriacea 


ers 
olis robustis 4~6°™ longis. Pedunculi 3-6-fasciculati crassi 6-10™™ longi. 
Flores diametro 6. 5°", bracteis crassis opacis, exterioribus late orbicularibus 
2 longis uninerviis siccitate retroflexis et margine involutis, interioribus 1.5°™ 
longis margine crispato- plicatis. Calycis tubus campanulatus glaber, limbus 
rm altus 12™ latus carnosulus extus strigillosus in alabastro petala usque ad 


r1l2 BOTANICAL GAZETTE [FEBRUARY 


duas partes includens, lobi oblongo-ovati herbacei nervosi supra furfuracei 
subtus strigillosi jam ante anthesin e basi arcte retroflexi. Petala glabra 

ongo-obovata 2.5—3™ longa late unguiculata carnosula rosea. Stamina in 
sicco rubescentia, antheris pendulis dolabriformibus acute calcaratis 8™™ 
longis filamenta subaequantibus. Ovarium diametro 9™™, centro in rostrum 
oblongo-conicum 5™™ longum apice denticulatum producto, stylo gracili 1 
longo, stigmate punctiformi. Baccae non suppetunt.— 2. grandifiorae 

emsl. proxima. 

Linas propter exsiccationis pressionem diruptus limbum in herbario 6- 
fissum lobis appendiculatum simulat. 

Ad pascuorum margines prope La Palma, Prov. San José, Costa Rica, 
alt. 1550™, Aug. 1898, Zonduz, n. 7363 Pl. Guat., &c., qu. ed. Donn. Sm. 
(n. 12435 herb. nat. Cost.) 


Anguria ovata Donn. Sm.—Tota glaberrima. Folia omnia 
simplicia integra membranacea concoloria ovata cuspidato- 
acuminata dimidio longiora quam latiora ad basin rotundam levi- 
ter retusa, nervis basilaribus 5, lateralibus utrinque 3. Flores 
masculini. pauci subcapitati. Calyx lineari-cylindricus. Petala 
utrinque dense furfuracea. Antherae rectae lineares, appendice 
glabra. 

Caulis gracilis striatus. Folia 8.5-9°™ longa viridia, venulis reticulatis 
diaphanis, petiolis 3-5 longis. Cirrhi tenues striati. Pedunculi masculini 
Striati 12~-15°™ longi apice brevissime racemosi et 8—12-flori, pedicellis 1-2™™ 
longis. Calyx basi rotundus apice haud constrictus 14™™ longus 3™™ latus, 
dentibus obtuse ovatis 1™™ longis. Petala suborbicularia 6™™ longa 4-5" 
lata exunguiculata enervia lateritia. Antherae to™™ longae 1.5™™ latae, 


appendice 0.5™" longa. Flores feminini fructusque desunt.— A. ongipedun- 
culatae Cogn. ptoxima. 


In dumetis ad Las Vueltas, Tucurrique, Costa Rica, alt. 635™, Feb. 1899, 
Tonduz, n, 13006 herb. nat. Cost. 


Gurania Tonduziana Donn. Sm. (§ F. 1. Cogn. in DC. Monogr. 
Phan. 3: 694.)— Folia circumscriptione cordiformia paulo 
longiora quam latiora 5-nervia profundissime 7-partita, seg- 
mentis § interioribus oblanceolato-linearibus, exteriore utroque 
oblongo et aurito.. Pedunculi masculini foliis superati, floribus 
brevissime spicatis. Calycis pubescentis segmenta linearia tubo 
2-3-plo longiora. Antherae oblongae, appendice minuta del- 
toidea. 


1901] UNDESCRIBED PLANTS FROM CENTRAL AMERICA 13 


Caulis robustus striatus cum petiolis cirrhis pedunculis pubescens. Folia 
24 longa, pagina superiore scabriuscula, inferiore pallidiore pubescente 
conspicue reticulato-nervosa, margine subundulata remote minuteque spinu- 
loso-denticulata, nervo basilari utroque exteriore imum sinum marginante, 
segmentis acuminatis, terminali 21°" longo ultra medium 5™ lato basi 13™™ 
lato ceteris interioribus paulo majore, utroque exteriore inequilaterali 6.5°™ 
longo 3™ lato, sinibus inter segmenta rotundis a sinu basilari 2-3°™ distanti- 
bus, hoc sub-rectangulari 3°" profundo 5°™ lato. Pedunculi 17°™ longi, spica 
8™™ longa. Calycis tubus ovalis 1°™ longus 6™ latus, segmenta parce 
pubescentia 22~25™™" longa basi1™™ lata enervia miniata. Petala conniventia 
lineari-lanceolata g-1o™™ longa 2™™ lata enervia extus dense glandulari- 
papillosa intus furfuracea. Antherae 7™ longae 2™™ latae tertia parte repli- 
Catae, connectivo loculis angustiore, appendice vix 0.5™" longa. Flores 
feminini fructusque ignoti.— . Makoyanam Cogn. florum indole acce- 
dens foliis insigniter recedit. 

“In sylvis ad Shirores, Talamanca, Costa Rica, alt. 1oo™, Feb. 1895, 
Tonduz, n. 9332 herb. nat. Cost 


Sciadophyllum systylum Donn. Sm.— Ferrugineo-pubescens. 
Foliola 7-9 elongato-oblonga longe acutissimeque caudato- 
acuminata basi obtusa subcoriacea praeter costam tandem glab- 
rescentia, nervis patulis. Umbellae in racemum simplicem 
cylindricum dispositae. Styli in columnam quam bacca tertia 
parte breviorem omnino coaliti, stigmatibus 5-6, seminibus 
abortu paucioribus. 

Totum pube detergibili stellata interdum tomentulosa vestitum. Stipulae 
Coriaceae lineares 57.5% longae. Petiolus teres 16-27 longus, petiolulis 
Canaliculatis, foliolo intimo maximo 14-2 5°™ longo 4-7™ lato, cauda 2.5-3.5°" 
longa, petiolulo 4-7 longo, foliolis lateralibus cum petiolulis sensim decres- 
centibus, infimo utroque 6-14.5™ longo, petiolulo 2-2.5™ longo, nervis lat- 
eralibus utrinque 12-18 prope marginem arcuate conjunctis. Racemi 20— 
35°" longi, ramis satis approximatis 1.5—4.5°™ longis, pedicellis circa Io et 
5-8"™" longis. Baccae ovali-globosae 3-4™" longae 5—6-angulares, juniores 
pube stellata punctatae, stylis 2-2.5™™ longis, stigmatibus vix 0.5™™ longis 
tadiantibus, seminibus 3™° longis, 1-3 saepius abortivis. Flores ignoti.— 
Species stylis longis totis coalitis insignis. 

In sylvis ad El Alto de La Palma, Costa Rica, alt. 1542™, Aug. 1898, 
Tonduz, n. 7395 Pl. Guat. &c., qu. ed. Donn. Sm. (n. 12488 herb. nat. Cost.), 


eopanax pycnocarpum Donn. Sm.—Stellato-pubescens. Folia 
Simplicia integra crassa pube punctulata mox glabrescentia 


114 BOTANICAL GAZETTE [FEBRUARY 


nitidaque ovalia utrinque rotundata triplinervia longe petiolata. 
Capitula fructifera spiciformia oblongo-globosa pedicellis longi- 
ora inter maxima in racemos 3-nos abbreviatos pauciramos dis- 
posita, baccis compactis stylis liberis 6-7, seminibus abortu sae- 
pius paucioribus, albumine aequabili. 

Folia 16-20 longa 8-11™ lata, nervis subangulo angusto ascendentibus, 
basili utroque exteriore 8™ a basi distante, lateralibus utrinque 3 ultra medium 
ortis, venis transversis inter marginem et nervum utrumque basilarem subtus 
conspicuis circa 10, petiolis pubescentibus 1o-12™ longis. Racemi pubes- 
centes 7—-9™ longi, ramis crassis 8-11™™ longis, inferioribus alternis, supremis 
4-nis, capitulis 2.5—3.2°™ longis 1.8-2.2™ latis, rhachi 1~1.5°™ longa, bracteolis 
cuspidato-ovatis ciliatis. Baccae subglobosae pressione mutua angulatae, 
nitidae in sicc. pallide flavescentes numerosissimae 8™™ longae 6™™ latae, 
stylis stellatim recurvatis 1.5"" longis, seminibus oblongis 3—gonis 5™™ longis 
2™™" crassis. Flores deficiunt.—Ad O. cafitatum Decne. et Planch. foliis ad 
O. macrocephalum Seem. capitulis accedit. 

“in sylvis ad Copey, Prov. Cartago, Costa Rica, alt. 1800", Febr. 1898, 
Tonduz, n. 11933 herb. nat. Cost. 


Chomelia microloba Donn. Sm.—Inermis. Folia oblongo-ovata 
aut- -elliptica obtuse acuminata basi acuta aut obtusa coriacea 
praeter nervorum axillas subtus barbatas glaberrima. Pedunculi 
terminales bini gracillimi, cymis bifidis multifloris, floribus ebrac- 
teolatis. Calyx glaber, ore subintegro. Corollae extus pubescentis 
tubus infundibuliformis calyce triplo longior, lobis induplicato- 
valvatis. 


Ramuli recentiores cum stipulis cuspidato-triangularibus 3-4™™ longis 
persistentibus et petiolis 4-6™" longis pubescentes. Folia 5.5—10™ longa 
3-4.5°™ lata, juniora e basi rotunda magis ovata, nervis lateralibus utrinque 
4-5. Pedunculi glabri 2.5—3.5°™ longi, cymis glabris 3.5 latis circa 19—-25- 
floris, floribus sessilibus. Calyx cylindricus 3™™ longus 1.5™™ latus, denticulis 
vix 0.3" longis obtusis. Corollae intus glabrae tubus 8—10™™ longus, lobi 
obtuse ovati 3™" longi. Antherae semiexsertae lineares 2™ longae ad tertiam 
partem longitudinis affixae. Ovarium calycis duas partes aequans, stigmatibus 
1™" longis, ovulis linearibus 1™ longis. Drupa in speciminibus suppetenti- 
bus deficit.—Ab omnibus congeneribus calyce subtruncato differt. 

“In sylvis litoralibus ad Santo Domingo de Golfo Dulce, Costa Rica, Febr. 
1896, Tonduz, n. 7048 Pl. Guat., &c., qu. ed, Donn. Sm. (n. 9874 herb. nat. 
Cost. 


Igor] UNDESCRIBED PLANTS FROM CENTRAL AMERICA its 


Faramea trinervia K. Sch. et Donn. Sm.—Folia nitida obo- 
vato- -oblonga vel oblonga contracto-acuminata auriculata subses- 
silia, nervo utroque basilari percurrente, lateralibus angulo 
subrecto a costa divergentibus. Cyma foliaceo-bibracteata tri- 
chotomo-decomposita, floribus ebracteolatis. Calycis limbus 
denticulatus discum aequans. 

Glaberrima. Folia pergamenea diaphana laete viridia 20-25™ longa 
medio 6—9.5°™ basi 1.5—3°™ lata, suprema lineari-oblonga 11™ longa 3™ lata, 
costa utrinque prominente, nervo a basi utrinque prodeunte conspicuo api- 
cem attingente a margine 5-10™™ distante, nervis lateralibus primariis utrin- 
que circa 14, nervum percurrentem attingentibus, petiolis 2-3" longis. 
Stipulae 6-8™" longae in vaginam semiconnatae parte libera rotundatae 
aristatae. Pedunculi singuli ramulos terminantes 4—6,5°™ longi apice brac 
teis ante anthesin caducis instructi. Cyma 4.5—6.5™ alta, ramis late patulis 
complanatis, iis infimis 4~-5.5°™ longis, pedicellis accrescentibus 5—11™™ lon- 
gis. Calycis limbus ovarium aequans usque ad medium acute denticulatus. 
Ovarium post anthesin pedicello paulo crassius cum disco aequilongo trun- 
cato adjecto 1.5™° longum. Corolla et fructus desunt.—A ceteris speciebus 
haec nervatione insigniter differt. 

In sylvis ad Boca Zhorquin, Falamanca, Costa Rica, Mart. 1894, Zonduz, 
n. 8571 herb. nat. Cost.; in sylv eee Talamanca, alt. 200", Apr. 
1895, Tonduz, n. 9583 herb. nat. rs 


FARAMEA TRINERVIA Suerrensis Donn. Sm.— Folia obovato- 
vel oblongo-elliptica in caudam gracilem contracta infra medium 
in petiolum angustata, suprema lanceolata. 

Frutex 3-4™altus, ramosus. Foliorum cauda 1.4—2™ longa, petioli 1-1.5°%™ 
longi. Bracteae lanceolatae tenuissime elongatae 6™ longae 12™™ latae. 
Florum nondum satis evolutorum pedicelli 2—5™™ longi, calycis limbus 1™ altus, 
dentibus triangularibus, corolla intense cyanea in sicc. nigrescens 6"™ longa 
tubulosa usque ad mediam paene lobata, filamenta longiuscula prope basin 
corollae inserta, antheris 3™™ longis exsertis, ovarium obovatum cum disco 
Conico adjecto 2™™ longum. Fructus cyaneus transversim depresso-ovalis 
15™" longus 11™ crassus, semine conformi 11™™ longo 8™™ crasso subtus 
usque ad medium transversim bipartito, testa pallide scariosa. 

vIn sylvis profundis ad fundum Suerre dictum, Llanuras de Santa Clara; = 
Comarca de Limén, Costa Rica, alt. 300, Febr. 1896, Donn. Sm., 0. 6589, 
Pl. Guat., &c., qu. ed. Donn. Sm 


Parathesis glabra Donn. Sm.—Omnibus in partibus maculata 
€t praeter corollam glabra. Folia oblongo-elliptica acuminata 


116 BOTANICAL GAZETTE [FEBRUARY 


in petiolum attenuata integerrima subdiaphana utrinque dense 
maculata, nervis crebris patulis. Paniculae axillares foliis 
breviores, pedunculo filiformi, pedicellis ad apicem ramorum 
primariorum subumbellatis. Calycis segmenta elongato-triangu- 
laria. Corollae segmenta linearia. Antherae filamentis bis 
longiores prope basin affixae per totam longitudinem dehis- 
centes. ; 

Folia 8-14 longa 3.5—7™ lata maculis punctata et striatula, nervis inter 
se 4-6™™ distantibus, petiolis canaliculatis 0.8-1.5°" longis. Paniculae cum 
pedunculo 2.5—4°™ longo adjecto 5-7.5™ longae, ramis solitariis 7-15™™ longis, 
pedicellis 3-6-fasiculatis 5—7™" longis, bracteis bracteolisque lanceolato- 
linearibus parvis, alabastris lanceolato-conicis. Calycis segmenta 1™™ longa. 
Corollae tubus 1™™ altus, segmenta e basi 1™™ lata attenuata usque ad 7™™ 
longa staminibus bis longiora revoluta intus praesertim marginibus sordide 
pubescentia. Ovarium conicum calyce brevius parce maculatum, stylo 4.5™™ 
longo. Fructus desideratur.—Avrdisiam ramifioram Oerst. habitu simulans. 

YIn sylvis ad Copey, Prov. Cartago, Costa Rica, alt. 1800™, Febr. 1898, 
Tonduz, n. 11714 herb. nat. Cost. 


MACROSCEPIS CONGESTIFLORA Donn. Sm., Bor. Gaz. 25:149. 
1898.— Sub hoc nomine eandem plantam atque eodem numero 
indicatam incaute descripsi, quam typicam M. pletstanthae 1. c. 
20:543. 1895, jam edideram. 


Markea leucantha Donn. Sm.—Praeter filamenta omnibus in 
partibus glaberrima. Folia coriacea nitida obovato-oblonga apice 
rotundata a medio in petiolum attenuata. Flores apice ramorum 
annotinorum aggregati pauci. Calyx parvus teres coriaceus, 
lobis oblongo-ovatis brevissime cuspidatis. Corolla calyce 5- 
plo longior, tubo extra calycem sensim ampliato late infundibu- 
lari. Genitalia exserta. 

Frutex epiphyticus 2-3™ longus verrucosus, ramulis hornotinis ‘quadran- 
gularibus sub apice ramorum plerumque ortis. Folia plerumque geminatim 
aut ternatim approximatis 7-15 longa 2.5-5.5°™ lata, recentiora magis 
oblanceolata et acutiuscula, nervis lateralibus utrinque 5-8 subtus tantum 
conspicuis, petiolis crassis 4-12™" longis. Pedicelli brevissime racemosi in 
speciminibus suppetentibus 3-8 circa 10™™ longi, bracteis subulatis 3™™ lon- 
gis deciduis. Calyx intus nervosus 11-13™" longus usque ad medium fissus. 
Corolla alba (Cooper in schedula) 5-nervia reticulata 5.3-6.3°™ longa, tubo 


oe erm ince 


aN, ~*~ “ 


a 


tgor] UNDESCRIBED PLANTS FROM CENTRAL AMERICA it7 


3.8-4.6" longo quam faucium latitudo bis longiore, lobis semiorbicularibus. 
Stamina ad 8™™ supra basin corollae inserta 4~-4.8™ longa, filamentis ad basin 
versus incano-barbatis, antheris paulo infra mediam affixis 6™" longis 2.5™™ 
latis. Ovarium ovoideum 4" altum atque latum, stylo 4.3-5"™ longo, stig- 
matis obconoidei lamellis semiorbicularibus. Bacca desideratur.—Species 
calyce parvo insignis. 

Estrella, Prov. Cartago, Costa Rica, alt. 1800™, Jun. 1887, Coofer, n. 5887 
Pl. Guat. &c., qu. ed. Donn. Sm.; Prope Juan Vifias, Prov. Cartago, alt. 1140", 
Maj. 1890, Tonduz,n. 1845 herb. nat. Cost. 


Alloplectus macranthus Donn. Sm.--Omnibus in partibus 
strigilloso-pubescens. Folia maxima oblongo- vel obovato- 
elliptica apice acuminata in petiolum attenuata et decurrentia 
inaequilatera subintegra. Pedicelli racemosi bracteis breviores 
calycem subaequantes, floribus maximis numerosissimis coarcta- 
tis. Calycis segmenta breviter connata subaequalia oblonga 
obtusa integra. Corolla calyce bis longior erecta late infundi- 
bularis, limbo obliquo, lobis inaequalibus rotundatis denticulatis. 

Caulis pollicis crassitudine in arborum truncis scandens. Folia 23-45% 
longa 10-19 lata, nervis lateralibus utrinque 7-8, petiolis crassis rugosis 
Canaliculatis 4—7™ longis. Racemi aetate provectiore recurvi 2—4°™ longi. 
Bracteae herbaceae ovales aut oblongo-ellipticae 3-4, 5 longae. Pedicelli 
2.5-3™ longi, demum subsecundi. Calyx herbaceus, segmentis circa 2.5™ 
longis 8-1o™™ latis. Corolla puberula ex schedula Tuerckheimiana extus 
Candida intus maculato-aurantiaca 6°" longa, tubo 4.2 longo a basi saccata 
6™" lata in fauces 2° latas sensim ampliato, lobis posticis 1°™ anticis 7™ 
longis. Filamenta in vaginam brevem dilatata, antheris liberis oblongis 6™™ 
longis. Disci glandula solitaria. Ovarium dense pubescens oblongo- 
ovatum 6™™ longum, stylo 1.6™ longo, stigmate concavo. Fructus calyce 
aucto cinctus ovalis 1.7°™ longus 1™ crassus, placentis rubris, seminibus 
oblongo-ellipticis 1™" longis.—Ab A. circinnato Mart. foliis et inflorescentia 
Proximo recedit praecipue corolla infundibulari altero tanto longiore. 

In fundo Cubilquitz vocato, Depart. Alta Verapaz, Guatemala, alt. 350”, 
Jul. 1900, A. von Tuerckheim, n. 7642 Pl. Guat. &c., qu. ed. Donn, Sm.— 
Huic probabiliter referenda sunt specimina a Zonduz in Costa Rica lecta et 
Sub n. 13042 distributa. Haec pedicellos densissime aggregatos brevissime 
racemosos bracteas calycesque erubescentes praebent, corollis tamen carent. 


Columnea sulfurea Donn. Sm. (§Eucorumnea Benth. et 
Hook. )—Pilis aspersa. Folia disparia subtus vinosa elongato- 
elliptica utrinque acuminata inaequilatera basi valde obliqua. 


118 BOTANICAL GAZETTE [FEBRUARY 


Pedunculi solitarii aut bini petiolo longiores calyce breviores. 
Calycis segmenta vinosa attenuato-linearia tubi corollini duas 
partes aequantia. Corolla flava infundibulari-tubulosa leviter 
arcuata, limbo postico quam tubus breviore quam anticus por- 
rectus altero tanto paene longiore, galea truncata. Ovarium 
dense pilosum. 

Epiphytica, caule crasso parum ramoso erubescente, internodiis 1.5-3.5°™™ 
longis, superioribus et petiolis pedunculis calycibus densius rubro-pilosis. 
Folia discoloria utrinque sparsim bulboso-pilosa ad basin latere altero obtusa 
altero valde exciso acuta, uniuscujusque paris folio florali 7.5-11.5° longo 
2.5-4™ lato, minore conformi 3-4°™ longo, petiolis 3-7™™ longis. Pedunculi 
plerumque singuli 9-15™" longi. Calycis segmenta sublibera 2.5°" longa. 
Corolla ex sched. Tuerckheimiana sulfurea sparsim pilosa 6.8 longa basi 
5 ea a, 7™™ lata, tubo basi gibbo, limbo postico 3™ longo, galea 1.5™ 
longa atque lata integra, lobis lateralibus angulo recto subtriangularibus 1™ 
longis obtusis, limbo antico lineari-oblongo 1.5°" longo. Stamina 5.8 longa, 
antheris oblongis 3™™ longis, staminodio 4™™ longo. Disci glandula solitaria 
crassa bidentata. Ovarium ovoideum 2™™ altum, stylo rubro 6.5™ longo ad 
apicem versus pubescente, stigmate bilamellato. Fructus ignotus. 

“Cubilquitz, Depart. Alta Verapaz, Guatemala, alt. 350", Febr. 1900, /7. 
von Tuerckheim, n. 7646 Pl, Guat. &c., qu. ed. Donn. Sm. 


COLUMNEA MICROCALYX macrophylla Donn. Sm.— Folia mem- 
branacea pubescentia oblongo-elliptica vix acuminata basi cordata 
inaequilatera, uniusque paris majus usque ad 5°™ longum et 2™ 
latum, alterum triente minus. Calyx virescens corollae 6.7™ 
longae quintam partem aequans. 


In sylvis prope Las Vueltas, Tucurrique, Costa Rica, alt. 650-700™, Dec. 
1898, Zonduz, n. 12932 herb. nat. Cost 


Napeanthus repens Donn. Sm.— Herba repens acaulis strigillo- 
so-pubescens. Folia Opposita subrosulata petiolata tenuimem- 
branacea oblongo-elliptica utrinque acuminata dentata. Pedicelli 
fasciculati filiformes. Calycis segmenta sublibera 3-nervia. 
Corolla rotata, tubo brevissimo, faucibus explanatis, limbo alte 
fisso subbilabiato. Stamina 4, loculis ovoideis divergentibus 
distinctis. Ovarium corollae tubum aequans. Capsula mem- 
branacea sphaerica. 


sunset ens ihc ttese. cess 


etn Mi 


Igo1 | UNDESCRIBED PLANTS FROM CENTRAL AMERICA 119 


Folia 4-9" longa 2-3.5°" lata plerumque inaequilatera supra densius 
pubescentia subtus pallidiora et purpureo-reticulata, dentibus magnis crebris- 
que. Pedicellicomplures 2—3.5°™ longi inferne marginati. Calycissegmenta 
lineari-lanceolata 2™™ longa dense pubescentia exreticulata. Corolla alba 
parce pubescens diametro circa 1°, faucibus 3™” latis, segmentis parum 
aequalibus ovalibus 3-4™" longis, Stamina glabra, filamentis 1™™ longis 
inferne marginatis, antheris reniformibus supra medium affixis reversis 0.5™" 
latis ante anthesin leviter cohaerentibus, loculis poro laterali subrotundo 
dehiscentibus. Ovarium pubescens, stylo 3.5™™ longo, stigmate parum bifido. 
Capsula diametro 1.5™™ calyce immutato inclusa. 

“Tn rupibus praeruptis humidissimis, Cubilquitz, Depart. Alta Verapaz, 
Guatemala, alt. 350™, Jun. 1900, von Tuerckheim, n. 7647 Pl. Guat. &c., qu. 
ed. Donn. Sm 


Amphilophium oxylophium Donn. Sm.—Pilis singulis dense 
molliterque ochraceo-tomentosum. Stipulae foliaceae. Folia 
conjugata cirrho carentia, foliolis breviter petiolulatis ovato-cor- 
diformibus contracto-acuminatis infra indumentum supra tuber- 
culatis subtus lepidotis. Calycis limbus tubum aequans lobos 2 
interiores plus quam duplo superans, lobis exterioribus triangulari- 
bus induplicatis reflexis. Corolla glabra ultra mediam labiata, 
labio inferiore fisso. Ovarium lepidotum glabrum. 

Caulis cum petiolis thyrsoque robustus et floccoso-tomentosus. Stipulae 
1o-12™™ longae deciduae. Petioli 2-2.5°™ petioluli 8-12™ metientes. Foliola 
7-12™ longa 5—75°™ lata supra pilis simplicibus adpressis vestita subtus pilis 
ramosis densius tomentosa et pallidiora. Thyrsus contracto-cylindricus in 
Speciminibus suppetentibus vix evolutus 1°™ !ongus, bracteis bracteolisque 
lineari-lanceolatis 1-1. 5°™ longis. Calycis totius lepidoti tubus subglobosus 
8™™ longus dense pilosus, limbus utrinque parce pilosus, lobis 5 exterioribus 
5-6™" longis, interioribus lati-rotundatis 3™™ longis extus parce pilosis. 
Corolla 2.5°™ longa ad apicem versus minutissime lepidota intus prope inser- 
tionem staminum linea lepidota incrassata, tubo 1° longo, labii inferioris 
laciniis 13™™ longis. Stamina majora 13™" minora to™™ metientia, thecis 
2.5™™ longis, staminodio 3™ longo. Discus pulvinaris cum stylo 1.5 longo 
glaber, stigmatibus semiorbicularibus, Capsula ignota. 

ucurrique, Costa Rica, alt. 630™, Jan. 1899, Zonduz, n, 13045 herb. nat. 

ost. 


Lophostachys Guatemalensis Donn. Sm.—Folia nascentia 
incano-pubescentia aetate provectiore praeter petiolum nervosque 


120 , BOTANICAL GAZETTE | FEBRUARY 


glabrescentia disparia lanceolato-elliptica longe falcato-acuminata 
triente inferiore in petiolum arcuatim attenuata subtus pallidiora. 
Spicae terminales subsessiles  solitariae, bracteis lanceolato- 
linearibus 5-nerviis, bracteolis spathulato-linearibus abrupte 
cuspidatis 2-nerviis. Calycis segmenta exteriora aequalia spathu- 
lato-oblonga cuspidato-acuminata, postico basi 7-nervio medio 
5-nervio, antici usque ad medium fissi laciniis 3-nerviis. Stamina 
omnia perfecta. 
Fruticulus decumbens dichotomo-ramosus, ramulis quadrangularibus. 
_ Folia 5-8™ longa 1.5-3.5°" lata supra saepe minutissime lineolata subtus ad 
costam nervosque pubescentia, petiolis pubescentibus 1-1.5™ longis, axillis 
saepius folia duo minuta producentibus, Spicae foliis reductis suffultae 
3-4°™ longae, bracteis g-10™™ longis 2.5-3™™ latis triente superiore falcato- 
acuminatis et ciliatis ad nervos pubescentibus, bracteolis 9™™ longis inaequi- 
lateralibus ciliatis, altera angustiore. Calyx 1.5°™ longus sicut bracteolae 
erubescens et pulchre reticulatus ciliatus ad basin versus utrinque incano- 


longi segmentis ovalibus 6™” longis. Stamina medio tubo affixa, antica 17" 
longa, posticorum 13™™ longorum antheris unilocularibus polleniferis, Discus 
i™™ altus, ovario oblongo-ovoideo 4™™ longo, stylo capillari 2.7° longo, ovulis 
ovalibus 1™ longis. Capsula non adest.— Ceteris speciebus adhuc descriptis, 
omnibus Brasiliensibus aut Peruvianis, haec bracteis bracteolis calycibus 
praecipue discrepat. 
/ Casillas, Depart. Santa Rosa, Guatemala, alt. 1300", Jan. 1893, Heyde et 
Lux, n. 4382 Pl. Guat. &c., qu. ed. Donn. Sm 


Hernandia didymantha Donn. Sm.— Ramuli cum foliis peti- 
olisque glaberrimi. Folia oblongo-ovata a medio linea arcuata 
sursum angustata basi obtusa aut rotundata penninervia. Pani- 
culae folia superantes praeter involucella totae incanae, ramis 
secundariis bifloris, bracteis 4 oblongis. Flos masculinus soli- 
tarius 3-merus, pedicello filiformi. Flos femininus 4-merus, 
involucello truncato pyramidali subclauso, fructifero sphaerico 
drupam subaequimagnam arcte includente, semine globoso. 

Arbor 15™ alta, trunco excelso, ramis inferioribus reflexis, superioribus 
erecto-patentibus, floribus porraceis (ex schedula et icone photographica Pit- 
terianis). Folia 12-18" longa 4.5-7™ lata tenuiter coriacea teralibus 


1901 | UNDESCRIBED PLANTS FROM CENTRAL AMERICA r21 


utrinque 7-8, petiolis glabris canaliculatis 3-6°™ longis. Paniculae circa 
5 terminales et ex axillis foliorum superiorum decrescentium prodeuntes cum 
: pedunculo ae iy 3 he psy el ae longae corymbiformes densiflorae, 
, inferioribus 2-3 longis. Bracteae 
jam ante anthesin reflexae 8™™ longae 3.5™™ latae apice rotundatae. Floris 
masculini pedicellus filiformis 7™™ longus, altero anantho brevi rarissime 
exstante, segmenta 6 oblongo-elliptica 5™™ longa obtusa, glandulae 6 liberae 
subsessiles, antherae oblongae 1.5™" longae filamenta aequantes. Floris 
feminini pedicellus 1™ longus, involucellum coriaceum parce pubescens 
siccitate nigrescens 3™™ altum atque latum, ore integro 1™ lato, segmenta 8 
uti glandulae 4 masculinis similia, stylus sursum incrassatus et quadrangularis 
4™" longus, stigma obliquum subintegrum nudum. Involucellum fructiferum 
crasse coriaceum 2.2°"-diametrale, drupa 1.9-diametralis ecostata (ut vide- 


V Hf. Guianensi Aubl. proxima recedit praesertim foliis penninerviis, flore 
masculino singulo graciliter pedicellato, involucello jam sub anthesi paene 
clauso, drupa minore. 

Ad litora Atlantica prope Punta Mona, Costa Rica, Sept. 1898, Pittier, 
n. 12682 herb. nat. Cost. 


Brosimum heteroclitum Donn. Sm.—Scandens. Folia mem- 
branacea glabrescentia oblongo-elliptica cuspidato-acuminata 
basi obtusa aut acutiuscula crenulato-serrulata, stipulis bracteisque 
a basi lata filiformibus. Receptacula in axillis duabus supremis 
approximatis solitaria sessilia maxima, masculinorum loculis I-3, 
ovulis abortivis in quoque loculo pluribus. 


Suffrutex in arborum truncis scandens et radicans, ramis virgatis 30-40™ 
longis glabris. Folia nascentia pubescentia, vetustiora supra punctis minutis” 
albidis scabriuscula costa nervisque pubescentia 10-16™ longa 4.5-6.5™ lata, 
nervis lateralibus utrinque 6-8, venis transversis subparallelis, petiolis 1-3 
longis sicut stipulae bracteaeque 6™ longae pubescentibus. Receptacula 
masculina globosa 1—-1.3-diametralia demum ovalia et usque ad 2™ longa 
Crasse Corticata ore bracteolis crassis ovatis munita, ovulis abortivis lateraliter 
affixis. Cetera desunt. Species habitu scandente et ovulis rudimentariis 
compluribus valde anomala. Receptacula vetustiora staminibus derasis tan- 
tum ee petin nt. 

“Jiménez, Llanuras de Santa Clara, Costa Rica, alt. 250", meet: 1896, 
Donn. Sm., n. 5117, Pl. Guat., &c., qu. ed. Donn. Sm, 


Pilea ptericlada Donn. Sm.— Dioica. Glabra. Folia oblongo- 
elliptica utrinque acuminata in petiolum brevem cuneato-attenuata 


122 BOTANICAL GAZETTE [FEBRUARY 


supra trientem inferiorem grosse crenato-serrata, nervis basilari- 
bus trientem superiorem attingentibus et costa subtus explanatis 
fuscis, stipulis quam petiolis paulo brevioribus. Pedunculi femi- 
nini foliis breviores et cymarum axes alati. 

Herbacea. Caulis e basi lignosa radicante assurgens in exemplis obviis 
6-15 altus, nodis approximatis. Cujusve paris folia aequalia aequilatera 
triplinervia 5-11°" longa 3-5°™ lata summa apice obtusiuscula supra cysto- 
lithis fusiformibus et punctiformibus farcta subtus plerumque nuda, foliis 
inferioribus minoribus, serraturis obtusis, costa nervisque supra immersis, 
stipulis orbiculari-ovatis 5-8™™ longis binerviis, petiolis 6-10™™ longis. Pedun- 


ramulisque patulis circa 1™ latis sicut pedunculi cystolithis obsitis. Peri- 
anthium sessile fructifer bracteolam cystolithis lineatam subaequans, seg- 
mentis parum inaequilongis, achenia ventricoso-ovata 2™" longa perianthium 
dimidio superante rubro-punctulata et marginata. Planta masculina ignota. 
Secundum methodum Weddellianam juxta P. elegantem Wedd. locari debetur. 

“In sylvis udis Atirrensibus, Prov. Cartago, Costa Rica, alt. 600", Apr. 
1896, Donn. Sm. n. 6779 Pl. Guat., &c., qu. ed. Donn. Sm.; Suerre, Llanuras 
de Santa Clara, Costa Rica, alt. 300", Apr. 1896, Donn. Sm. n. 6780 Pl, Guat., 


etc., qu. ed. Donn. Sm.—Eadem planta, ut videtur, sub nn. 172 et 199 Endres 


in herb. Kew. exstat. 


Costus sanguineus Donn. Sm.— Pallide ferrugineo-villosus. 
Folia discoloria supra bulboso-pilosa oblanceolato-oblonga acut- 
issime acuminata basi minute rotundata subsessilia. Strobilus 
lanceolato-ovoideus, bracteis pilosis oblongo-ovatis superne infra 
apicem carinatis, infimis apice foliaceis. Calyx prima tertia 
parte acute lobatus. Corolla sanguinea, labello pubescente 
oblongo apice angustata truncata 5-lobulato, ceteris segmentis 
anguste lanceolato-oblongis, connectivo triangulari labellum 
paulo superante. Ovarium pilosum. 

Ochreae laciniis scariosis usque ad 2°™ longis marginatae saepius reticu- 
lato-striatae cum vagina extus villosae et intus glanduloso-punctulatae. Folia 
subtus sericea et ad costam longe villosa 14-22™ longa supra medium 4.5— 
6.5°™ lata basi 8-1o™™ lata, nervis omnibus a basi prodeuntibus. Strobilus 
7-10™ longus circa 3° crassus, bracteis coriaceis sanguineis circa 4™ longis 
20-23" latis, bracteola 1 pubescente lineari-oblonga 18™™ longa 6™” lata 
acuminata, floribus sub quaque bractea singulis. Calyx campanulatus 8-9" 
altus pilosus striatus. Corolla purpureo-coccinea (von Tuerckheim in sched.) 
5.56 longa, labello 3.3°" longo 1.5°™ lato a triente superiore deorsum paulo 


it Min 


eee, meta 


Igor] UNDESCRIBED PLANTS FROM CENTRAL AMERICA 123 


angustato, ——* oblongis 3™™ longis rotundatis, ceteris segmentis 26—28™™ 
ongis 7~9™™ latis, filamento g-1o™™ lato, connectivo 7™™ longo in caudiculam 
marginibus revolutam producto, anthera 8™™ longa, loculis 1™ inter se distan- 
ibus. Ovarium breviter oblongum 5™™ longum, stigmate semilunari 3™™ lato 
ciliolato, appendice dorsali ovali apice retusa. Capsula ignota.— C. sficato 
w. proximus differt praecipue indumento et florum indole atque colore. 
vCubilquitz, Depart. Alta Verapaz, Guatemala, alt. 350™, ;Maj. 1900, von 
Tuerckheim, n. 7686 Pl. Guat. &c., qu. ed. Donn. Sm 


IsCHNOSIPHON Mor.ak (Eggers, Bot. Centralbl. 53: 307. p/. 
2) leiostachya Donn. Sm. —Spicae glabrae, bracteis magis elon- 
gatis.— Ceteroquin exempla Centrali-Americana cum specimine 
typico Ecuadoriensi in herb, Kew. asservato prorsus congruunt. 

San Pedro Sula, Depart. Santa Barbara, Honduras, alt. 300”, Sept. 1888, 
C. Thieme, n. 5519 Pl. Guat. &c., qu. ed. Donn. Sm.; Las Vueltas, Tucur- 
rique, Prov. Cartago, Costa Rica, alt., 635", Dec. 1898, Zomduz, n. 12 
herb. nat. Cost. 


Calathea dasycarpa Donn. Sm. (§ EucaLarHea Koern.)—Folia 
glabra oblonga 3-4-plo longiora quam latiora in cuspidem graci- 
lem acuminata basi rotundata in articulum cuneato-producta, 
folio inflorescentiam suffulciente caulinis simili longe petiolato. 
Pedunculus petiolo brevior bifidus. Spicae pro ratione parvae 
oblongae interdum basi furcatae, bracteis distichis vix ac ne vix 
imbricatis dimidiato-ovatis cymbiformibus coriaceis ad margines 


barbatis, floribus paucis binis. Corollae tubus sepalis 3-plo 


longior intus villosissimus. Ovarium pilosum. Fructus mono- 
spermus, 

Folia 60-80" longa prope medium 18-19 lata ad costam subtus pubes- 
centia, cuspide lineari 1.5—2.5° longa, petiolis cum articulo glabro 5-6" 
longa adjecto 43™ longis pilosiusculis. _Pedunculus 21% longus basi bractea 
1o™ longa apice bractea 3.5™ longa suffultus, ramis 3~-5°™ longis. Spicae 
6-9" longae, rachis internodiis inferoribus 1o-18™" longis, bracteis 10-12 
demum late patulis circa 2™ longis 9™™ latis, bracteolis 2 late oblongis Io - 
14™™" longis, exteriore bicarinata et psoe ta rotundatis 3-lobulata, interiore 
minore unicostata integra, floribus plerumque 4 minute pedicellatis. Sepala 
lanceolata 7-9™ longa. Corollae eee luteae (Pittier in sched.) tubus 
25™™ longus intus valde pluri-costatus. Ovarium obovatum 3™” longum dense 
longeque adpresso-pilosum, ovulo abortu unico triquetro, Fructus ovalis 
tom longus 5™™ latus pilosus, pericarpio membranaceo nervoso, semine 


° 


124 BOTANICAL GAZETTE [ FEBRUARY 


atro-purpureo oblongo 4.5™™ longo corrugato. Corolla cum androecio mar- 
cida tantum suppetit. 

“In sylvis ad oras Rio Hondo prope Madre de Dios, Comarca de Limon, 
Costa Rica, alt. 200", Nov. 1896, Péttier, n. 10350 herb. nat. Cost.; Agua 
Buena, Cafias Gordas, Comarca de Limon, Costa Rica, alt. 1100", Feb. 1897, 
Pittier, n. 11136 herb. nat. Cost. 


Calathea lasiostachya Donn. Sm. (§ EucatarHea Koern.) — 
Folia glabra elongato-ovata apice acuminata inferne rotundata 
ima basi cuneata, folio inflorescentiam suffulciente conformi 
dimidio minore petiolis usque ad articulum vaginatis bis longiore. 
Pedunculi 3-ni petiolum subaequantes. Spicae elongato-oblongae 
sordide villosae, bracteis stricte distichis late patentibus con- 
duplicatis membranaceis. Ovarium glabrum. Fructus dis- 
permus. 

Caulis robustus elatus. Articuli cum vaginis pedunculisque pilosiusculi. 
Folia 86° longa 23 lata a triente inferiore sursum angustata, articulis 9°” 
longis, petiolis longissimis, vaginis scabriusculis. Folium sub inflorescentia 
4o*™ longum 15 latum, petiolo cum articulo 3°" longo adjecto 21™ longo. 
Pedunculi 15~22°™ longi ad apicem versus villosi. Spicae 11.5-17°™ longae 
5.5-6™ latae, lateribus subparallelis. Bracteae 22-32 sublaxe imbricatae 
dimidiato-ellipticae subfalcatae 3.5°™ longae 1.5°™ latae intus sericeae, omnes 
conformes, bracteolis lineari lanceolatis et linearibus circa 2™ longis. Ovarium 
nitido-atrum oblongum 6™™ longum 4™™ latum 3-loculare abortu 2-ovulatum. 
Fructus laevis pallidus ellipticus 12™" longus 6™™ crassus, pericarpio per- 
gameneo intus valde reticulato, valvis 3 inaequilatis, minoribus latere septatis, 
seminibus atris 8™™ longis arillosis. Perianthium deficit. 

“In sylvis ad oras Rio Hondo prope Madre de Dios, Costa Rica, alt. 200", 
Nov. 1896, P2ttier, n. 10344 herb. nat. Cost. 


Calathea Verapax Donn. Sm. (§ MonosticHE Koern. )—Glabra. 
Acaulis. Folia inaequilatera elliptica vel ovato-elliptica acumi- 
nata ima basi in articulum brevem producta petiolis subaequi- 
longa. Scapus nudus petiolum superans usque ad medium 
bractea radicali inclusus basi petiolo et bracteis cinctus. Spica 
obovato-elliptica, bracteis membranaceis coloratis numerosis- 
simis lanceolatis filiformi-attenuatis subaequilongis. Corollae 
tubus sepalis linearibus dimidio longior, lobi lanceolati. 

Rhizoma bracteas 7-20™ longas apice denticulata petiolos 2 scapumque 
involventes emittens, petiolo altero toto’ fere vaginato alterum nudum 


a 


ee ae 


1901] UNDESCRIBED PLANTS FROM CENTRAL AMERICA 125 


includente. Folia 20-30°" longa 9.5-14™ lata, petiolis cum articulo 1-2 
longo adjecto 29-33 longis. Scapus 35-55°™ longus. Spica 8-12™ longa circa 
4°™ lata, rhachi 5°" longa, bracteis in sicco rubescentibus erecto-patentibus, 
infimis 6.5° longis 1.8°™ latis, supremis 6° longis 9™™ latis, bracteolis 3 late 
oblongis 1.7°" longis, exteriore valde bicarinata, floribus 2-4-nis. Sepala 
2.1°™ longa 2.5™™" lata. Corollae coccineae (ut videtur), tubus 3°" longus, lobi 
1.5% longi 5™™ lati acuti. Androecii labellum 1° longum 7™™ latum, lobus 
cucullatus appendice filiformi 4™™ longa instructus, staminifer anthera 2.5™™ 
longa superatus. Stylus incurvus, stigmate valde inflexo. Ovarium glabrum 
oblongo-obovatum 3-loculare. Fructus non adest.—Ad C. Petersenii Eggers 
habitu accedens spica et florum fabrica recedit. 

“Rubelcruz, Depart. Alta Verapaz, Guatemala, alt. 1000", Maj. 1887, von 
Tuerckheim, n. 1269 Pl. Guat. &c., qu. ed. Donn. Sm.; Sacolal, Depart. 
Alta Verapaz, alt. 1000", Apr. 1889, Donn. Sm.,n. 1779, Pl. Guat. &c., qu. 
ed. Donn. Sm 


Callisia grandiflora Donn. Sm.— Folia a basi rotundata ses- 
sili elliptico-lanceolata. Cyma elongata cincinnalis, pedicellis 
compluribus, floribus 3-meris maximis. Stamina fertilia 3, fila- 
mentis sursum barbatis, loculis divergentibus connectivo amplis- 
simo brevioribus. Stamina sterilia 3 nuda. Capsula 3-valvis, 
loculis monospermis. 

Praeter ocreas ciliatas glabra. Folia 6-12 longa 1.5—2.5™ lata acutis- 
sima, Cyma paniculiformis 6-11™ longa, rhachi flexuosa, axibus primariis 
1.5-2.5°" longis, lateralibus 7—20"™ longis, bracteis inferioribus foliaceis 
1-5 longis, superioribus depauperatis sicut bracteolae minimae subspatha- 
ceis, pedicellis circa 5—g-fasciculatis 7-10™" longis. Sepala herbacea albo- 
punctata oblongo-elliptica 6.5™™ longa 2.5™™ lata sub anthesi reflexa. Petala 
ex schedula Tuerckheimiana alba in sicc. pallide flavescentia oblongo- 
elliptica g™™ longa 4™™ lata obtusa. Stamina fertilia 6-7"™" longa, pilis 
aureis longissimis densissimis, antheris cinnabarinis transversim ovalibus 
1.5" latis utrinque praesertim apice bifidis, loculis oblongis 0.5" longis 
subdivaricatis. Stamina imperfecta 2-3™" longa, loculis connectivo parvulo 
subdiscretis. Ovarium ovale 1™™ longum stylum aequans, stigmate capitel- 
: _ Lae = ai been eee longa abortu saepius bisperma, semini- 

oblongis 4™™ longis.—Species staminibus 
et ae et iieceatarite sbhonnale ad pieereraceiad sect. Descantariam 
accesen 

“in wee et frutectis prope Cubilquitz, Depart. Alta Verapaz, Guate- 
mala, alt. 350™, Mart. 1900, H. von Tuerckheim, n. 7684 PI. Guat. &c., 
qu. ed. Donn. Sm 


oe eee Mp. 


BRIECLR ARTICLES: 


NITRATES AS A SOURCE OF NITROGEN FOR SAPRO- 
PHYTIC FUNGI 


The readiness with which different classes of plants make use of 
nitrogenous inorganic material has been variously demonstrated. It 
is well understood that salts of nitric acid, and to a less extent ammo- 
nia, serve as nutrient material for the higher plants, and that free nitro- 
gen and salts of nitrous acid are worked upon by certain bacteria in 
such a way as to make them available as a nitrogen food supply. But 
Pfeffer* states that phanerogams and saprophytic fungi are unable to 
assimilate nitrites. 

With a view of testing this latter point certain fungi were grown in 
culture solutions containing ammonium chlorid, potassium nitrate, 
potassium nitrite, hexamethylenetetramine, and peptone. The cul- 
ture was prepared with 200% chemically pure water obtained by redis- 
tilling from a solution of potassium permanganate until free from 
every trace of ammonia, 10% cane sugar, 0.5 magnesium sulphate, 
o.1™ acid potassium phosphate, and a trace of ferric chlorid. To 10 
of the sterilized solution was added 0.1 of one of the nitrogen con- 
taining substances. One drop of this preparation placed on a cover 
glass was inoculated with the fungus and the cover glass inverted 
over the cell and placed in the thermostat for cultivation at 28° C. 
The fungi used for inoculation were Aspergillus flavus and Botrytis 
vulgaris. Throughout the experiment care was taken to have the 
solutions and apparatus thoroughly sterilized, and all of the salts used 
were chemically pure. 

Duplicates of each were prepared, also checks using the solution 
without the nitrogen containing compound. The cultures were exam- 
ined at intervals for from one to three days. At the expiration of the 
time unquestionable results were obtained. In potassium nitrate and 
potassium nitrite thé fungi grew with apparently equal vigor, Aspet- 
gillus developing well formed fruit bodies. Hexamethylenetetramine 
showed itself an excellent source of food, since both fungi grew an 

* Physiology of Plants, translated by Ewart, 406. 1899. 

126 [FEBRUARY 


1901 | BRIEFER ARTICLES 127 


fruited in it. In peptone the fungi grew rapidly and luxuriantly, 
proving it to be a favorable source of nitrogen.— Mary H. Smiru, 
Botanical Department, Cornell University. 


NON-SEXUAL PROPAGATION IN OPUNTIA. II. 


A VERY interesting Opuntia which has recently come to my notice 
in studying the various propagative methods of the Cactaceae is O. 
arbuscula Engelm., a small, more or less arborescent form, densely 
branched, and reaching a height of about 15. This plant sets an 
abundance of fruit which appears to mature well, but which upon 
examination is found to contain very few good seeds. So laden is the 
plant with its fruit that its branches, as a rule, bend over so as almost, 
if not quite, to touch the ground, In this position there takes place a 
process analogous to “layering,” new shoots of an apparently primitive 
character arising from the decumbent branches, which also give off 
roots into the soil. The same formation of primitive shoots occurs in 
joints detached from the parent plant. This is also true of fruits, from 
the sides of which both stems and roots may often be found forming, 
so often, in fact, that this must be regarded as the rule rather than the 
exception. We have here the case of a structure, modified primarily 
for sexual purposes, turned finally to use in a non-sexual way, to 
accomplish, broadly speaking, the same end. 

Still another method of propagation, perhaps not very common, 
yet apparently not infrequent with this species, is by the formation of 
adventitious shoots on the roots. The roots are, in this form as in the 
majority of the Cactaceae, divided into two systems, as already described.’ 
On the absorptive roots, which run just below the surface, there arise, 
at some distance from the main plant, adventitious shoots of a char- 
acter far more primitive than those formed on fruits or fallen joints. 
The leaves of these shoots are in some cases over 10" long, green, and 
succulent. By the time these shoots reach the height of about 2, the 
root connecting them with the parent plant dies, thus leaving them 
independent at an early stage. 

The distribution of this species is very well defined. It occurs 
almost exclusively in those slight depressions in the plain, which in 
time of hard rains are washed by broad and shallow streams of surface 


* Bor. Gaz. 30 : 348 seg. 1900. 


128 BOTANICAL GAZETTE [FEBRUARY 


water. Here it grows in colonies, following the lines of the depres- 
sions. The soil in such places is much finer and less pebbly than that 
on the slight elevations near by, but experiments in transplanting have 
demonstrated that this distribution is not due to soil characters. It 
seems probable that the fruits and joints are washed down by the 
stream, and settle at various places along the course. From the 
single plants so started colonies soon are formed, through the agency 
in part, it may be supposed, of root propagation and “layering.” — 
CaRLETON E. Preston, Harvard University. 


4 
i] 


— a “errr 


CURREN FF LIFERAAAIRS, 
BOOK REVIEWS. 
A new school botany.’ 


AT least three things must be considered in making an estimate of a 
text-book, namely, its style, its reliability, and its pedagogical standpoint. 
Professor Bailey’s style is too well known to need description or commenda- 
tion. He is one of our clearest and most forceful writers. The general 
facts of botany are fairly well established and are common property, so that 
the preparation of an elementary text involves merely selection from a great 
mass of well-known material. In the book before us, therefore, there is no 
occasion to discuss style or reliability, although in the latter feature Profes- 
sor Bailey is as great a sinner as the rest of us. In regard to the pedagogi- 
cal standpoint, however, he has raised a distinct issue and this deserves 
statement and some discussion. The author has had an extensive experience 
with teachers and schools, and his verdict is that “the schools and the teach- 
ers are not ready for the text-book which presents the subject from the 
view-point of botanical science.” To discover the explanation of this state- 
ment by means of his book, it becomes evident that the author does not 
believe in the organization of botanical material so that some conception of 
the science as a whole may be developed. From his point of view the study 
need not develop the idea of relationships, or need not be used to illustrate 
principles. The selection of material is to be made from forms and phe- 
nomena which are familiar, and which are related to the experiences of the 
daily life. All of this means that in the judgment of the author the average 
recent botanical text is not adapted to the majority of teachers and of schools 
as they are, but overshoots them. 

At least two objections to this view have doubtless had weight with those 
botanists who have prepared texts from a different standpoint, namely, the 
Conviction that pupils of secondary-school age are ready for some organi- 
zation of a science, and the further conviction that teaching can only be 
improved when some pressure is brought to bear upon teachers to become 
properly trained. It should be said that the author distinctly disclaims any 
Criticism of existing text-books, but recognizes the need of one adapted to 
actual rather than to ideal conditions. 

An illustration of the result of not keeping hold of some little thread 
of — may be found in Professor Bailey’s chapter XXv, entitl 


* BAILEY, L. H.: an ceremings 2 text for schools. 8vo. pp. xiv-+356. 


New York : pei leant aces 1900. 


t90r] 129 


- 


130 BOTANICAL GAZETTE [FEBRUARY 


‘Studies in cryptogams.”’ In our judgment this chapter will be unintelligible 
or misleading to those who have had -no other preparation for it than that 
obtained from the preceding chapters. This is no criticism of the chapter 
as to its contents, but as to its pedagogical soundness. 

Another position taken by the author deserves attention. He says: 
‘There are other ideals than those of mere accuracy. In other words, it is 
more important that the teacher be a good teacher than a good botanist. 
One may be so exact that his words mean nothing.”’ The writer sympathizes 
with the thought in Professor Bailey’s mind, for he has encountered these 
oppressively accurate and insistent teachers, whose mania for precision kills 
inspiration; but he doubts whether teachers in the secondary schools need 
any encouragement to be inaccurate. It would seem evident that reasonable 
accuracy, as opposed to pedantic accuracy, and inspiration should coexist in 
the teacher. If there is anything for which science stands, and in which it 
should train even the very young, it is a reasonable accuracy. 

The four general subjects presented in the book are the nature of the 
plant itself ; the relation of the plant to its surroundings ; histological studies ; 
and determination of the kinds of plants. From the pedagogical standpoint 
the author regards the third as the least important. The book is full of sug- 
gestive material for the teacher, and the illustrations are very recite 

he experiment suggested by Professor Bailey is worth the trial, and n 
teacher should be so confident of his own methods as not to await the out- 
come with keen interest.— J. M. C 


The Umbelliferae. 

JUST TWELVE YEARS after the appearance of their first Revésion of 
North American Umbelliferae, Drs. Coulter and Rose have brought out a new 
monograph of the same group, which appears to be a model of what such 
work should be.? 

From the time of Linnaeus to the treatment of our Umbelliferae by Tor- 
rey and Gray, eighty-seven years, thirty authors wrote on the group, pro- 
ducing forty-one books and papers, which contained 195 new species Or 
names. In the next forty-seven years, up to the appearance of the Revision 
of Coulter and Rose, twenty-seven writers, in fifty-seven contributions, intro- 
duced 258 new names or species in the group. And in the last twelve years 
nineteen persons, contributing forty-three papers, have added 108 new species 
and names. e Monograph now issued (as it chances, on the last day of the 
century) describes and places 332 native species and 39 which are considered 
as introduced, or a total of 371, in contrast with 233 included in their earlier 
Revision. No comment is needed on this as an indication of the rapidity 
with which the understood components of our flora are changing. 

OULTER, JOHN M., and Rose, J. N.: Monograph of the North American 
Ar oobi Contr. U. S. National Herbarium 7: 1-256. pls. 7-9. figs. 1-05. 19000 


Sy oe 


a 


1901 } CURRENT LITERATURE 131 


In addition to what appear to be serviceable keys to the genera and 
species, that which is now believed to be their necessary bibliography and 
Synonomy, adequate descriptions, and a full citation of material examined, 
the work contains a large amount of tabular and statistical matter, which if 
not of interest to the ordinary seeker after the name of a plant, at least 
shows the painstaking care that has been bestowed on the study. A 
interesting feature is a table of data concerning the specimens which have 
served for illustration, and as all of the sixty-two native and sixteen intro- 
duced genera are figured, both as to the appearance of their fruit and its 
Cross section, this information is of no little importance for those who in 
future may have to familiarize themselves with what these genera now 
Stand for. 

That all but four of some 800 references have been verified not only 
Shows the industry of the authors but ensures the trustworthiness of their 
Statements in this, a feature which is too often lamentably misleading to the 
men who compile from unverified citations. 

The most conspicuous changes are as follows: Coloptera C. & R. is found 
to represent true Cymopterus, and the Cymopterus aggregate of the former 
Revision is distributed under four genera, two of them new (Aulosfermum and 
Rhysopterus), and two of them Nuttallian (Phellopterus and Pteryxia) ; Peu- 
cedanum L. is found to have no indigenous species in North America, and 
this greatest of our umbelliferous genera becomes known as Lomatium Raf., 
certain groups of species heretofore included being recognized as genera, as 
Cynomarathrum Nutt. and Euryptera Nutt.; Centel/a L. is recognized as dis- 
tinct from Hydrocotyle; Deweya T. & G. is restricted to its type species, and 
a new genus, Drudeophytum, established to include the other species variously 
described under Deweya, Velaea, and Arracacia; and Sphenosciadium Gray is 
taken out of Sedinum. As a result of these and other changes, the following 
§eneric names disappear from our flora: Coloptera C. & R.. Crantzia Nutt., 
Cryptotaenia DC., Discopleura DC., Leptocaulis Nutt., Osmorhiza Rat., 
Peucedanum L., Phellopterus Benth., Selinum L., Tiedemannia DC., an 
Velaea DC. 

All-in-all, though changes are not unlikely to occur that may relatively 
Soon cause us to look upon this as merely work of the last century, it appears 
to be of such a character that the twenty-first century will still see it at the 
elbow of every advanced and attentive student of the Umbelliferae of our 
country.— WILLIAM TRELEASE. 

MINOR NOTICES. 

THE THIRD FASCICLE of the second volume of Pittier's Flora of Costa 
Rica3 has appeared. The first fascicle contained the Polypetalae (excepting 

3PITTIER, H.—Primitiae Florae Costaricensis. Vol. Il, pp. 219-294- Piper- 
ace, by Casimir de Condelle. San José de Costa Rica. 1899. $1. 


132 BOTANICAL GAZETTE [FEBRUARY 


Polygalaceae, Sapindaceae, Meliaceae, Leguminosae, and Melastomaceae), by 
John Donnell Smith. The second fascicle included the Gamepetalae (except- 
ing Rubiaceae, Compositae, Solanaceae, Convolvulaceae, Acanthaceae, and 
Labiatae), by the same author. The present fascicle presents the Piperacee, 
by Casimir de Candolle, who calls attention to the close affinity of the flora 
to that of South America. The two genera are Piper and Peperomia; the 
former containing eighty-three species, fifty-one of which are described as 
new ; the latter forty-three species, twenty-one of which are new.—J. M 


FESSOR Kari” SCHUMAN has begun the publication of Blihende 
ppc (Lconographia Cactacearum), with the assistance of the Deutsches 
een-Gesellschaft. Each part is to contain four colored plates, with 
ee text, and is sold for four marks. It is expected that about three 
parts will be issued each year. The first part contains Echinocactus micro- 
spermus Web., Echinopsis cinnabarina Lab., Echinocereus subinermis Salm- 


. Dyck, and Echinocactus Anisitsii K. Sch., the last being a new species from 


Paraguay. The plates are beautifully colored illustrations of the plants in 
bloom, made from nature by Frau Dr. T. Giirke. The text, by Professor Schu- 
man, needs no comment. The publisher is J. Neumann, Neudamm, Bran- 
denburg, Germany.—J. M. C 


Dr. A. J. Grout’s Mosses with a hand lens‘ describes in nontechnical 
language 100 of the mosses of the northeastern United States which can be 
recognized with some degree of certainty by the use of a simple lens. The 
identification in many cases must be limited to the genus, the specific differ- 
ences being too recondite for observation in this way. Miss Thayer’s excel- 
lent drawings will be quite as helpful to the student as Dr. Grout’s text. 
key based upon the more apparent structural characters, and one based upon 
habitat, would need to be tested before pronouncing judgment upon them, 
but they are here and there unavoidably somewhat vague, which always 
detracts from the value of a key. The only danger from the use of such a 
book will be that beginners will not heed sufficiently the author’s cautions, 
and will be too sure of their determinations. But if properly used the book 
will stimulate interest in the mosses and lead on to more exact study. 

The glossary has concise and accurate definitions, elucidated by admira- 
ble illustrations. It would have been more serviceable had it been arranged 
in one alphabetic sequence instead of being divided into several. One must 
first know to what his strange word is applicable before he can 1 tell in what 
group to look for it— C. R. B. 


‘Grout, A. J.: Mosses with a hand lens. A nontechnical handbook of the more 
common and more easily recognized mosses of the northeastern United States. Illus- 
treated by Mary V. Thayer. 8vo. de xii+74. pls. 8 figs. go. New Yo rk: Th 
Author, 360 Lenox Road, Flatbush. 


. 


Sy 


1901 | CURRENT LITERATURE 133 


NOTES FOR STUDENTS. 

IN TWO PAPERSS Dr. Bessey states that some recent observations of his 
have lead him to think that the greater portion of the state of Nebraska is 
capable of supporting a tree vegetation. He claims that the absence of trees 
is due to the prairie fires, and that now wherever given a chance the tree area 
of the state is spreading. — H. N. WHITFORD. 

ACCORDING TO MIYAKE ® the majority of the leaves of Japan evergreens 
are found to contain more or less starch in winter, only one third of the 
species being devoid of it altogether. The starch content in winter, how- 
ever, is considerably decreased; this decrease begins in November, the 
minimum of starch is found in January, and the amount increases again in 
February. The author also shows that starch is actually manufactured in 
winter, though much less than in summer.—H. C. CowLEs. 

NEMEC asserts’ that in certain plant parts, ¢. g., the root, where transmis- 
sion of a stimulus occurs, there are embedded in a special plasma fibrils, the _ 
clusters of which, by proper staining, may be made easily visible with mode- 
rate magnification. These fascicles correspond on opposite sides of the 
partition walls and presumably are continuous or at least in contact through 
it. Experiment has shown that conduction of a stimulus is more rapid in the 
direction of these fibrils than across them, and tha: after their degeneration 


‘this difference in rate disappears. The perceptive region of the root, he 


declares, lies mostly in a special group of cells in the root cap. These are 
characterized by a very fluid plasma, and permanent starch grains which 
easily sink through it to rest on the ectoplasm. “Fibrils extend from these 
cells to the region of curvature. In some roots this group of cells becomes a 
special organ, which may be compared in principle with the organs of equilib- 
rium (vesicles with statoliths) in certain lower animals. (Cf. Noll, rev. in 
Bor. Gaz. 30: 134. Igoo0.)—C. R. B. 

Mr. JAmes A. TERRAS, of Edinburgh, has examined the conditions 
under which the winter buds of Aydrocharis Morsus-rane germinate. This 
plant propagates itself by buds, formed at the extremities of the subaqueous 
runners, in which is stored an abundant supply of reserve proteid, apparently 
a fluid albumin. The autumnal buds separate as soon as mature and sink to 
the bottom, where they rest till the following spring, or longer if covered by 
mud to the depth of two or three centimeters. It seems that this cover is 
effective merely because it cuts off the light, any other opaque screen like- 
wise preventing germination. Indeed if merely much shaded the buds do 

*The forest and forest trees of Nebraska. Reprint from Report of Nebraska 


Board of Agriculture, pp. 79-102. 1899. 
€ natural spreading of timber areas. Forester 6: 240-243. 1900. 


* Bot. Mag. 14: 44-49. 1900. 7 Biologisches Centralblatt 20: 369. 1900. 


134 BOTANICAL GAZETTE [FEBRUARY 


not develop. Experiments with colored liquid screens show that yellow and 
orange rays are most efficient, as Heald found in studying the germination of 
moss and fern spores. Heat apart from light is incapable of inciting to 
development ; and no cou of food, either carbohydrate or nitrogenous, 
awakens to activity. No zymogen was found in resting buds, and Terras 


activity, leading to the development of a zymogen, which in its turn is con- 
verted under the influence of the light into a zymase by which the utilization 
of the stored food becomes possible.—-C. R. B. 

IN HER RECENT WORK on Lavatera,® Miss Byxbee describes a process of 
spindle development as follows: In the young pollen mother cells the cyto- 
plasm consists of a fibrous network and a granular substance. As division 
approaches, the network surrounding the nucleus pulls out parallel to the 
membrane, forming a felt of fibers, and at the same time the granular sub- 
stance of the cytoplasm collects in a wide dense zone about the nucleus. 
* The nuclear wall now breaks down, and the fibers outside begin to grow into 
the nuclear cavity and mingle with the linin threads, which appear to have 
increased in quantity. This central mass of fibers now grows out into several 
projections, which become the cones of the multipolar spindle. Two of 
these cones become more prominent than the others, which they finally 
absorb, and the result is a bipolar spindle. Just how this absorption of the 
smaller cones is brought about is not made clear either in the description or 
in the figures. 

The work is well illustrated by four beautiful lithographic plates. While 
the results differ in certain minor details from previous work on the subject, 
it confirms the more important points that have already been worked out in 
such forms as Equisetum, Cobaea, Passiflora, Gladiolus, etc. The paper is 
an addition to the very interesting series of contributions on spindle forma- 
tion recently issued from the Botanical Laboratory of the University of 

rnia, 


ing’s strong solution, with an excess of acetic acid, palladium 


acid was added, were used for fixing; saffranin, gentian violet, and orange 
G were rie for staining. — A. A. Lawson. 

THE GEOLOGICAL SuRVEY of New Jersey has just issued a publication? 
which contains some valuable information concerning the forests of that 
state. In this C. C. Vermeule discusses the physical conditions of the forests 
of the state, and gives some field notes on forest conditions. Gifford Pinchot 

8 BYXBEE, EpiIrH SUMNER: The on of the karyokinetic spindle in the 
pollen mother cells of Lavatera. Cal. Acad. Sci. III. Bot. 2: 63-81. p/s. 10-73. 1900- 
® Annual Report of the State Geologist for was Report on forests. Geol. Surv- 

of New Jersey, pp. ix+-327, pls. 27, with maps. 


Igor | CURRENT LITERATURE 135 


writes on the effects of fire on forest production, and on the plains, and adds 
some silvicultural notes on the white cedar. The last two topics are incor- 
porated in the present volume from former reports of the survey. 

y far the most useful part of the report from an ecologic standpoint is 
a paper discussing the relation between forestry and geology, by Dr. Hollick. 
This is a revision of a paper of the same title published in the January and 
F ‘ebruary (1899) numbers of the American Naturalist. \t contains a discus- 


the tension zone lies between these. Dr. Hollick draws the conclusion that 
“the coniferous zone is destined to be ultimately obliterated, or only to exist 
over limited areas, often’ for the negative reason that in such areas the con- 
ditions may not be favorable for other ty pes of vegetation.” 

In part three of the report John B. Smith treats of the réle of insects in 
the forest; and part four contains an article by John Gifford on the forestal 
conditions and silvicultural prospects of the coastal plain of the state, with 
remarks in reference to other regions. Other valuable features of the report 
are a large number of half-tone reproductions from photographs, and maps 
Showing geological formations, distribution of rainfall, and distribution of 
forest areas.— H. N. WHITFORD. 


ITEMS OF TAXONOMIC INTEREST are as follows: C. H. BIssELL (Rhodora 
2: 225. 1900) has described a new variety of Zézia aurea from Connecti- 
cut.—Two new genera of Hymenogasters have been described recently : 
Arcangeliella, by F. CAVARA (Nouvo Giorn. Bot. Ital. 7: 117-128. p27, 
1900), from the coniferous forests of Vallombrosa, Etruria; and Martedlia. 
by O. MartrroLo (Malpighia 14:39-110. AZ. 7. 1900), from Sicily.—L. 
PETRI (Malpighia 14: 111-139. pls. 2-g. 1900) has described a new genus 
of Gasteromycetes from Borneo, C/athro aster by name.—H. T. A. Hus 
(Zoe 5: 61-70. 1900) has published a preliminary synopsis of the west coast 
Species of Porphyra, recognizing thirteen species and varieties, four of which 
are described as new.—S. B. PARISH (idem, 71-76) has begun a series of 
papers entitled “Contributions to Southern California Botany,” the first con- 
taining new species or varieties under Sphacralcea, Gilia, Galium, Eupatorium, 
and Sidens.—T. S. BRANDEGEE (idem, 78-79) has described a new species of 
Tapirira from Lower California.— Miss ALICE Eastwoop (idem, 80-90) 
has described new Californian species under Sa/ix, Chrysopsis, Helianthella, 
Sphacele, Mimulus, Aphyllon, Asclepias, Cleomella, Peucedanum, and Nav- 
arretia.—C. L. POLLARD (Proc. Biol. Soc. Washington 13: 184. 1900) has 
described a new Helianthus (H. agrestis) from Florida.— SCRIBNER and 
MERRILL (Division of Agrostology, Circular 27, Dec. 4, 1900) have described 


136 BOTANICAL GAZETTE [FEBRUARY 


two new species of Eafonia from the southern states. — RUDOLPH SCHLECHTER 
(Mém. de l’Herb. Boiss. 21: 1-78. 15 N. 1900) has published a monograph of 
the Podochilinae (a group of orchids including four genera), much enlarging 
the borders of Podochilus, recognizing forty-seven species in it, and describ- 
ing five as new; describing a new genus, Lodogyne; and recognizing six 
species in Th elasés, and two in Oxyanthera.—J. M 


Messrs. D. H. Scott and T. G. Hitt have published an account of 
the structure of Jsoetes Hystrix, a terrestrial species, dealing entirely with 
the vegetative organs. Some of the results are as follows: - 

Stem.— There is some evidence of a single apical cell; the stele is not 
composed of united leaf traces, but is cauline, as in the simpler lycopods; 
two cambiums are developed, in some cases the outer, in other cases the 
inner first; well-differentiated phloem is always present in the intracambial 
zone, being continuous with the leaf traces. 

Leaf.-— The vascular bundle in the lamina has exarch structure, the pro- 
toxylem lying next the phloem; the phloem contains true sieve tubes with 
transverse and lateral sieve plates, on both of which callus is formed ; growth 


is intercalary, except at first ; the labium and velum are derived from tissue - 


above the sporangium, and not from sterilized sporogenous tissue 

Root.— The stele has a monarch structure throughout, the differentiation 
of the xylem beginning with the development of a single tracheid ; the apex 
is distinctly layered, the initial groups giving rise to plerome, and to inner 
and outer cortex, 

In the conclusion of the paper the authors discuss the systematic position 
of the genus, presenting strong arguments in favor of its affinity with the Lyco- 
podiales, in which group it seems to have some real affinity with Selaginella, 
but not close enough to include the two in the same family. ‘“ The relation- 
ship of Isoetes to the Lepidodendree is probably a nearer one.” The 
authors regard the genus as one reduced from a much more complex type, 
and in no sense a primitive form of the leafy sporophyte. They would regard 
it ‘as a group that has long hovered on the limit of terrestrial and aquatic 
life, some of the forms becoming wholly submerged, while a few have definitely 
betaken themselves to dry land, a large proportion leading a more or less 
amphibious existence,” — J. M. C. 


ONE OF THE greatest ecological investigations of the day has been 
delayed if not permanently checked by the untimely death of. the brilliant 
young Scotch botanist, Robert Smith, of Dundee. Inspired largely by 
Flahault, he attempted to do for Scotland what the latter is doing for south- 
-ern France, viz., make a detailed ecological survey of the country. Since 
1896 Smith has worked unceasingly at his task, and had published but the 


7 Annals of Botany 14: 413-454. pls. 27, 2g. 1900. 


eS eee es en oer ee | fee eer eer were = i) Soh SSS ae eee 


1901] CURRENT LITERATURE 137 


first two installments when death put an end to his labors. In 1894 
Flahault conceived the idea of making an ecological map of France,” espe- 
cially with regard to the forests and agricultural areas. In 1897 the first 
sheet, corresponding in a way to the topographic sheets of our national geo- 
logical survey, was published.’ The maps are made on the scale of 1: 200,000, 
and each plant association is represented by a given color. Flahault has 
adopted twenty-two conventional color tones, which also in a general way 
show the topographic relief, lowlands having light and highlands dark 
colors. Contour lines are used as on ordinary topographic sheets. Smith 
studied with Flahault at Montpellier and then turned his enthusiastic atten- 
tion to his native country. 

In 1899 Robert Smith published an interesting paper on the study of 
plant associations,“ in which was given a historical summary of plant society 
studies from Humboldt down to Warming and Flahault, together with sug- 
gestions for use in field work. The two sheets which the author published 
give evidence of the most careful work, and cause us to regret that we shall 
not soon see any more. The Edinburgh sheet deals largely with lowlands 
and hence with cultivated areas. The littoral vegetation consists of marsh, 
dune, and rock plants, all of which the author regards as halophytic. The 
dominating forest trees are oaks on the plains and hills, pines and birches on 
the mountains, and alders in the swamps, though but little natural forest 
remains. In the higher areas are many dry, medium, and wet heath asso- 
ciations. The North Perthshire sheet has to do with a mountainous district. 

he maps are finely executed in colors that show strong contrasts. In 
America no comprehensive work like that of Smith or Flahault has yet been 
done, though the excellent mapping of our forest reserves under Gannett’s 
supervision, published in the nineteenth and twentieth annual reports of the 
director of the United States Geological survey, does a similar grade of work 
for the forests. Professor Geddes*® has given a very appreciative sketch of 
Robert Smith, which shows how his loss was felt at home.—H. C. CowLes. 

THE CURIOUS PARASITIC Balanophoracee have always excited interest, 
but especially so since the appearance of the papers of Treub (1898) and 
Lotsy (1899) describing the strange ovulate organ and apogamous embryo of 
species of Balanophora. Our knowledge of the group has now been extended 
by a paper just published by Dr. Lotsy,** in which he describes a species of 

**SMITH, ROBERT: Botanical Survey of Scotland. I. Edi vie District. I 
North Perthshire District. Scot. Geog. Mag. 16: 385-416, 441-467. I 

* Bull. Soc. Bot. France 41: 56-94. 1894. "3 Annales de agit 1897. 

4 Nat. Sci. 14: 109-120. 1899. Scot. Geog. Mag. 16: 597-599. 1900. 

* Lorsy, J. P.: Rhopalocnemis phailoides Jungh., a morphological-systematical 
study. Ann. Jard. Bot. Buitenzorg II. 2: 73-101. pls. 3-74. 1900 


138 BOTANICAL GAZETTE [FEBRUARY 


Rhopalocnemis. The plant body is tuber-like, frequently as large as a man’s 
head, is entirely devoid of even a trace of foliar organs, and is parasitic upon 
the roots of various trees. It seems to pass several years underground and 
comes to the surface but ashort time before the development of flowers. The 
thick spicate flower clusters burst through the outer layers of the tuber- 
ous body, the individual flowers being well protected by peculiar peltate 
scales. 

The carpellate flower consists of a syncarpous pistil, made up of two to 
five carpels, and inclosing a structure which Lotsy calls a free central 
placenta, and once, presumably by inadvertence, a nucellus. In any event, 
the structure is the enlarged tip of the axis of the flower, which soon com- 
pletely fills the cavity of the ovary. At this stage one or more hypodermal 
cells of this axis tip enlarge, and without division are transformed into 
embryo sacs, one of which germinates in the usual way and passes through 
the ordinary ante-fertilization stages. The oe bk cola this axial structure 


with its embryo sacs as a placenta without ovules; but, even aside from the 
fact that a placenta is nothing ria ar the reviewer sees no reason 
for regarding the structure other than a terminal cauline ovule without 


integuments. A very large primary fs SE pre is formed in the 
usual way, but the author never observed a pollen tube, nor could he by 
repeated artificial pollination induce pollen tubes to develop. Under these 
circumstances Balanophora has learned to develop an embryo apogamously 
from the micropylar polar nucleus, but Rhopalocnemis is unable to do so, 
and hence has become practically a seedless plant. In just one case was 
Lotsy able to secure a few seeds, and even in them few stages of developing 
embryos were discovered, but enough to assure him that they had come from 
the egg, and probably a fertilized egg. 

The staminate flowers are no less singular, each one consisting of a single 
structure which by courtesy may be called a stamen, but is probably a 
transformed axial structure. In its enlarged extremity numerous imbedded 
sporangia are developed, centrally as well as peripherally. These sporangia 
do not organize definite wall layers as in ordinary angiosperms, and have no 
method of dehiscence other than the breaking down of the superficial tis- 
sues. It would seem to be the rarest chance, therefore, if a pollen grain 
should ever reach a stigma, which in fact has usually lost all power of retain- 
ing pollen grains. The pollen grains are completely organized, and the two 
male cells appear, both of them finally assuming, along with the tube nucleus, 
an elongated, vermiform appearance, which according to Lotsy is merely 
preparatory to disorganization. 

he twelve elaborate and handsomely colored plates present every detail 
observed, as well as the condition of the preparations. It is unfortunate that 
Dr. Lotsy writes in English, as his unfamiliarity with the language makes his 


ae 


TT ARE meng nee 


1901] CURRENT LITERATURE 139 


meaning a matter of inference rather than statement. While the well- 
informed English reader finally comes at his meaning, the paper must be 
baffling to the foreigner who is compelled to translate——J. M 

PROFESSOR HARPER has given us a detailed description of the sexual 
reproduction of Pyronema.” The account is of especial interest, for it pre- 
sents another instance of the fusion of multinucleate gametes, and is an impor- 
tant corroboration of Stevens’ studies on A/bugo bliti with respect to the 
behavior of cytoplasm and nuclei under such conditions. 

The oogonium of Pyronema, as is well known, puts forth a conjugation 
tube (trichogyne) whose tip fuses with the antheridium. e tube is sepa- 
rated from the oogonium by a cross wall before its fusion, and this septum is 
later absorbed, when the protoplasm from the antheridium passes into oogo- 
nium and fertilizes the shee Mcheiiaie 

The oogonium and a Itinucleate f the start. The con- 
jugation tube likewise contains many nuclei, but these break down before 
fertilization. The number of nuclei in the oogonium is variable but there 
may be an many as two hundred. These gather in the central] region of the 
Structure, forming a closely packed collection at the time of fertilization. 
A great many sperm nuclei enter the oogonium through the conjugation tube, 
but they are hardly likely to equal the female nuclei in number. e sperm 
nuclei are attracted to the central mass of female nuclei, and shortly after- 
wards are found fusing in pairs with these elements. ome nuclei are 
always left over unmated, and these may be recognized for a long time by 
their smaller size. There is evidence that they finally break down. 

The ascogenous hyphae spring directly from the fertilized oogonium, and 
as they develop the oogonium becomes rapidly emptied of its protoplasm 
and is finally left as a hollow cyst. The development of the asci is not essen- 
tially different from Peziza, Ascobolus, Erysiphe, and other types. The 
young ascus is the second cell of a curved branch. It contains two nuclei 
that fuse, and these are not sister nuclei. The fusion nucleus gives rise by 
successive mitoses to eight nuclei accompanied by the beautiful asters that 
Harper has described for several other forms, and the ascospores are cut out 
of the cytoplasm by these asters in the characteristic manner. 

The ascocarp of Pyronema is a compound structure involving several sys- 
tems of ascogenous hyphae from as many fertilized oogonia. The elements 
become so mixed that it is impossible to separate them. However, the — 
§enous hyphae may be readily distinguished from the vegetative mycelium 
that forms the envelop of the ascocarp and the paraphyses in the hymenium. 
Harper points out that the characters most distinctive of the ascogenous 
hyphae are large nuclei, many times larger than those of the vegetative 
mycelium. 

‘7 HARPER, R. A.: Sexual reproduction in hatin confluens and the caashaiver 
of the cman Ann. of Bot. 14: 321. pls. 7 1900, 


140 BOTANICAL GAZETTE | FEBRUARY 


This study of Pyronema has important bearings in various directions and 
these are discussed at length. Along the lines of Harper’s previous studies 
they serve to clinch more strongly his argument, supporting the views of De 
Bary, that the Ascomycetes have sexual organs. Indeed the opinions of Van 
Tieghem, Brefeld, and Dangeard seem to have passed below the horizon of 
the present day outlook, so conclusive is the evidence presented from work in 
various fields, among the lichens, the Laboulbeniales, the Perisporiales, and 
the Pezizales. 

Pyronema is of especial interest because it presents characters somewhat 
intermediate between the simple fusion of the antheridium and oogoniun in 
Sphaerotheca, and the complex apparatus with the trichogyne found in the 
lichens and Laboulbeniales. Still it is very remarkable that such diverse 
conditions should appear in a single group, and the question seems very 
fair whether or not the Ascomycetes are a phylogenetic unit. However, 
the trend of investigation indicates complexities in life histories as well as 
structures among the fungi far greater than were at first imagined, and it is 
quite possible that widely different results may have quickly arisen under the 
pressure of peculiar life conditions. 

The study of multinucleate gametes has opened an interesting line of 
investigation, and promises results that may materially modify our views of 
the evolution and differentiation of en cre ae among the Phycomycetes, 
and perhaps the Ascomycetes.— B. M. Davis. 


IN A RECENT PAPER Wager gives an interesting account of his observa- 
tions on Euglena viridis as they bear on the functions and relations of the 
eye spot and flagellum. After a brief résumé of the general morphology of 
the cell he takes up the vacuole system and gullet. He states definitely that, 
contrary to previous observations, the principal vacuole opens directly into 
the gullet, and therefore is an excretory reservoir. Just at this point Wager 
takes issue with the zoologists’ claim of holozoic nutrition. Using powdered 
carmine in the culture medium he failed to find a single grain entering the 
gullet. He also brings forward tentatively Kawkine’s explanation that the 
gullet is an absorptive region because paramylum grains are smallest in close 
proximity to it. Wager’s views on the structure of the eye spot antagonize 
some older ideas and support others. In brief, the eye spot is composed of 
granules, bright red in color, imbedded in plasmatic network. The granules 
are in a single layer and with no regular arrangement. On treatment with 
alcohol, a reaction similar to that of disintegrating chlorophyll grains is 
obtained; hence the eye spot coloration is a derivative from chlorophyll, 
The origin of the spot de novo is in doubt. The flagellum and its close 
physical connection with the eye spot takes up the next paragraph. Nothing 
is known of its mechanism, however. Its structure is simple; a single fila- 
ment with bifurcate base, bearing a swelling on one of the bifurcations. The 


£gor | CURRENT LITERATURE 141 


base is attached to the posterior side of the excretory reservoir. The swell- 
ing mentioned lies against and below the concave side of the eye spot. This 
fact leads to a consideration of the effect of light on Euglena. As is the 
case in all motile cells, strong light repels and a moderate light attracts. A 
bright light will cause the active cells to round up and.encyst, if the stimula- 
tion be continued for several days. In darkness the cells round up, lose fla- 
gella, and divide. In spectrum rays, over seventy per cent. are drawn into 
the green-blue field. These blue rays are those absorbed by the red eye 
spot. As to the function of the eye spot, Wager makes two suggestions. First, 
that the absorbed blue rays stimulate the eye spot, which in turn stimulates 
the swelling on the flagellum ; second, that by cutting off certain rays, the eye 
spot produces a definitely unequal illumination of the enlargement, and as a 
result, an attempt at orientation. Both hypotheses, however, he puts forward 
tentatively, subject to further and more careful investigation.—PHILIP 
WRIGHTSON 


R. A. ROBERTSON (Trans. and Proc. Bot. Soc. Edinburgh 21: 290-298, 
Pés. 3. 1900) has recorded and illustrated some interesting observations on 
variations in Lycopodium clavatum. A luxuriant patch of this plant growing 
in a wood became exposed by a great storm which removed nearly every 
tree, and upon 20 to 30 per cent. of the erect axes the observed variations 
were found. Of this varying material 87 per cent. had extra branching of 
the strobilus-bearing axes, 66 per cent, showed branching of the strobili, and 
in 9 per cent. the strobili were completely metamorphosed into leafy shoots. 
The author suggests that these variations are of interest in reference to the 
phylogeny of the Lycopodiales, the branching of the strobilus and the steriliza- 
tion of sporogenous tissue being supposed to have played a part in the evolu- 
tion of the group.—J. M. C 


THE LITERATURE of lenticels has been further supplemented by James 
A. Terras (Trans. and Proc. Bot. Soc. Edinburgh 21: 341-352. Aés. 2. 1900), 
who has written upon the relation between the lenticels and adventitious roots 
of Solanum Dulcamara. We concludes that these roots do not arise below 
or grow out through lenticels, as is apparently the case in the majority of 
plants, but that their origin is entirely independent of the formation of lenti- 
cels. Furthermore, he states that the protuberances on the surface of the stem 
are not lenticels, but result from the formation of a mass of secondary tissue 
which originates in the reaction of the phellogen to the pressure set up by the 
elongating root below it. The lenticels only appear after the protuberances 
are fully formed.—J. M. C 


NEWS. 


THe Fern Bulletin for January contains an excellent portrait of Professor 
L. M. Underwood. 

WE REGRET to learn of the death of Dr. S. J. Korshinsky, of St. Peters- 
burg, at the age of 40. He had done some excellent work, especially in 
problems concerned with the vegetation of the steppes. 

Dr. M. RacrgorskI, formerly professor of botany at the University of 
Cracow, and lately of Buitenzorg, has been appointed professor of botany and 
director of the botanical garden in the Dublany Agricultural Academy at 
Lemberg. 

ONE GARDEN scholarship is to be awarded by Director Wm. Trelease, of 
the Missouri Botanical Garden, before April 1 next. Applications must be 
in his hands before March 1, and examinations will be held at the garden 
on March 5. 

THE FRENCH ACADEMY OF SCIENCES has awarded the Desmaziéres prize 
to Abbé Hue for his ‘‘ Mémoire sur une Classification des lichens fondée sur 
leur anatomie,” and the Montaigne prize to M. J. Heribaud for his ‘Les 
Muscinées d’Auvergne.”’ 

THE VIENNA Academy of Sciences intends to send an expedition in 
Ig0I to study the flora of Brazil. Professor Dr. Richard von Wettstein, 
director of the botanical garden of the University, and Dr. Victor Schiffner 
of Prague, will accompany the party. 

THE BOTANICAL DEPARTMENT of the British Museum has purchased the 
Bescherelle herbarium of mosses and liverworts, the collection containing 
14,800 specimens of the former, and 3500 of ahe latter. The herbarium is 
very rich in types and authentic specimens. 

THROUGH the liberality of T. M. Baird, Jr., Esq., of Victoria, B. Cae 
tract of land on the coast of Vancouver island, opposite Cape Flattery, has 
been presented for a seaside botanical station of the University of Minnesota. 
The erection of a group of log buildings has been begun and a party of thirty 
or more botanists has been organized to open the work of the station next 
June. 

AT A MEETING of the Botanical Section of the Philadelphia Academy of 
Natural Sciences, held December to, 1900, the following officers were elected 
for the ensuing year: Director, THOMAS MEEHAN; Vice director, GEORGE 
M. BERINGER; Treasurer and Conservator, STEWARDSON BRowNn; Recorder, 

142 | FEBRUARY 


1901 } NEWS 143 


Joun W. HARSHBERGER; Executive Committee, GEORGE M. BERINGER, 
THOMAS MEEHAN, STEWARDSON BROWN, JAMES D. CRAWFORD, IDA A. 
KELLER. 

THE REPORT of the committee of the Society for Plant Morphology and 
Physiology on securing better reviews of botanical literature was presented 
at the recent meeting of the society at Baltimore. The results secured are 
of the greatest interest to all botanists. After correspondence with the editor 
and publisher of the Botandsches Centralblatt, a modification of this journal 
will be effected with the first number of the present year, and these changes 
are officially announced in the last number for 1900. Hereafter the Centra/- 
blat¢t proper will contain only reviews and the list of new literature, while the 
Bethefte will’contain only original articles, and will appear as heretofore at 
irregular intervals, These may be subscribed for separately; the price and 
size of the former will remain as at present, 1/28 per year. A committee of 
three from the society, Drs. Farlow, MacDougal, and Ganong, together with 
one botanist from the central states and one from the Pacific coast, to be 
_ named later, will be asked to nominate American editors, who will cooperate 
with the German editors, Drs. Uhlworm and Kohl. Inasmuch as the editors 
of the Centra/blatt have shown themselves so willing to take up the sugges- 
tions made by this committee it should now be a matter of pride, and it cer- 
tainly is a matter of self-interest as well, for American botanists to give the 
fullest support, financial and scientific, to this bibliographic journal. It 
should remain unique and become as perfect in its notices of literature as it 
is possible to make it. Separates, not titles merely, should still be sent to 

r. Uhlworm, Cassel. 

THE following action was taken by the employés of the Department of 
Agriculture in reference to the death of Mr. Thomas A. Williams: It is with 
feelings of profound sorrow and regret that we, the employés of the Depart- 
ment of Agriculture, learn that death has removed from among us our 
beloved companion and coworker, Thomas A. Williams, a classmate and inti- 
mate associate of many of our number. His broad knowledge of scientific 
matters, his keen appreciation of nature, his kind and forbearing disposition 
have endeared him to all. He was known to his associates as an indefatigable 
worker and investigator. 

Resolved, That in the death of Professor Thomas A. Williams, science and 
agriculture have suffered a great loss, and we, his associates, a dear friend, 
whose self-forgetfulness in his kindly consideration for the feelings of others 
and uniform cheerfulness, often under conditions of severe physical suffering, 
revealed a lovable character of the highest Christian type. The loss to the 
Division of Agrostology is irreparable. In the performance of his official 
duties he had proved himself an excellent executive and an organizer of 
unusual merit, and his relations with his associates in office were always such 


ae es oe 


144 BOTANICAL GAZETTE | FEBRUARY, I9gOI 


as to command the highest esteem and respect. He never shirked a duty, 
and however difficult the undertaking the work performed by him was done 
most creditably. His loss will be felt most keenly by his associates in the 
division, and his memory will remain with them as one whose exemplary 
life and steadfastness of putpose they should strive to emulate. 

Resolved, Further, that we tender to his bereaved family our heartfelt 
sympathy in their great loss and invoke for them the blessing of the Heav- 
enly Father, who alone can heal the broken heart and give Jasting comfort. 

Resolved, That copies of these resolutions be sent to the family of the 
deceased and to the various daily and scientific journals.— Committee on 
Resolutions for the Department of Agriculture. 


FROM ADVANCE sheets of the twelfth annual report of the director of the 
Missouri Botanical Garden we make the following extracts: 

The garden has been maintained through the past year on about the same 
lines as for some years past. Though the revenue has not yet been increased 
through sales of real estate, the power to make such sales now makes it 
possible to spend on the garden the entire current revenue, which adds 
annually ten or fifteen thousand dollars to the available funds. The exten- 
sions which have been contemplated for some years past have now been 
begun. The improvements made this year consist in the grading of some 
twenty acres of land, and in planting the border about this tract. Two rather 
large ponds, connected by a meandering brook, were made, and the ground 
was given an easy fall from the streets to these bodies of water. All of the 
material used in this border plantation is. representative of North American 
plants, and it is proposed to plant on this tract a collection of 181 arborescent, 
269 fruticose, and some 1400 herbaceous species, representative of the 
North American flora, and arranged essentially in the familiar sequence of 
families of the ‘‘Genera Plantarum” of Bentham and Hooker. The garden 
now contains 9194 species and varieties of plants, of which 5547 are annuals 
or hardy perennials, and 3647 are cultivated under glass. A further exten- 
sion of the plant houses has been made by the erection of a tower at the 
northwestern corner of the system, in which small but representative collec- 
tions of succulents and of acacias and acacia-like plants have been planted 
out in a natural manner. The material incorporated in the herbarium during 
the year comprises 8415 sheets of specimens. The additions to the library 


456 pamphlets presented or sent in exchange for garden publications. Of 
serial publications the garden now receives 1117, all but 66 by exchange- 
The garden has materially assisted the botanical work in the public schools 
by supplying material. Several pieces of research work are about ready for 
publication. 


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Hot Springs Physicians Use Per- 
sonally and Prescribe for Patients 


>| BUFFALO 
+; LITHIA 
WATER 


IN THESE CONDITIONS AND FOR ALL URIC ACID POISONING. 


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Dr. = C. Laird, Resident Physician, Hat Springs, N. Cc. J ae 
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SUT, RHEUMATIC GouT, RHEUMATISM, BRIGHT’S DISEASE, RENAL 
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SE SAPOLIO. 


a ae 


Vol. XXXI MARCH, Igo1 ‘No. 3 


hie | THE 


BOTANICAL GAZETTE 


EDITORS 
JOHN M. COUETER anp CHARLES R. BARNES, 


WITH OTHER MEMBERS OF THE BOTANICAL STAFF 
OF THE UNIVERSITY OF ‘CHICAGO 


ASSOCIATE EDITORS 


{we ae pre UR 
Purdue University 
CASIMIR Dec DECANDOLLE 


J. B. DETONI 

“ati of tees 
ADOLF ENGL 

eee of Berlin 
LEON GUIGNARD 

£ Ecole de ersig Paris 

ROBERT A. HARP 

Cniversity " Wisconsin 
JINZO MATSUM MUR 

: imperial canes Tokyo 


FRITZ NOLL 

University hei Bonn 
VOLNEY M. ea 

Ont: sitet ” ‘iichigen 
ROLAND Pies 

ae Pigebelss 

WILLIAM pega ASE 

Missouri et Garden. 
38 gina tee oe 

pest = Fe cixotnes 

EUGEN. baat 

Oni ibe of Copenhagen 
VEIT WITTROCK 

Royal Academy id Setences 


CHICAGO, ILLINOIS 


| Publisher by the Gnibersity of Cities 


€hbhe Bniversice of Cbicage meted 


ann re re ene 


SEN SPE eee Lane eR 


i 


Botanical Gazette 


A Montbly Fournal Embracing all Departments of ae ena Science 
Subscription per year, umbers, 40 Cents 
The gosiecren a ies must be paid in advance. No numbers are sent ie the expiration 

time paid for. No reduction is made to dealers or agents 


FOREIGN AGENTS: 
t Britain — Wm. WeEsLEY & Son, 28 Esse 
St., Strand, London. 18 Shillings. 


Vol. XXXI, No. 3 


ermany — GEBRUDER BORNTRAEGER, Berlin 
. 46, ns SB eee 17a, [> 18 Marks 


Issued March 16, 1901 


CONTENTS 


THE PHYSIOGRAPHIC ECOLOGY OF CHICAGO AND bie pea A oe OF THE 


L BOTANICAL LABORATORY. pent vir THIRTY-FIVE 
FIGURES). Henry Chandler Cowles 


145 
SOME RECENT PUBLICATIONS AND THE NOMENCLATORIAL PRINCIPLES THEY 
MP RESENT. A0..2; Pita 293.5 Ooo ee Boi Seon, See 
BRIEFER ARTICLES. 
OBSERVATIONS UPON THE FEEDING PLASMODIA OF FULIGO SEPTICA hie ONE 
FIGURE). John W. Harshberger - 198, 
SWARM era FORMATION IN HYDRODICTYON UTRICULATUM Roth. A. G. Timberlake 203 
CURRENT inser TURE. 
BOOK REVIE ‘ = z 2 Z ' “ ‘ ~ 204 
GOEBEL’S Disieanee 
“MINOR NOTICES - . - - - - : é % oe 
NOTES FOR STUDENTS - - - - - : : - 208 
NEWS ‘ . f = es ‘ ‘ : - - - 216 


aa g of plain text or text with sw engravings. The actual cost may v the figures given, 
will depend upon the amount in re-making the pages into forms, press work, paper, binding 
€parates containing half-tones may be expected to cost somewhat more a9 the rates given, the 
ineiae depending upon the number of cuts and the amount of work required upon them. 
Number of copies 50 100 150 = 
I oc. for 4 pagesorless.. . . . $1.60 $2.00 $2.25 $2.50 
Letter tos, fur e pa ra Age (og aig ue a aan 2.25 2.75 3-15 3-50 
Lette er-press, for 16 pages et eG 4.00 5.00 5.80 6.50 
Single plates (1 dei ble = eee: ingle 1.00 1.35 i aia 
Overs, with title (paper like GAZETTE cover), 1.50 2.00... 2.50 300 
ea ea 


Manuse ‘uscripts. — Contributors are requested to write snip and i 8 names with particular care, 
aud in citations to Lalli t 
Editor of the Botanical 


he form shown in the pa ae es of t he GazerrE. Manuscripts should be sert to 
Book So 
and Pam) for ait Me aca eo tees same addre 
, will be replaced ioaty only when claim is made within thirty days after receipt of the 
ener folowing. 


emittances should be made payable - ene order of The University of aoa oe! ne 
mel rréspon ndence regarding subscripti tio 
University of Chee on on "Il. 


any cond-class mail matter. ] 


Every Botanist 


Should be familiar with 
the prominent works of 


GEBRUDER BORNTRAEGER 
Publishers. 


Die Glykoside. Chemische Monographie der Pflanzengly- 
koside nebst systematischer Darstellung der kiinstlichen 
Glykoside von Dr. I. I. L. von Ryn, Director der Reichsver- 
suchsstation in Mastricht. 8vo. Cloth, $2.50. 

Das Werk giebt—wite es bisher noch nirgends geschehen — eine eingehende 
chemische Behandlung der Glykoside—nicht nur eine kurzgefasste Zusammen- 
stellung der chemischen Eigenschaften dieser Kérperklasse, sondern die Darstet- 
lungsmethode, die Griinde, welche zur Aufstellung der Constitutionsformeln 
gefihrt haben ete., so dass das Buch in chemtsch-pharmaceutischen wie pharma- 
hologischen Kreisen sowie unter den Studirenden und sonstigen Freunden der 
phytochemischen Forschung sicher mit grosser Freude begriisst werden wird. 


Die Harze und die Harzbehalter. Historisch- 
kritische und experimentelle, in Gemeinschaft mit zahlreichen 
Mitarbeitern ausgefihrte Untersuchungen von Proressor DR. 
A. Tscuircn, Director des pharmaceutischen Institutes der Uni- 
versitat Bern. Mit 6 Tafeln. 8vo. Half calf, $5.00. 

Das Werk stellt zum ersten Mal das gesammte Material dieser wichtigen 
Gruppe von Phlanzenproducten kritisch durchgearbettet dar. Die streng wassen- 
sthaftlichen Untersuchungen werden auch fiir die Praktiker, besonders fiir de, 
die sich mit Harzen und Harzproducten beschdftigen, von Interesse sein, da sede 
rationelle Technik ja auf wissenschaftlicher Grundlage ruht. 


Write for free Catalogue; postpaid. Address: 


Gebruder Borntraeger, Publishers, 
Bertin, S. W. 49. 


SCHONEBERGERSTRASSE 17a 


The Posi 


URING the coming year, THE binge TER, the piniaee nce mete y magazine of the 
American Bol a Rie tease vill be ore ing and va arr oe 
wh 


ever befor No oO cares ma trees, or life i apr woods, or who 

ested in the movement bg encourage the preserva sips al care of the forest, onl 
be without it. :: :: Among the con ein rsare: GIFF( ain Chief of the U.S 
Division of Forestry; Dr. B. E. FER 7, Dean of the New Y k State College of 
Forestry; HENRY enone Connie of the U.S. "Geological Survey 
GirrorD; Pror. HENRY S. GRAVE ES, of the Yale Forest School; Dr. C. A. Scitexer, 
of Biltmo ore, NaGe Ho JAMES WILSON, Secretary aoe Fea PR Wo. R. 
sche of Stan ford University, Cal.; PROF . SH R, of Harvard Unitas 

and many others of n ree au thority on their speci alti Hon Beside s a number of 
eeatdees articles, neon issue of the magazine will c cord of legis pits 
touching the geil of a country’s forests (of ihe ‘6 there will | probably be a great 
deal i the coming year), with beet comment, and r of recent pu ublica- 
tion ia pe 3 most rr ent exper Each n ne is handsomely i illustrated. 
H : For sample ete “send a ae cent stamp t 


‘he forester S09 14th St., &. w., washington, D. €. 


np oo in one payment. The Aescetetiens is engaged in work which is ‘of the reateat i im- 
portance, and for the progress of which all the moral and financial support obtainable is 
needed. To — the Association address the SECR ETARY, 202 iar Street, ‘S. Ww. Wash 
ington, : 


ee 


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f 28 Essex Street, Strand, eo London, England. 
a a a Oe ea) 


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RRS SSS SSS SSS SSS 


[JUST PUBLISHED] 


Send 12 cents, U. S. stamps, for 


sears & Son’s Botanical Catalogue, 1901 


— 


i, Bein CONTENTS: 
No. ie 138 of Transactions of Scientific Societies Periodicals , 
J THE a Bibliography §_ History Biographies and Portraits f 
Herbal: Early Botanical Science Linnaeus 2 
f N. ATUR AL £ erpdais y Bo 
Handbooks 
7 HISTORY and f : rere J 
SCIENTIFIC Microscopy Morphology an ysiology f 
J Ooo f Encyclopaedic works Classification Nomenclature ./ | 
= il Plants 
f CIRCULAR i 3 Cryptogams Phanerogams Fossil Plan 
f Natural distribution of Plants (Floras) 0 
ft BOTANY Agriculture and Horticulture to the end of the J8th century 7 
as More than 3300 works, JS Gardening Landscape Gardening 
classified under 42 f The Flower and Ornamental Garden { 
df headings Husbandry Tropical Agriculture Commercial Plants *. 
eee Medical Botany Forestry -—CDiseases of Plants J 


William Wesley & Son, Booksellers, 


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Announcements of 7Be UNIVERS 


OF CHICAGO PRESS 


The Assembly of Gods; or, The Accord of “3 
son and Sensuality in the Fear of Death. 

OHN LYDGATE. Edited by OSCAR yee 

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ies and Simile in the Minor oo 
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Parts five and six in preparatio 


RELIGION AND tL dace tpmestetd 


The Present Status of ls Epil nema” 
the Genuineness of the saline By 
BERNHARD Nba ati faerie D : hes ie 
8vo, paper, 50 

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VOLUME XXXI NUMBER 3 


BOTANICAL” GAZETTE 
MARCH, roor 


THE PHYSIOGRAPHIC ECOLOGY OF CHICAGO AND 
VICINITY; A STUDY OF THE ORIGIN, DEVELOP- 
MENT, AND CLASSIFICATION OF PLANT SOCIE- 
TIES, 


CONTRIBUTIONS FROM THE HULL BOTANICAL LABORATORY 
XXIV. 


HENRY CHANDLER COWLES. 
[Concluded from p. 108.] 
2. THE POND-SWAMP-PRAIRIE SERIES. 


A. The pond.— There are all gradations between rapid streams 
and completely undrained ponds, and corresponding with these 
various gradations are characteristic plant species. It will be 
convenient to subdivide the series under discussion into two 
parts, the first dealing with undrained ponds and swamps, the 
second with half-drained ponds and swamps. 

No two floras can be more unlike in species or in adaptations 
than are the typical brookside and swamp floras. Though each 
type may be called hydrophytic, so far as the water is concerned, 
the vegetation is really hydrophilous in the first case but pro- 
nouncedly xerophilous in the second. Peat bogs which may be 
taken as the type of undrained swamps have a remarkable assem- 
blage of xerophytic adaptations, such as leathery or hairy leaves, 
and special structures for water absorption. Schimper*® believes 

*3SCHIMPER : Pflanzengeographie, p. 18. 

145 


146 BOTANICAL GAZETTE | MARCH 


that these structures are due to the difficult absorption in peaty 
soil, the humus acids and the lack of oxygen being detrimental 
to normal root activities. For similar reasons the normal soil 
activities of bacteria and fungi are lessened, and as a result of 
this relative lack of decay great quantities of peat accumulate. 
All of these peculiarities of peat bogs may be referred to the 
lack of drainage, since the stagnant conditions prevent oxidation 
and the removal of the humus acids. The lack of drainage is of 
course due to topographic conditions. Peat bogs and undrained 
lakes, therefore, are features of a young topography, since several 
agencies combine to cause their rapid destruction. Rivers may 
work back and tap the undrained lakes or inlets may fill them up. 
Probably the most important agent in the death of undrained 
lakes, however, is the vegetation, as will be seen later. The 
great abundance of lakes and ponds in the young glaciated 
regions as compared with older regions to the south is a striking 
proof of their short life. 

In the immediate neighborhood of Chicago typical peat bogs 
are scarce. They find their best development in the depressions 
of the dune region, where they may be called abundant. Wher- 
ever a sag between two dunes is low enough to retain moisture 
for the greater part of the season, the conditions favor the 
development of an undrained swamp flora. If the depression is 
so low that the water level outcrops throughout the year, then 
there is an undrained pond or lake. The first flora in this latter 
case consists of plants that are able to exist with little or no 
change in the water of the pond except through rain and evap- 
oration. Among these plants the alga Chara takes a promi- 
nent place. The water lilies (Nymphaea and Nuphar) are an 
exceedingly important constituent of this first vegetation, as is 
also Utricularia, which is represented by several species. The 
above species, together with others, play a great part in filling 
up lakes, since their remains accumulate with almost no decay. 
Chara in particular is a soil former of great importance. The 
rapidity with which these filling processes are carried on is 
striking; in pools of known age among the rubbish heaps of 


Igor | PHYSIOGRAPHIC ECOLOGY OF CHICAGO 147 


Jackson park the author has noticed accumulations of Chara 
peat amounting to one or two inches per year. 

B. The undrained swamp.—It is obvious that the processes 
outlined in the preceding paragraph must eventuate in the death 
of the lake or pond involved and its replacement by a marsh, 
entirely apart from ordinary erosive activities. Indeed, as has 
been stated, these activities are relatively unimportant here; 
this fact is shown by the absence of ordinary sediments from 
most peat beds. As the aquatics make the pond shallower 
and shallower they make it more and more unfit for themselves 
and fit for their successors, viz., those plants which grow along 
pond margins. Among the first plants of this type are various 
sedges (Carex), also the bulrush (Scirpus lacustris), though this 
latter species is more characteristic of the half-drained margins 
than of those under discussion here. Other marginal plants of 
our peat bogs are Menyanthes trifoliata and Potentilla palusinis. 

The vegetation that follows may be called typical of peat 
bogs. The dominant plants are usually shrubs, especially the 
leather leaf (Cassandra calyculata) ; this plant may be so abun- 
dant as to give tone to the landscape. /%g. 19 shows some Cas- 
sandra islands in a sedge swamp. It is clear that the islands 
represent places where in the original lake the water was shal- 
ow. The present remnant of the lake is shown at the left. Not 
only have the sedge zones advanced upon it from all sides, but 
centers of sedge growth appear also in shallow places in the 
lake itself. Just as the sedge zone encroaches upon the lake, 
when conditions become favorable, so the Cassandra zone 
advances on the sedges. Again atree zone advances on the 
shrubs, as will be seen farther on. The zonal arrangement of 
plant societies that has just been seen is a feature of most peat 
bogs, and is due to the symmetry of lake and bog conditions. 
It will be observed that along the lake margin the zones advance 
toward a common center, while on the islands the advance is 
from a center. Eventually, of course, the marginal and island 
zones will merge. o 

Besides Cassandra many other plants are commonly found 


Ser's5 
Vs, 


Tond 


148 BOTANICAL GAZETTE [MARCH 


in the shrub zone. Other shrubs are the swamp blueberry (Vac- 
cinium corymbosum), the cranberry (Vaccinium macrocarpon), the 
dwarf birch (Betula pumila), the alder (Alnus incana), the hoary 
willow (Salix candida), and the poison sumach (Rhus venenata). 
Characteristic herbs, especially in the open places, are the pitcher 
plant (Sarracenia purpurea), the sundew (Drosera rotundifolia), 


Fic. 19.—Typical peat bog in a depression between established dunes at Miller. 
Relict of the original pond at the left. Sedges (light-colored vegetation) are encroach- 
ing on the lake, while paral, mainly Cassandra, are encroaching on the sedges. Cas- 
sandra islands toward the right. Advance of conifers on Cassandra (seen in its begin- 
nings on the islands) ae n at the extreme right. 


various orchids, as Calopogon pulchellus, Pogonia, and Cypripe- 
dium; sedges, as Eriophorum and Dulichium; Woodwardia Vir- 
ginica, and Elodes campanulata. One of the most typical plants 
of these places is the peat moss, Sphagnum. 

The flora just mentioned has many interesting features which 
are well known and may be passed over briefly. The highly 
xerophytic character of this plant society has already been 
noticed, and the reasons for it briefly given. The xerophytic 
structures are well illustrated in the leathery leaves of Cassandra 
and the absorption and storage adaptations of Sphagnum. 


Igor | PHYSIOGRAPHIC ECOLOGY OF CHICAGO 149 


Many bogs of this type are very spongy and unstable, whence 
the name quaking bogs; this feature is due to the rapid growth 
of the vegetation and the absence of ordinary inorganic soils for 
a considerable depth. The similarity of the peat bog vegetation 
throughout the northern hemisphere is one of its most striking 
features. Not only the adaptations but the species themselves 


- 20.—Tamarack swamp in an undrained portion of the Calumet flood plain 
at ‘fie. Peat bog herbs and shrubs in the foreground. 


are similar over vast areas; the conditions are unique and the 
flora also. None of our plant societies, not even the lakeward 
dune slopes, have sucha pronounced northern flora as do the peat 
bogs. No contrast could be more striking than that between 
the southern vegetation of the flood plains and the northern 
flora of the bogs. 

fig. 19 shows that a coniferous vegetation, now represented 
by but two or three small trees at the centers of the islands, is 
to follow the Cassandra. Such an advance of conifers on 
Cassandra is shown in the background at the right. The most 
typical conifer in such cases is the tamarack (Larix Americana) ; 
With this the arbor vitae ( Zhaya occidentalis) is sometimes found. 


150 BOTANICAL GAZETTE [MARCH 


Larix and Thuya swamps reach but an imperfect development 
in our region and little need be said about them. The shade 
in these forest swamps is so dense that bare patches of soil are 
often seen. The vegetation consists largely of shade plants, 
among which may be mentioned Mnium and other similar mosses, 
Coptis trifolia, Cornus Canadensis, Viola blanda, and Impatiens. The 
tamaracks appear to be succeeded by the pines (Pinus Strobus or 
P. Bankstana), and they in turn by oaks, as the soil becomes 
drier and better drained, and thus more adapted to deciduous 
trees. /ig. 20 shows a tamarack swamp near Miller, Ind. 

Not all peat bogs have a history like the above. Just as 
some flood plains are forested and others not, so some peat 
bogs grow up to shrubs and trees, while others are dominated, 
for a long time at least, by herbs and grasses. Fig. 27 shows a 
swamp of this character. Bulrushes are seen to be encroach- 
ing upon the water lily vegetation, while back of the bulrushes, 
instead of Cassandra, is a zone with sedges and grasses and 
scattered willows. Among the species, other than sedges and 
grasses in a plant society like this are Viola sagittata and V. lan- 
ceolata, Potentilla Anserina, Fragaria Virginiana, Parnassia Caro- 
lintana, Sabbatia angularis, Gentiana crinita, Gerardia purpurea, 
Castilleta coccinea, Aletris farinosa, Iris versicolor, Sisyrinchium 
angustifolium, Hypoxys erecta, Xyris flexuosa, Triglochin maritima. 
The shrubs in such places are chiefly Sadix glaucophylla, Cornus 
stolonifera, Potentilla fruticosa, Hypericum Kalmianum. The con- 
ditions that determine this type of bog, as contrasted with the 
Cassandra type, are not clear. The soil is hard, compact, 
shallow, and usually sandy ; it may be that this type develops in 
shallow depressions, while the type with spongy, quaking ground 
develops in deeper depressions. This second type much more 
closely resembles the half-drained swamps in its flora than does 
the Cassandra type, although so far as drainage is concerned it 
agrees with the Cassandra bogs. 

There is yet a third type of swamp which still more closely 
resembles the half-drained swamp in its flora. It is found along 
the edge of the Calumet valley near Dune park, also at West 


1901 | PHYSIOGRAPHIC ECOLOGY OF CHICAGO 151 


Pullman. In this case the soil is rather deep and rich, in which 
respects there is agreement with the first type rather than the 
second. Grasses and sedges, but of a more luxuriant type, domi- 
nate here also, and with them are found such plants as Cepha- 
lanthus occidentalis, Aspidium Thelypteris, Onoclea sensibilis, Saxifraga 
Pennsylvanica, Caltha palustris, Viola blanda, Polygala sanguinea. 


Fic, 21.— Shallow, undrained swamp (peat bog) at Dune park. Jn the foreground 
the relict of the original pond, with water lilies; then in order, encroaching zones 0 
bulrushes, sedges, willows, and pin The hei in the background are on an estab- 
lished dune, and are not sisting on the swampy soil. 


Sphagnum occasionally occurs here, as it never does in the 
second type. Here again there is doubt as to the determining 
conditions, but it may be that things can be explained by the 
difference in the drainage. The ultimate fate of the second and 
third Swamp types is not known, The relative absence of trees 
and shrubs is certainly natural and in no wise due to man. 


152 BOTANICAL GAZETTE [MARCH 


Possibly local prairies will be the final type, or it may be that 
the forest will come in. Fig. 2z, which shows pines encroaching 
upon the grassy areas, favors the latter view. So do some of 
the facts seen in the Calumet valley. 

All of the peat bog types have a characteristic marginal 
flora, 2. ¢., the vegetation at the margin of the original lake is 
essentially alike in all cases. These plants, as well as those of 


Fic. 22.— Encroachment of bulrushes on Calumet lake, showing how plants may 
destroy lakes. 


Cassandra bogs, are the same over wide areas. The most 
common members of the bog margin flora are the sour gum 
(Nyssa sylvatica), the aspen (Populus tremuloides), [lex verticu- 
lata, Pyrus arbutifolia (including var. melanocarpa) , Spiraea salict- 
folia and S. tomentosa, Rubus hispidus, Gaultheria procumbens, 
Osmunda cinnamomea, O. Claytoniana, O. regalis, Betula papy’ t- 
fera, and Polytrichum commune. This vegetation originates out- 
side the swamp, and may be regarded as xerophytic ; however, 
it often encroaches upon the swamp as the latter develops. At 
Thornton there isa dead swamp which is now almost entirely 
occupied by this xerophytic bog margin flora, only a few of the 
original swamp plants now remaining. Near Morgan park is a 


1901] PHYSIOGRAPHIC ECOLOGY OF CHICAGO 153 


bog margin flora without a bog; a shallow trench has been dug and 
in this trench there have appeared various peat bog plants, e. ¢., 
Sphagnum. These considerations show that bog margin floras, 
though associated with most bogs, are not necessarily genetically 
connected with them. 

A word may be said about undrained swamps among the 
active dunes. The conditions here, of course, are far more 
severe than in ordinary peat bogs and only a few species are 


Fic. 23.— Pond at Waukegan almost destroyed by bulrushes. 


able to endure in such a habitat. The most typical herb is /uncus 
Baltcus littoralis. Seedlings of the cottonwood, as well as the 
long-leaved and glaucous willows, germinate in these wet depres- 
sions. Reference will be made to these plants in connection 
with the dunes. 

In the morainic portions of our territory there are few if any 
peat bogs as described above, although they are usually more 
typical of moraines than of other topographic areas. On account 
of the clay soil which characterizes the morainic uplands there 
are many patches of swampy woods throughout the district. 
Shallow depressions of this type in sandy soil would not have a 
swamp developed. Morainic forest swamps are characterized by 
several trees, viz.: the bur oak, swamp white oak, and scarlet 


154 BOTANICAL GAZETTE [MARCH 


oak (Quercus macrocarpa, Q. bicolor, and Q. coccinea), the red 
maple (Acer rubrum), the elm (Ulmus Americana) , and the ash 
(Fraxinus Americana). Other species are Cephalanthus occidentalis, 
Salix discolor, Ribes floridum, Cardamine rhomboidea purpurea, 
Ranunculus septentrionalis. This vegetation is ultimately sup- 
planted by the mesophytic forest. A vegetation allied with that 


24.—Typical grass prairie near Pullman. This prairie has been reclaimed 


Fic 
feanniadig from Lake Calumet, and has passed through bulrush and sedge stages. 


of swamps is the amphibious ditch flora with such plants as Vas- 
turtium palustre, Penthorum sedoides, Proserpinaca palustris, Lud- 
wigia palustris, Polygonum Hydropiper, etc. 

Calumet lake and Grand Calumet river may be taken as 
types of half-drained waters. We have here conditions that are 
midway between those of peat bogs and those of ordinary rivers. 
The vegetation is subject neither to the currents of the rivers 
nor to the stagnant conditions of the peaty lakes, and hence the 
luxuriance of the flora is far greater than in either of the other 
instances. The aquatic vegetation is rich both in species and 


Ig0I } PHYSIOGRAPHIC ECOLOGY OF CHICAGO 155 


individuals. Here is to be found a great wealth of alga vegeta- 
tion, including such forms as Cladophora, Spirogyra, Oedogon- 
ium, Hydrodictyon. Among the floating plants are Riccia, 
Ricciocarpus, Spirodela, Lemna, and Wolffia. There are also a 
large number of attached plants, including many species of Pota- 
mogeton, Ranunculus aquatilis, Brasenia, Nelumbo, Myriophyllum, 
Ceratophyllum, Elodea, Vallisneria, and Naias. This rank 
growth of vegetation fills the lake up rapidly, since the currents 
are not sufficient to carry off the plant remains. There isa rapid 
advance of marginal plants upon the lake, a phenomenon that is 
shown in fig. 22, where the scattered bulrushes (Scirpus lacustris) 
are seen to be soon followed by a dense bulrush society. With 
or soon after the bulrushes are a number of marginal plants, 
especially 7ypha latifolia, Pontederia cordata, Sparganium eurycar- 
pum, Sagittaria variabilis and S. heterophylla, Zizania aquatica, 
Phragmites communis, Acorus Calamus, and Eriophorum cyperinum. 
fig. 23 shows a stage in which a lake has been all but destroyed 
by a rank bulrush vegetation. 

C. The prairie—Sedges encroach rapidly upon the bulrushes 
as the new soil becomes raised more and more above the lake, 
and grasses in turn encroach upon the sedges, forming a prairie. 
fig. 24 shows an expanse of grassy prairie which has developed 
through these successive stages from Calumet lake. Skokie 
marsh and Hog marsh are undergoing transformations of this 
character also. Sometimes with the prairie grasses are a number 
of coarse xerophytic herbs, largely composites (Si/phium lacini- 
atum, S. terebinthinaceum, S: integrifolium, Lepachys, Solidago 
rigida, Aster, Liatris), with some legumes (Amorpha canescens, 
Petalostemon, Melilotus, Baptisia), Eryngium, Dodecatheon, 
Phlox, Allium cernuum. A Silphium (compass plant) prairie is 
shown in fig. 25. The prairies of our area are in the basin of 
the glacial Lake Chicago, and hence all may be referred to a 
lake or swamp origin, exactly as prairies are developing from 
Calumet lake today. This explanation of the prairie, an 
undoubted explanation for the cases in hand, must not be 
applied to the great climatic prairies farther west. Whether 


156 BOTANICAL GAZETTE [MARCH 


the Chicago prairies will ever become forested is a question not 
easily answered. There are signs of it in some places, as at 
Stony island, but this topic needs more detailed treatment than 
can be given here. 

The processes outlined in this section are rapid. The meso- 
phytic prairie or forest develops from the lake or marsh, while 
the region as a whole still retains a young topography. Thus 


Fic. 25.— Prairie at Pullman in which the compass plant (Silphium) grows with 
the grasses. This prairie is much older and drier than that shown in fg. 24. 


this mesophytic assemblage, like that of the ravine slope, is 
bound to pass away, though its life tenure is much longer. 
Sooner or later river action will enter; there will be developed 
ravines, xerophytic bluffs, and ultimately flood plains, again 
with a mesophytic flora. A broad survey then shows a rapid 
development to a somewhat prolonged temporary climax, and 
finally after ravine and bluff vicissitudes there appears the true 
and more enduring climax of the mesophytic flood plain. 


¥ 3. THE UPLAND SERIES. 
A. The rock hill—While all of a land area is eventually 
worked over by stream activities and can thus be referred to the 


~~ —.......-- 


1901] PAYSIOGRAPHIC ECOLOGY OF CHICAGO 157 


river series, other activities are at work in a young topography. 
The swamp series which has just been discussed is one illustra- 
tion. So also there are hills which are not due to erosive 
processes, but to other causes, notably in our region morainic 


"IG. 26.— Slope of limestone ledge at a island, showing mosses and higher 
plants aubaiie themselves in the crevice 


hills and sand hills. There are rock hills also which are. not 
connected with the present erosion cycle. All of these hill 
types have their peculiar vegetation features and must be dis- 
cussed apart from river activities, since they have an interesting 
history before they are attacked by stream erosion. 

We may speak first of rock hills, which in the vicinity of 


158 BOTANICAL GAZETTE [MARCH 


Chicago are quite rare and consist entirely of dolomitic Niagara 
(Silurian) limestone. Not only are hills of this limestone quite 
rare, but surface outcrops of any kind are uncommon because of 
the heavy drift. Hence the rock vegetation of the Chicago 
area is not very important. Perhaps the most interesting out- 
crop is at Stony island, where it is quite easy to trace the 
various stages in the development of the vegetation. This 
rock, like most limestone, is subject to chemical as well as 
mechanical erosion, but is much more resistant than most lime- 
stones on account of its strongly dolomitic character. The first 
vegetation that gets a foothold is composed of lichens, but the 
lichen flora appears to be rather sparse, perhaps because of the 
chemical nature of the rock, since lichens are commonly sup- 
posed to shun calcareous soils. The relative poverty of lichens 
may be due, however, to the easy solution of the surface rock 
layers and the consequent difficulty in retaining a foothold. 
The limestone is considerably jointed and fractured and there is 
in consequence a rich crevice vegetation, composed of several 
mosses, especially Ceratodon and Bryum, and also various 
grasses. Fig. 26 shows a vegetation of this nature, and among 
the other crevice plants is an abundance of Solidago nemoralis. 
Other species growing in the crevices or on the first soil which 
is formed on the rock face are Potentilla arguta, Verbascum T; hap- 
sus, Heuchera hispida, Poa compressa, etc. At Thornton there is 
a rock outcrop which gradually recedes from the surface, and it 
is possible to tell by the vegetation where the rock surface 
dips considerably under the surface of the soil. Where the soil 
is shallow the dominant plant is Poa compressa, but as the soil 
layer deepens it becomes gradually replaced by Poa pratensis. 
Similarly at Stony island crevices can be distinguished in a 
covered horizontal rock surface by a sudden change from the 
xerophytic plants of the shallow soil, that hides most of the 
rock, to the mesophytic plants of the deeper soil which lies 
over the crevices. 

’ Through rock decay and the accumulation of organic matter 
a considerable soil comes to be developed where there was at 


Igor | PHVSIOGRAPHIC ECOLOGY OF CHICAGO 159 


first an outcrop of bare rock. The opportunity for a shrubby 
vegetation eventually arrives, especially in the crevices. Fig. 
27 shows such a vegetation getting a foothold. Among the 
shrubs in such places are the chokecherry (Prunus Virginiana), 


- . ~ 
- ‘ es 
f ty 5 ial ‘ie a 8, 5 


27. ~~ Limestone ledge at Stony island, showing vegetation farther advanced 
than in a 26. The crevice shrubs here are chokecherries (Prunus Virginiana). 


ninebark (Physocarpus opulifolius), poison ivy (Rhus Ti oxicoden- 
dron), Rosa humilis, sumach (Rhus typhina), hop tree (Ptelea tri- 
foliata), wild crab (Pyrus coronaria). Still later the way is open 
for a tree vegetation, at first xerophytic, but ultimately meee: 
phytic, as the author has frequently observed in the Alleghanies. 
There can be no doubt but that a temporary mesophytic climax 
can be reached even on rock hills, though the probability of this 


160 7 BOTANICAL GAZETTE [MARCH 


is much greater where the hill is composed of limestone than in 
the case of sandstone or granite. 

B. The clay ill—— Morainic hills are common in the Chicago 
region and almost without exception they are covered with a 
mesophytic forest, in which the dominant trees are usually the 


Fic. 28.—Typical upland clay (morainic) forest at Beverly hills. The dominant 
trees here are red oaks (Quercus rubra), though a white oak (Q. a/éa) is shown at the 
extreme right. 


white oak ( Quercus alba), the red oak (Quercus rubra), and the 
shell-bark hickory ( Carya alba). This is easily the dominant 
forest type of the Chicago region, and is remarkably character- 
istic of morainic areas. The soil in all cases is a glacial clay or 
till, heterogeneous in composition, but rich in food salts. Of 
all our plant society life histories these are about the most diffi- 
cult to unravel and it is due to the favorable conditions under 
which they have developed. After the continental glacier left 
this region for the last time, it was doubtless on these low 


Sf aa ee 


1901] PHYSIOGRAPHIC ECOLOGY OF CHICAGO 161 


morainic hills that the first mesophytic forests were developed. 
And they have been developed for so long that almost no traces 
of their history are left behind; we have only the completed 
product, the mesophytic forest. 

Where these mesophytic forests are disturbed we may per- 
haps get some notion of what took place in the first postglacial 
centuries. On the clay banks along the drainage canal and also 
on recent river bluffs, one may follow in rapid succession a series 
of plant societies leading to the forest. There is here no pro- 
nounced lichen or moss stage as on rock hills, but the first vege- 
tation consists of xerophytic annuals and perennial herbs. 
Xerophytic shrubs, especially Salix and Populus, soon appear. 
It is not long before there is an extensive thicket formation with 
an herbaceous undergrowth. Humus accumulates with great 
rapidity and we soon have almost a mesophytic vegetation in 
which the dominant thicket species are likely to be the aspen 
(Populus tremuloides\, wild crab (Pyrus coronaria), red haw 
(Crataegus punctata, C. coccinea, etc.). Such a thicket is the 
immediate forerunner of the oak-hickory type of mesophytic 
forest. When a forest of oak and hickory is cut down or 
destroyed by fire it returns after a comparatively short interval, 
but the first stages in the clearing are thicket stages much like 
those just described. Of course it takes much longer to develop 
a forest from naked clay soil than from a forest land that has 
been cleared. Whether the stages that led up to the first post- 
glacial forests are such as have been described is very doubtful. 
It is much more likely that the first forests were of slow growth 
and were coniferous in character, such as are found farther 
north. Fig. 28 shows a typical morainic hill forest of the above 
type. Here the dominant tree is the red oak; a white oak is 
seen at the right. 

Among the shrubs of these morainic forests there may be 
mentioned, apart from the crabs and haws, the hazel (Corylus 
Americana) , and various species of Viburnum. Many herbaceous 
plants are found, among which are Podophyllum, Claytonia, 
various species of Aster, Trillium, Geranium maculatum, Viola 


162 BOTANICAL GAZETTE [MARCH 


pubescens, Anemone nemorosa, etc. Sometimes the bur oak ( Quer- 
cus macrocarpa) is the dominant tree in these morainic forests, 
though in such cases the habitat is usually more moist or else 
the drainage is less perfect. A bur oak forest is shown in Fig. 
29. The transition from this type to the morainic swamp for- 
ests, already mentioned, is an easy one, and bur oaks are often 


1G. 29.—Typical forest of low morainic clay soil, made up chiefly of bur oak 
(Quercus macrocarpa). 


found with the swamp white oak and other species characteristic 
of such places. , 

In spite of the abundance of the type of morainic forest 
described above, it is scarcely probable that it is anything more 
than a very slowly passing forest stage. The fact that in all 
directions from Chicago the ultimate forest type on morainic 
uplands is not the oak-hickory but the maple-beech forest leads 
us to expect that here. This latter type seems to be of a higher 
order in all respects. It is found in richer soil where the humus 
content is very great. Seedlings of the beech or maple can 
easily grow in the relatively light oak forest, whereas oaks cannot 
grow in the denser shade of the maple or beech. Furthermore, 


| 


1901] PHYSIOGRAPHIC ECOLOGY OF CHICAGO 163 


oak forests have been seen with a pronounced undergrowth 
of beech. It would seem that one of the chief factors in deter- 
mining the order of succession of forests is the light need of 
the various tree species, the members of the culminating forest 
type being those whose seedlings can grow in the densest forest 
shade. There are evidences that the oak forests about Chicago 
are being succeeded by the beech or maple. The best instance 
of this which the author has seen is on the low moraines along 
the Desplaines river west of Deerfield. The sugar maple (Acer 
saccharinum) has already been mentioned as a character plant of 
the temporary mesophytic forests of ravines. Here we see it in 
the more permanent forest of the morainic hills. The beech 
(Fagus ferruginea) is much rarer than the sugar maple, though it 
is a rather important constituent of the mesophytic forests about 
Chesterton. Why the beech-maple forest has lagged so far 
behind in the region about Chicago is a question not yet settled. 
If these forests elsewhere have had an oak stage it indicates 
that the development here is very slow. 

Though the forests just described, whether of the oak-hick- 
ory or the maple-beech type, are of a high degree of permanence, 
it can be seen that this permanence is but relative. Sooner or 
later stream action will enter these districts and base leveling 
processes will begin on a more rapid scale. But for these activi- 
ties the lowering of hills would be very slow indeed, so slow as 
hardly to interfere at any point with a luxuriant development of 
the vegetation. The destruction of these morainic forests by 
stream erosion is well shown near the shore north of Evanston 
and also along Thorn creek. ig. 78 shows a morainic island 
in a flood plain, the sole remnant of an extensive stretch of 
upland mesophytic forest. We must therefore regard upland 
- forests as temporary also, though they endure for a much 
longer time than do the temporary mesophytic forests of the 
ravines. 

C. The sand hill—A third type of upland is found in the 
sand hills, but since most of these in our district are of dune 
origin, their treatment will be deferred until later. 


164 BOTANICAL GAZETTE [MARCH 


B. The coastal group. 
I, THE LAKE BLUFF SERIES. 

The plant societies that have been discussed hitherto may be 
found in many if not‘in most inland districts. The societies that 
follow, on the other hand, are best worked out only in connec- 
tion with the coasts of oceans or great lakes. Theoretically a 
bluff may be composed of any kind of rock or soil, but those of 
our area are composed of morainic clays, and the life histories 
that follow will not hold good in other conditions. It may be 
noted here that there is a short stretch of rocky shore with litho- 
phytic algae at Cheltenham, but there is nothing that in any way 
approaches a rock cliff. 

Wherever a sea or lake erodes rather than deposits, there is 
commonly developed a sea cliff of greater or less dimensions. 
The material which is thus gathered may be deposited elsewhere 
in the form of beaches and later the wind may take up the sands 
from the beach and form dunes. The Chicago area gives splen- 
did examples of these two types of sea activity; to the north of 
the city is an eroding coast line with its bluffs, and to the south 
and southeast is a depositing coast with extensive areas of beach 
and dune. 

The lake bluffs at Glencoe give an excellent opportunity for 
the study of the life history of a sea-cliff vegetation. There 
can be almost no other ‘habitat in our climate which imposes 
such severe conditions upon vegetation as an eroding clay bluff. 
The only possible rival in this regard is a shifting dune, and even 
here the dune possesses some points of advantage so far as the 
establishment of vegetation is concerned. In the first place, the 
conditions as to exposure are almost identical with those of a 
dune: the heat of midday and of summer and the cold of night 
and winter are extremely pronounced; the intensity of the light 
and the exposure to wind make the conditions still more severe: 
In other words the only plants that can grow on these lake bluffs, 
at least in the earlier stages, are pronounced xerophytes. Again 
the character of the soil is unfavorable, for while the clay is wet 
in the autumn, winter, and spring, it dries out in the summer and 


1901] PHYSIOGRAPHIC ECOLOGY OF CHICAGO 165 


becomes almost as hard as rock. In the heart of summer the 
conditions for vegetation are no better on the hard dry slopes 
of a clay bluff than on the hot, dry sands of a dune. Finally as 
to instability: it is doubtless the constant shifting of the sand 
which in the last analysis accounts for most of the poverty of 
the dune vegetation. It is similar on clay bluffs, for when the 


Fic, 30.—Sea cliff along the eroding shore at Glencoe, exposing the morainic 
clay. Vegetation almost entirely absent. Projecting turf mats at the top show the 
tenacity with which the vegetation holds its ground in the face of the erosive forces 


Waves undermine the cliff at its base, the action of gravity 
Causes great masses of material to fall down from the entire cliff 
face. Furthermore, when the clay is saturated with water, great 
portions of the cliff face slide down, entirely apart from the 
action of the sea or lake. At no time, then, is an eroding bluff 
any more stable than a naked dune. 

It becomes evident from a survey of the bluff conditions 
that all vegetation is impossible so long as active erosion by the 
lake continues. Not only this, but vegetation at the top of the 
bluff is soon destroyed. Fig. 30 shows a naked cliff of this 


166 BOTANICAL GAZETTE [MARCH 


character ; at the top there can be seen overhanging turf, giving 
evidence both of the destructive action of the lake and also of 
the tenacity with which a grass mat holds its place in the pres- 
ence of adverse conditions. Near the center of fig. 3z may be 
seen a white oak which was almost overthrown by the erosive 
activities, but which has been preserved through the cessation 
of erosion at this point. The gully shown near the center of 
jig. 30 is seen in closer view in fig. 7; the absence of vegetation, 
save that which has slid down from above, is very striking. 

If for any reason the lake activities at the base of the cliff 
are stopped, an opportunity is offered for the development of 
vegetation. At Glencoe the cliff erosion has been checked to 
some extent by artificial means, and one can see various phases 
of cliff life within a small area. When the erosion at the base 
of the bluff ceases, conditions become much more stable, though 
landslide action may still occur. In time the slope gradient 
becomes so low that the cliff soil is essentially stable ; when this 
time arises vegetation develops with great rapidity in spite of the 
xerophytic conditions which are still as pronounced as before. 
It is very obvious, therefore, that it is the instability of the 
eroding cliff and not its xerophytic character which accounts 
for the absence of plant life. 

The first vegetation is commonly made up of xerophytic 
herbs, both annual and perennial. Among these are the sweet 
clover (Melilotus alba), various annual weeds, various species of 
aster, especially A. laevis, Equisetum hyemale, various grasses, 
etc. oon there develops a xerophytic thicket vegetation, 
such as is shown in fig. 37. This may be called the shrub stage 
of the captured cliff, and among the dominant species are the 
juniper and cedar (Funiperus communis and ¥. Virginiana), Salix 
Slaucophylla, the osier dogwood (Cornus stolonifera), Shepherdia 
Canadensis, various sumachs (Rhus. typhina and R. glabra). The 
following tree stage is dominated by various poplars (Populus 
tremuloides, P. grandidentata, P. monilifera), the hop hornbeam 
(Ostrya Virginica), the white pine (Pinus Strobus), the red cedar 
(Funiperus Virginiana), and some of the oaks (probably Quercus 


, 


tgor] PHYSIOGRAPHIC ECOLOGY OF CHICAGO 167 


rubra and Q. coccinea tinctoria). Fig. 32 shows a tree-clad cliff’ 
in which most of the above trees are to be found. 

Whether a mesophytic forest would develop on a lake bluff 
is something of a question. It seems likely that semi-xero- 
phytic trees will dominate there for a long time to come on 


Fic. 31.—Sea cliff at Glencoe, at a place where lake erosion has ceased. Shrubs 
(largely celiat and willows) prominent as well as herbs. Absence of lake erosion 
also indicated by the e gentle slope, as compared with fg. 70. The leaning oak at the 
top bears wie to former erosive forces 


account of the xerophytic atmospheric conditions. Particularly 
at the top of the bluff do the conditions remain severe, by 
reason of the great exposure and also the dryness of the soil. 
If the lake should recede for some distance, a mesophytic forest 
could certainly develop on the bluff before it is reduced to any- 
thing like the common level. This is shown onthe ancient lake 
bluff at Beverly hills. Here there is an old cliff about forty 


168 BOTANICAL GAZETTE {MARCH 


feet above the country level, representing a lake bluff of the 
Glenwood stage of Lake Chicago.* This bluff has long had a 
mesophytic forest on its slopes, and yet it will be many centuries 
before the erosive forces remove all traces of this ancient sea 


FIG, 32.—Sea cliff at Glencoe, where lake erosion has been absent for a long 
period. Xerophytic trees and shrubs, especially conifers, dominate, e. g., white pine, 
red cedar, juniper. 


cliff. A still more striking case is to be seen north of Wauke- 
gan, where an ancient lake bluff, higher than that at Beverly 
hills and only a mile back of the present lake shore is tenanted 
by a high grade type of mesophytic forest. 

It will be instructive to make a few comparisons between lake 
bluffs and other plant societies. Closest to the lake bluff ina 


a and ALDEN: The geography of Chicago and its environs. Chicago, 


1901 } PHYSIOGRAPHIC ECOLOGY OF CHICAGO 169 


physiographic sense is the river bluff. When a stream has 
banks of clay the conditions seem decidedly similar and yet the 
flora is not the same. A comparison of the lake bluffs at 
Glencoe with the bluffs along Thorn creek shows that some 
species are common, notoriously Ostrya, Rhus, Quercus, Popu- 
lus. Yet the differences are still more striking, for the bluffs 
along Thorn creek do not show Sahx glaucophylla nor Shep- 
herdia; most striking of all, however, is the entire absence of 
conifers. When we compare the lake bluffs with the rock 
bluffs of the Illinois river we find that the resemblances are 
greater than the differences, since the river bluffs have conifers, 
though even here some of the lake bluff forms are absent. 
When, however, we compare the Glencoe bluffs with the dunes, 
we find that all of the dominant shrubs and trees of the bluff are 
found also on the dunes ; not only this, the dominant bluff forms 
are dominant on the dunes also. 

The facts of the preceding paragraph are pregnant with sig- 
nificance. One obvious corollary is that given similar soils but 
dissimilar conditions of atmospheric exposure, as at Glencoe and 
Thorn creek, the vegetation is unlike. Another and more strik- 
ing corollary is that given the most dissimilar soils possible, viz., 
the Glencoe clay and the dune sand, we still have similar vege- 
tation, because the atmospheric conditions are the same in the 
two cases. The evidence of the Illinois river bluffs is less clear; 
they are more xerophytic than the bluffs along Thorn creek, but 
whether this is chiefly due to rock as against clay or to greater 
exposure is not certain. At all events these facts show that it is 
not enough to know about chemical or physical conditions in 
the soil. We cannot divide plants into those of clay, rock, and 
sand, but must take into account that most plants have a wide 
range of life so far as soil is concerned, provided the atmos- 
pheric conditions are congenial. The chief exception to this 
statement seems to be found not in the original soils but in the 
Superimposed humus. There are many plants that require 
humus for their occurrence in nature, but it makes no difference 
whether the subsoil is rock, sand, or clay, provided alone that 


170 BOTANICAL GAZETTE | MARCH 


the humus is present in sufficient quantity. It is by reason of 
this last fact that the mesophytic forest can appear in all con- 
ditions in this climate, since the meeeeny tic forest is associated 
to a high degree with humus. 


2. THE BEACH-DUNE-SANDHILL SERIES. 


A. The beach.—The author has previously discussed in con- 
siderable detail the dynamics of the dune societies,?5 and it will 
not be necessary to do more here than to summarize the chief 
conclusions, and add a few new data. Before long it is expected 
that a paper will appear giving the changes that have taken 
place since the first observations were made in 1896. 

The beach in the Chicago area is xerophytic throughout. 
There is nothing analogous to the salt marshes of the Atlantic 
coast, nor to the hydrophytic shores farther north along Lake 
Michigan. The lower portion of the beach is exposed to alter-’ 
nate washing by the waves and desiccation in the sun, and is 
devoid of life. The middle beach, which is washed by winter 
waves, though not by those of summer, has in consequence a 
vegetation of xerophytic annuals, the most prominent of which 
is Cakile Americana. The upper beach is beyond present wave 
action, and is tenanted by biennials and perennials in addition to 
the annuals. /ig. 33 shows a beach of this type, the lower 
beach being smooth and even, the middle beach covered with 
débris, while the upper beach has a scattered perennial vegetation. 

The beach at the base of cliffs shows similar subdivisions, 
though the zones are much narrower as a rule. The vegeta- 
tion, too, is much the same, though some forms, as Strophostyles, 
have not been seen as yet on the beaches of the dune district. 
At the foot of cliffs there often occur alluvial fans of sand, 
which have been deposited by the torrents during and following 
rain storms, These fans have a comparatively rich vegetation 
and species sometimes occur here that are not found elsewhere 
on the beach. 

75 COWLES, H. C.: The ecological relations of the vegetation on the sand dunes 
of Lake Michigan. Bor. Gaz. 27: 95-117, 167-202, 281-308, 361-391. 1899. 


Igor] PHYSIOGRAPHIC ECOLOGY OF CHICAGO 171 


B. The embryonic or stationary beach dunes.—Wherever plants 
occur on a beach that is swept by sand-laden winds, deposition 
of sand must take place, since the plants offer obstacles to the 
progress of the wind. If. these plants are extreme xerophytes 
and are able to endure covering or uncovering without injury, 
they may cause the formation of beach dunes. Among the 


Fic. 33.— Beach at Dune park, showing the smooth and naked lower beach, the 
ene beach with its line ot débris, the upper beach with scattered shrubs, and the 
dune 


dune-forming plants of this type are Ammophila arundinacea, 
Salix glaucophylla and S. adenophylla, Prunus pumila, and Populus 
monilifera. The shapes of these beach dunes vary with the 
characteristics of these dune-forming plants. Ammophila dunes 
are extensive but low, because of strong horizontal rhizome prop- 
agation. Prunus and Populus dunes are smaller but higher, 
because of the relative lack of horizontal propagation and the 
presence of great vertical growth capacity. Dunes are formed 
more slowly in protected places, and here the dune-forming 
species may be plants that are ill adapted to the severest beach 
conditions, such as the creeping juniper. 


172 BOTANICAL GAZETTE [MARCH 


C. The active or wandering dunes. The dune complex.—The 
stationary embryonic dunes on the beach begin to wander as 
soon as the conditions become too severe for the dune-forming 
plants. The first result of this change is seen in the reshaping 
of the dune to correspond with the contour of a purely wind- 
made form. The rapidity of this process is largely determined by 
the success or failure of the dune-formers as dune-holders. The 
best dune-holders are Calamagrostis, Ammophila, and Prunus. 

There are all gradations between a simple moving dune and 
a moving landscape; the latter may be called a dune-complex, 
The complex is a restless maze, advancing as a whole in one 
direction, but with individual portions advancing in all directions. 
It shows all stages of dune development and is forever changing. 
The windward slopes are gentle and are furrowed by the wind, 
as it sweeps along; the lee slopes are much steeper. The only 
plant that flourishes everywhere on the complex is the succulent 
annual, Corispermum hyssopifolium, although Populus monilifera is 
frequent. The scanty flora is not due to the lack of water in the 
soil, but to the instability of the soil and to the xerophytic air. 

The influence of an encroaching dune upon a preexisting flora 
varies with the rate of advance, the height of the dune above the 
country on which it encroaches, and the nature of the vegetation. 
The burial of forests is a common phenomenon. The dominant 
forest trees in the path of advancing dunes are Pinus Banksiana 
and Quercus coccinea tinctoria. These trees are destroyed long 
before they are completely buried. The dead trees may be 
uncovered later, as the dune passes on beyond. 

In the Dune park region there are a number of swamps upon 
which dunes are advancing. While most of the vegetation is 
destroyed at once, Salix glaucophylla, S. adenophylla, and Cornus 
stolonifera are able to adapt themselves to the new conditions, by 
elongating their stems and sending out roots from the buried 
portions. Thus hydrophytic shrubs are better able to meet the 
dune’s advance successfully than any other plants. The water 
relations of these plants, however, are not rapidly altered in 
the new conditions. It may be, too, that these shrubs have 


190r | PHYSIOGRAPHIC ECOLOGY OF CHICAGO 173 


adapted themselves to an essentially xerophytic life through 
living in undrained swamps. Again it may be true that inhabi- 
tants of undrained swamps are better able to withstand a partial 
burial than are other plants. 

Vegetation appears to be unable to capture a rapidly moving 
dune. While many plants can grow even on rapidly advancing 
slopes, they do not succeed in stopping the dune. The move- 
ment of a dune is checked chiefly by a decrease in the available 
wind energy, due to increasing distance from the lake or to barri- 
ers. A slowly advancing slope is soon captured by plants, because 
they have a power of vertical growth greater than the vertical 
component of advance. Vegetation commonly gets its first foot- 
hold at the base of lee slopes about the outer margin of the com- 
plex, because of soil moisture and protection from the wind. 
The plants tend to creep up the slopes by vegetative propaga- 
tion. Antecedent and subsequent vegetation work together 
toward the common end. Where there is no antecedent vegeta- 
tion, Ammophila and other herbs first appear, and then a dense 
shrub growth of Cornus, Salix, Vitis cordifolia, and Prunus Virgint- 
ana, Capture may also begin within the complex, especially in 
protected depressions, where Saiz longifolia is often abundant. 

D. The established dunes—No order of succession in this 
entire region is so hard to decipher as is that of the estab- 
lished dunes. There are at least three types of these dunes 
so far as the vegetation is concerned, and it is not yet possible 
to figure out their relationships. The continuation of the con- 
ditions as outlined in the preceding paragraph results in a forest 
society on the lee slope, in which is found the basswood, together 
with a most remarkable collection of mesophytic trees, shrubs, 
and climbers, which have developed xerophytic structures. These 
dunes are evidently but recently established, as is shown by the 
absence of a vegetation carpet; furthermore the slopes are 
almost always steep. 

Again, there are forest societies in which the pines dominate, 
either Pinus Banksiana or P. Strobus. These arise from a heath, 
composed in the main of Arctostaphylos and Juniperus. The 


174 BOTANICAL GAZETTE [MARCH 


heath appears to originate on fossil beaches or on secondary 
embryonic dunes or other places where the danger of burial is not 
great. It will be noted that both the heath and the pine forest 
are dominated by evergreens. These societies commonly occur 
near the lake or on lakeward slopes, which are northern slopes 
as well. On these coniferous dune slopes there is to be found 
another notable collection of northern plants, resembling eco- 
logically the peat bog plants already mentioned. Heaths and 
coniferous forests also occur on sterile barrens and in depressions 
where the conditions are unfavorable for deciduous forests. A 
slight change in the physical conditions may bring about the 
rejuvenation of the coniferous dunes, because of their exposed 
situation. This rejuvenation commonly begins by the formation 
of a wind sweep, and the vegetation on either hand is forced to 
succumb to sand-blast action and gravity. 

A third type of established dune is that in which the oaks 
predominate, and especially Quercus coccinea tinctoria. The oak 
dunes are more common inland and on southern slopes. Prob- 
ably the oaks follow the pines, but the evidence on which this 
is based is not voluminous. The pines certainly have a wider 
range of habitat than the oaks, occurring in wetter and in drier 
soil and also in more exposed situations. The mutual relations 
of the pines and oaks are certainly interesting and deserve some 
very careful study. Pine forests prevail on the north or lake- 
ward slopes and oak forests on the south or inland slopes. 
With the pines are other northern evergreen forms, such as 
Arctostaphylos, while with the oaks are Opuntia, Euphorbia, 
and other more southern types. The density of the vegetation 
on the north side is also in contrast with the sparser and more 
open vegetation of the south side. The cause for this radical 
difference on the two slopes is doubtless complex, but it is 
obvious that the north slope has greater moisture, shade, and 
cold, and probably more wind. Which of these is the more 
important is not certain, but the presence of the northern spe- 
cies seems in favor of cold or wind as the chief factor. 

There are a number of interesting sand hills and ridges at 


Igor] PHYVSIOGRAPHIC ECOLOGY OF CHICAGO IS 


some distance from the lake. Some of these are fifteen miles 
from the present lake shore, while others are found at various 
intervals nearer and nearer the lake. It has been found that 
these can be grouped for the most part into three series, repre- 
senting three beach lines of Lake Chicago, as the glacial exten- 
sion of Lake Michigan has been called. The upper and oldest 


a ar 


Fic. 34.— Portion of an ancient beach line (Calumet beach) at Summit, showing 
the characteristic oak vegetation, in this case chiefly bur oaks (Quercus macrocarpa). 


of these ridges has been termed the Glenwood beach, the inter- 
mediate ridge the Calumet beach, and the lower and younger 
ridge the Tolleston beach. The geographic relations of these 
beaches is well discussed by Leverett and also by Salisbury 
and Alden,?7 and nothing need be said here except as to the 
vegetation. In general these ridges and hills have a xerophytic 
forest flora, dominated by the bur, black, and white oaks 
( Quercus macrocarpa, Q. coccinea tinctoria, Q. alba). The propor- 
tions between these trees varies strikingly, though the bur or 


6 OP. cit. 55-85. °7 Op. cit. 31-51. 


176 BOTANICAL GAZETTE [MARCH 


black oak is usually the chief character tree. No satisfactory 
reason can yet be given for these variations, though the bur 
oak appears to be more abundant on the lower and less drained 
ridges, while the black oak is more abundant on the higher 
ridges. The shrub undergrowth is commonly sparse, and the 
most frequent members of this stratum are the hazel (Corylus 


35.— Portion of an ancient beach (Glenwood beach) near Thornton. The 
trees here are chiefly black oaks (Quercus coccinea tinctoria) ; the beach is higher, and 
the trees more luxuriant than usual. 


Americana), Rosa, the New Jersey tea (Ceanothus Americanus ), 
Salix humilis, the low blueberry ( Vaccinium Pennsylvanicum) , 
and the huckleberry (Gaylussacia resinosa). Among the com- 
moner herbs are Silene stellata, Antennaria plantagintfolia, Heu- 
chera hispida, Rumex Acetosella, Carex Pennsylvanica, Potentilla 
argentea, Poa compressa, Pteris aquilina, Ceratodon purpureus. In 
open places there are often almost pure growths of Poa or 
Potentilla. Figs. 34 and 35 show portions of these ancient 
beaches in which the oaks dominate; jig. 34 shows, perhaps, the 
more common condition, 7. ¢., a rather low beach with a sparse 
tree growth. 


1901] PHYSIOGRAPHIC ECOLOGY OF CHICAGO 177 


The future of the vegetation on the established dunes and 
beaches is somewhat problematical. From analogy with other 
plant societies in this region, and from established dunes in 
Michigan, we should expect a mesophytic forest, probably of 
the white oak-red oak-hickory type at first and then followed by 
a beech-maple forest. There are evidences that some such 
changes are now taking place. On many of the oak dunes, 
especially where protected from exposure, there is already a 
considerable accumulation of humus. Herbaceous ravine meso- 
phytes like Hepatica, Arisaema, and Trillium are already 
present, and with them mesophytic shrubs and trees, including 
the sugar maple itself, though the beech has not been found on 
the dunes of our area, as it has in Michigan. One might expect 
that the flora of the older Glenwood beach would have advanced 
more toward the mesophytic stage than has the flora of the 
younger Tolleston beach. Such, indeed, seems to be the case, 
especially at Glenwood, where the white oaks are more numerous, 
and the black oaks. much larger and more luxuriant. The humus 
is richer and most things look as if the age of this beach were 
notably greater than that of the Calumet or Tolleston beaches. 
This subject, however, needs much further investigation. In 
any event, one character of the sand hill stands out in bold 
relief, viz., its great resistance to physiographic change. Not 
only is its erosion slower than that of the clay hill, but the 
advance of its vegetation is vastly slower at all points along the 
line. The slowness of humus accumulation accounts for this, 


perhaps, more than all else. 


Ill. Summary and conclusion. 


In the present paper the author has endeavored to show the 
need for a classification of plant societies which shall form a 
logical and connected whole. Warming’s classification, based on 
the water content of the soil, is doubtless the best possible classi- 
fication, if but one factor is used. Graebner’s classification, based 
on soil characteristics, includes the advantages of Warming’s 
scheme, and adds desirable new features. 


178 “BOTANICAL GAZETTE [MARCH 


The physiographic theory here presented is the result of 
several years of field study devoted chiefly to testing the current 
theories and to developing new ones. The classification is based 
on the fundamental notion that a true theory must be “genetic 
and dynamic; the plant societies must be grouped according to 
origins and relationships, and the idea of constant,change must 
be strongly emphasized. 

The laws that govern changes in plant societies are mainly 
physiographic ; whether we have broad flood plains, xerophytic 
hills, or undrained swamps depends on the past and present of 
the ever-changing topography. Nor is topographic change 
haphazard. Modern physiography has made a logical classifica- 
tion of dynamic surface forces, and has found a progressive 
tendency toward a definite end. Denudation of the uplands and 
deposition in the lowlands results in an ultimate planation, known 
as the base level. Wherever hills are being eroded, or lakes 
filled, or coastal plains enlarged, it is obvious that there must 
be changing plant societies, in other words, a definite order of 
succession of plant groups. These changes, too, are cumulative ; 
a topographic form will have plants that are relicts of an older 
stage, as well as those that are typical of the new conditions, 
showing that the supplanting of one plant society by another is 
slow and gradual. The full effect of a given environment may 
not be felt till that environment has gone. 

Using ecological terms in place of those of physiography, 
soil conditions tend to become more mesophytic as the base level 
develops. A young topography is rich in xerophytic hills and 
in hydrophytic lakes and swamps. There may be local retro- 
gressions toward xerophytic or even hydrophytic plant societies, 
forming eddies, as it were, but the great movement is ever pro- 
gressive and toward the mesophytic condition. Though instances 
of vast planation are found in geological history, the ultimate 
mesophytic base level is seldom reached, since crustal move- 
ments interfere with physiographic processes. So far as plants 
are concerned, however, a physiographic terminology may still 
be used, since all possible crustal changes are either toward or 


ne _—— — 


he ae 


1901] PHYSIOGRAPHIC ECOLOGY OF CHICAGO 179 


away from the mesophytic, 7. ¢., progressive or retrogressive. 
Again, climatic changes doubtless occur; even here we may use 
the general terminology, since the new conditions either favor 
or retard the general mesophytic development. This leads to 
the general view that the climax type differs with the climate. 
While the general series of physiographic changes is much the 
same everywhere, the corresponding plant societies are vastly 
different. Ina desert climate most of the societies, including 
the climax type itself, are xerophytic. Finally, there is at least 
one point where physiographic-and ecological classifications must 
diverge. Changes. in vegetation often take place where the 
topography remains the same; in other words, a cycle of vege- 
tation may be shorter than a cycle of erosion. The following 
application of these principles applies only to the Chicago region. 

The typical erosion series is based on the life history of 
rivers, and this series is the most instructive ecologically. An - 
embryonic clay ravine is essentially a little desert, though this 
character is due more to the instability of the soil than to the 
ordinary xerophytic factors. Soon landslide action becomes 
much reduced, and a xerophytic flora may appear, though ina 
remarkably short time a rich mesophytic forest is developed. 
This forest is not permanent, but may be regarded as a tempo- 
rary climax. Rock ravines, whether of limestone or sandstone, 
commonly have more vertical slopes and drip with moisture, 
favoring the growth of extreme shade plants. The stages in 
limestone and sandstone habitats are essentially the same in 
spite of great physical and chemical differences in the rock. 

As a clay ravine widens, the exposure increases; xerophytic 
herbs appear at the top of the slope, and later farther down 
toward the stream bed. Xerophytic shrubs and finally xerophytic 
trees make their appearance, notoriously Ostrya Virginica. In 
the early stages of these xerophytic bluffs trees are often found 
that look back to the ravine for their origin, while under them 
are xerophytic herbs that are better suited to the new conditions. 
As the slopes of a mesophytic rock ravine pass to a xerophytic 
rock bluff, changes in the vegetation are most pronounced. A 


180 BOTANICAL GAZETTE [MARCH 


slope less xerophytic than that of clay becomes more xerophytic 
as it becomes a river bluff, and conifers are found as well as 
deciduous xerophytes. As the bluff slopes become more gentle 
through erosive action, a mesophytic flora may gradually replace 
the xerophytes. 

Before the growing valley possesses a permanent stream 
there may be developed in the torrent bed a vegetation of 
amphibious shade plants, and when the water becomes more per- 
manent one may find algae and other hydrophytes. Spring 
brooks are infrequent, but they have a characteristic vegetation, 
due doubtless to the presence of more light and water. The 
development of a flood plain vegetation is well shown on river 
islands. First a sand bar develops, then an annual flora, and 
later a perennial vegetation in which Salix dominates. The river 
constantly erodes above and deposits below, hence the islands 
migrate down the stream, showing the oldest plant societies at 
the upper end. Depositing streams gradually develop a flood 
plain which shows an interesting succession of societies. Beyond 
the true hydrophytes there is commonly seen a Salix zone, then 
a zone of Populus and other trees on the older flood plain, and 
finally there develops a luxuriant mesophytic flood-plain forest 
which as a whole is permanent, though local retrogressions may 
occur. In some of these flood-plain forests there are found 
interesting southern types of trees. Occasionally meadows 
occur on flood plains in place of forests. Retrogressive proc- 
esses are active on flood plains, such as terrace formation, which 
is due to further erosion; terrace development tends to favor 
xerophytes. New channels are also cut off, leaving portions of 
the old river as oxbow lakes; here hydrophytes of undrained 
swamps come in, and one often sees trees of the old river margin 
together with shrubs and herbs of the undrained swamps. These 
latter phases, however, are ephemeral and the mesophytic flood 
plain as a whole increases constantly in area. 

The vegetation of undrained areas has a remarkably xero- 
phytic stamp; this is possibly due to the unfavorable oppor- 
tunity for root activity in undrained soils. In any event, these 


| 
| 
| 
ig 
| 


ee 


1901] PHYSIOGRAPHIC ECOLOGY OF CHICAGO 181 


areas are features of a young typography and they are soon filled 
up by accumulating peat. Algae and other hydrophytes charac- 
terize the lake or pond stages, and largely by their partial decay 
the water becomes shallow enough to support the vegetation of a 
marginal swamp, particularly rushes and sedges. After these forms 
there appear the most remarkable plant societies of the entire 
series, characterized chiefly by Cassandra and other ericads with 
xerophytic structures. Following the shrub stage there is the 
tree stage in which the tamarack often dominates, though pines 
appear later and ultimately mesophytes. In these swamps one 
finds the most perfect examples of the regular succession of 
plant societies, and hence of zonal arrangement. These peat bog 
societies contain a most striking collection of northern plants. 
There are various diverging types of undrained swamps, some 
with shallow soil and a vegetation without extreme xerophytic 
structures, others in which an imperfect drainage may account 
for the facts observed. The marginal vegetation is the same in 
all cases and is remarkably characteristic; this flora is some- 
times seen after the lake and swamp floras have gone. Half- 
drained areas are characterized by luxuriance of the vegetation 
in the lake stage. The lake is followed in order by the bulrush, 
sedge, and grass stages, the latter being denominated the prairie 
stage. Whether this latter type passes into the forest is not 
certain; in any event, this mesophytic stage is not final, for the 
region must subsequently pass through the stages of the river 
series. 

Uplands, as well as swamps, have an interesting history 
before they are attacked by stream erosion. Limestone hills 
and outcrops show first a lichen vegetation, followed by mosses 
and crevice herbs and later by shrubs and trees. The stages 
on clay hills pass far more rapidly, indeed early stages are 
hard to detect, and one almost uniformly finds a mesophytic 
forest of oaks and hickories in these habitats. Where the 
forest conditions are disturbed, there is a rapid return through a 
series of herb and shrub stages to the same tree types. It is 
somewhat probable that the oaks will be followed by sugar 


182 BOTANICAL GAZETTE [MARCH 


maples and beeches, and this change is now occurring in some 
places. The mesophytic stage here also is not permanent, 
though it may endure for a long period. 

Clay bluffs along an eroding coast are subject to considera- 
ble change involving the rapid destruction of upland vegetations. 
The changes, too, are so rapid that practically no vegetation can 
develop on the bluff slopes. If the erosive activity of the lake 
ceases, there soon appears a vegetation of xerophytic herbs, fol- 
lowed by xerophytic shrubs and trees in which conifers play a 
large part. There is a notable resemblance between the flora of 
the clay bluff along the lake shores and the flora of the sand 
dunes, and also the flora of rock bluffs along rivers, while the 
resemblance is less close to the flora of clay bluffs along rivers, 
showing that soil conditions may often be less important than 
conditions of aerial exposure. 

The dune vegetation, presented in dean elsewhere, is sum- 
marized in this present paper. Nothing need be said in the way 
of further summary except to remark that dunes, like all other 
topographic forms in our climate, may ultimately develop a 
mesophytic forest, though the stages are far slower than in most 
of the other series. 


THE UNIVERSITY OF CHICAGO. 


SOME RECENT PUBLICATIONS AND THE NOMEN- 
CLATORIAL PRINCIPLES THEY REPRESENT. 
M. L. FERNALD. 

THE second edition of Mr. Heller’s Catalogue* presents, as 
a piece of presswork, a great improvement over its predecessor. 
For thus materially improving the dress of his work the author 
is certainly to be commended. The book also contains a very 
large increase of species over the first edition; and for bringing 
together hundreds of recently published names, where they can 
be readily consulted, the compiler should have the gratitude of 
students of systematic botany. In a work of this sort, neces- 
sarily accomplished largely by compilation, monographic treat- 
ment of recent and doubtful species can not and should not be 
expected; but there can be no question that the compiler of a 
check list or catalogue owes to the public the product of the 
best light he has upon the species with which he deals. Mr. 
Heller’s new Catalogue, especially, representing the so-called 
reform tendencies in American botany, should be judged pri- 
marily by the degree of adherence to or divergence from the 
principles which he has taken upon himself to exploit. This 
second edition, too, should be judged by the degree of readi- 
ness shown by its author to correct such obvious errors and 
inconsistencies in his preceding work as have been definitely 
called to his attention in print. 

It is a question which is the point of greater significance to 
Systematic botany—the hopeless tangle of nomenclatorial prin- 
ciples here exhibited, or the tendency, by no means new, to 
break through the traditional though necessarily vague barriers 
Separating the minor categories to which plant-variations may 
be assigned, namely, the species, variety, and form. That the 
author of this Catalogue, and numerous other American botanists, 

*Catalogue of North American plants north of Mexico, exclusive of the lower 
cryptogams. By A.A. HELLER. Second edition. Issued November 10, 1900. 
1901] 183 


184 BOTANICAL GAZETTE [MARCH 


like Jordan and others in Europe, have essentially abandoned 
these finer discriminations in classification, has been made per- 
fectly evident in their past work, and now we are informed that 
‘there is also a growing tendency to discard the use of varietal 
names, and to call all plants species which have characters 
enough to justify the use of a distinctive name.”* To the 
critical student, who is familiar with growing plants and the 
causes which so often control their variations, such action as 
this cannot appeal; and, though the author of the Catalogue is 
not alone in his standpoint, there are still many students whose 
conceptions of plants and their relationships can be expressed 
only by the retention of categories which are subordinate in 
rank to the species, 

However, even if, by putting essentially all variations within 
the genus upon a common level, the author chooses to obscure 
the minor degrees of relationship in plants, there should be no 
question of personal choice or opinion in judging the method so 
often adopted by him in order to increase the number of so-called 
species. In the introduction to the new Catalogue (as well as in 
the original edition) many new combinations of names are made; 
but, finding that space would not permit the publication there of 
all the changes he desired to make, the author has ventured a 
new private journal,3 the first issue of which is occupied by an 
appendix to the nomenclatorial changes begun in the Catalogue. 
It seems that the author has, or did have on November 10, 1900, 
some conception of the unsatisfactory methods he was employ- 
ing, for in apologizing for so proceeding he says: ‘‘The bare 
citation without discussion in most cases is undesirable, but lack 
of time forbids a more extended treatment of the different species | 
under consideration | italics ours}.”+ Does anyone suppose that 
by careful botanists such an apology can be accepted as a pledge 
of sincere desire to advance botanical science; or can it be that 

SHELLER, 2:¢, 3: 


3Muhlenbergia, a Journal of Botany. Edited and Published by A. A. HELLER, 
Lancaster, Pa. 


4+ FPLLERS 7. 63 22%, 


Igor] NOMENCLATORIAL PRINCIPLES 185 


the author conceives that his #me, the ‘‘lack”’ of which alone per- 
mits him to launch so many waconsidered species, is of far greater 
importance than that of the scores of other botanists who must now 
spend weary hours trying to unravel the snarls he has produced ? 
An example or two may make our point more clear. The names 
Prenanthes Serpentaria and Nabalus Serpentarius have been essen- 
tially interchangeable in American floras,’ and the name in either 
case has been made to cover until recently two very different 
species. Inthe //ustrated Flora, however, Dr. Britton has revived 
Cassini’s Nadbalus trifoliolatus for a well-marked northern plant, and 
has left the name WV. Serpentarius to cover (as it should) the thick- 
leaved species of more southern range with the ‘‘involucre more or 
less bristly-hispid.” Torrey and Gray described Nadalus Fraseri, 
var. darbatus, with the ‘“involucre (12—15-flowered) hirsute when 
young with long purplish hairs,”’ and in the Synoptical Flora, Dr. 
Gray, writing at a time when the name Prenanthes Serpentaria 
covered the northern plant with usually glabrous involucre, pub- 
lished P. Serpentaria, var. barbata, with the remark that ‘occa- 
sionally a few of these setose hairs are found on the involucre 
of ordinary P. Serpentaria, and in this variety [darbata] some 
heads are almost destitute of them.’’ Now the original Torrey 
and Gray specimen of this variety is in no way different from 
the species, Mabalus Serpentarius, as correctly interpreted by Dr. 
Britton. Nevertheless, we have in Muhlenbergia (1:8) the 
new combination Mabulus barbatus (T. & G.) Heller, although in 
the Catalogue both N. Serpentarius and J. ¢rifoliolatus are listed. 
Again, Zlex verticillata, forma chrysocarpa, noted by Dr. Robin- 
son in Rhodora (2:106), appears in the new Catalogue as 
“(var.] chrysocarpa Robinson.” The original specimen in the 
Gray Herbarium has never been borrowed by the author of the 
Catalogue, and it is perhaps elevated by him to varietal rank 
through carelessness; but now that it is listed as a variety it will 
be interesting to see how soon it will be erected to a species by 
one who believes in calling ‘‘all plants species which have 

SIn this paper these names may be thus accepted without discussion as to 
their status. 


186 BOTANICAL GAZETTE | MARCH 


characters enough to justify the use of a distinctive name,” and 
who, like him, knows the plant only from the descriptive phrase 
“fruit bright yellow.” These two cases are sufficient to show 
the character of work which can be done by one who believes 
in making all the new combinations. possible, when he is handi- 
capped by “lack of time’’ to consider his work, and who 
apparently holds it more important to launch a mass of ill-con- 
sidered (and often to him unknown) species than to publish 
only the results of critical and scholarly consideration. 

If in thus launching so many species (and occasional varie- 
ties) of which he can have little or no personal knowledge, the 
author were producing combinations consistent with the names 
in the remainder of his book, his reason would be obvious and 
to some extent justifiable. But only a slight examination of the 
names taken up is sufficient to show that he has had little con- 
ception of any clearly defined principle to govern his selection 
of names. Professedly the names in his work, like those in the 
Botanical Club Check List and in Britton and Brown’s //lustrated 
Flora, are based upon the principle of strict priority; but the 
result, as shown here perhaps even more than in those works, 
gives us little assurance that the publications on such a basis are 
bringing us the uniformity which has been so loudly proclaimed 
and which every one would so gladly welcome. 

In the first group of plants listed, for instance, the Pterido- 
phyta, the names essentially as defined by Professor Underwood® 
are taken up. Professor Underwood is one of the few authors 
among the radical botanists who has squarely faced the strict 
priority question, and in his selection of generic types he has 
attempted to follow the logic of his course to the bitter end. 
Thus, as the type of the genus he takes the first species described 
under the generic name, so long as the same plant does not 
belong to some previously defined genus. In such a case he 
logically takes for the generic type the first species which is 
clear from all previous genera. However much one may differ 
from him as to the expediency of such a course, it is indeed a 

° Our Native Ferns and their Allies. Ed. 6. 1900. 


Igor] NOMENCLATORIAL PRINCIPLES 187 


satisfaction to know that at least one follower of the Rochester 
Code is ready to show us the actual task and the enormous 
upsetting of names consequent upon a conscientious and logical 
working out of the principle of strict priority. 

In Mr. Heller’s Catalogue many of the names accepted are 
not those which can be used consistently by authors who are 
committed to the Rochester Code. When that code was pro- 
posed it was professedly with the purpose of establishing 
uniformity in our nomenclature. As an outgrowth of its adop- 
tion by some American botanists the Botanical Club Check List 
was issued, a list which aimed to give us the names which our 
northeastern plants must henceforth bear according to the 
rulings of strict priority principles. That publication gave us 
the first tangible result upon which to base our estimate of the 
workings of the code; and though by some thoughtful and con- 
servative students the book and the principles represented by it 
were carefully discussed, by other botanists the publication was 
hailed as “the sign that the day of ‘authority’ as such is ended, 
and the day of ‘law’ has begun,”? and we were informed that 
“even the most obscure botanist is nowadays entitled to know 
why an old plant comes out under a new name... . and that 
their [the compilers of the Check List] work is plain work, the 
plain and straightforward statement of facts.” ® 

It is pertinent, then, for ‘the most obscure botanist” to ask 
about some of the names now (at least at the time of this 


writing) in vogue among those who champion the Rochester 


Code, and we may be permitted to inquire of those who have 
been instrumental in bringing about the present ‘uniformity ” 
how they account for a few of the names in their pages. Ref- 
erence has already been made to Professor Underwood's treat- 
ment of the ferns partially adopted by Mr. Heller in his Cata- 
dogue. In Britton and Brown’s Jilustrated Flora, published in 
1896, 59 species of true ferns are recognized, and the names, we 
are told, are those authorized by the Rochester Code. But 
in Professor Underwood’s latest treatment more than 25 per 


7,8 Bessey, C. E.: Am. Nat. 29: 350. 


188 BOTANICAL GAZETTE [MARCH 


cent? of those very species appear under different names — still the 
names authorized by the Rochester Code. For the benefit of 
those not familiar with these works, but who have an interest in 
seeing the working of the same rule in the hands of one of its 
strong advocates, the fifteen northeastern species which have 


recently appeared under new names are here enumerated : 


Names in Britton and Brown’s 


Illustrated Flora (1896), based upon the 
Rochester Code. 


Onoclea Struthiopteris (L.) Hoffm. 
Dicksonia punctilobula (Michx.) A. 


Gray. 
Cystopteris bulbifera (L.) Bernh. 
Jragilis (L.) Bernh. 
- montana (Lam.) Bernh. 
Dryopteris Lonchitis (L.) Kuntze. 
: acrostichoides (Michx.) 
Kuntze. 
Braunii (Spenner) Under- 
wood, 
Phegopteris Dryopteris Robertiana 
Hoffm.) Davenp. 
Scolopendrium Scolopendrium (L.) 
Karst. 
Asplenium acrostichoides Sw. 
Pteris aquilina L. 
Pellaea Stelleri (S. G. Gmel.) Watt. 
Cheilanthes gracilis (Fée) Mett. 
Notholaena nivea dealbata (Pursh) 
Davenp. 


Names in Underwood’s 
Our Native Ferns and their Allies (1900), 
based upon the Rochester Code. 
Matteuccia struthiopteris(L.) Todaro. 
Dennstaedtia punctilobula (Michx.) 


Bernh. 
Filix bulbifera (L.) Underwood. 
Sragilis (L.) Underwood. 
‘“« montana (Lam.) Underwood. 
abi iocgeaati lonchitis (L.) Roth. 
acrostichoides (Michx.) 


chott. 
Braunii (Spenner) Law- 
son. 
Phegopteris Robertiana (Hoffm.) Un- 
erwood. 
Phyllitis scolopendrium (L.) Newm. 
Asplenium thelypteroides pee 
Pteridium aquilinum (L.) K hn. 
Cryptogramma Stellert ea Prantl. 


Chetlanthes Feet Moore. 
Notholaena dealbata (Pursh) Kunze. 


That the names recently used by Professor Underwood are 


more truly consistent with the strict priority principles than 
many names in other groups listed by Mr. Heller has been 
already emphasized. But why, we would ask, are there so many 
unexplained inconsistencies in this new Catalogue, especially 
when the attention of followers of the Rochester Code has 

° The true ferns alone are here considered, and the genus Botrychium is purposely 


omitted, since that genus has been subdivided by Professor Underwood to such an 
extent that comparative figures would have little definite significance. 


1901] NOMENCLATORIAL PRINCIPLES 189 


been called to a number of such erroneous matters? A very 
typical example of the inconsistent method (or lack of method) 
employed in the Catalogue is shown in the treatment of the 
genera Cheiranthus (Erysimum of authors), Evysimum (Sisym- 
érium of authors), and Stsymbrium (Nasturtium and Roripa’ of 
authors). Professor E. L. Greene* pointed out, in December 
'1896, the only logical and consistent course to be followed, accord- 
ing to strict priority principles, in the cases of Cheiranthus and 
Erysimum. Mr. Howell has followed his lead, and in a recent 
article the same point"? was further emphasized. There is, then, 
no reason why the authors of the second volume of the J/lus- 
trated Flora, published May 31, 1897, and of the Appendix, pub- 
lished June 20, 1898, should have been ignorant of Professor 
Greene’s logical article. But why did they ignore his conclu- 
sions and use names in a sense absolutely inconsistent with the 
*° The so-called reformers persist in writing Rorifa instead of the original form, 
Rorippa, and they say Bicucudla instead of the original form, Bikukulla, although the 
matter has been freely discussed in the past, and by this time oe ould be aware of 
the facts in the case. But, on the other hand, after using the name Aoniga, they now 
take up the original Xon7g. Ifin one case they adopt the iia spelling, why not 
in the others? Is this what they consider a consistent method, and does it appeal to 
them as “the plain and straightforward statement of facts?” We shoul 
inquire also about the name which, in the publications of the reformers, has recently 
taken the place of Mikania Willd. In the Botanical Club Check List we patie a name 
attributed to Necker and spelled Willoughbya, with the footnote remark “ 
HE 


we are informed that the plant was “ an ro 
1635-1672, English naturalist, b e spelled by Necker as above 
éaea}.” This spelling is thee eihly followed in Mr. a en 
Otto Kuntze enumerates in his Revisio Gener: lantarum some “ incorrect ways of 
writing the name” as follows: “ Wilke rie, Wig, Willughbeia, atlas 

beja, Willugbeia, Willughbeja, Villughbeia.” r Necker, himself, if he were living, 
would indeed be dazed, particularly as his name was unlike sh of those definitely 
asserted by Dr. Britton to be correct, and since, on the contrary, the true and original 
form Willugbaeya, is the first form enumerated by Kuntze as “ incorrect. 4 
these facts it would seem that to some botanists whose work is controlled by “law 
such divergence rate van original spelling is of slight moment. If so, will they be 
good enough to make clear why Mikania is rejected for Willugbaeya and its varia- 
tions (1790), when in 1789 eae Bice Willughbeja (compare Otto Kuntze 
above), a genus of the Afocynaceae 

* Pittonia 3: 128. 

FI, N. W. Am. 1 :38-56. 73 ROBINSON, B. L.: Bot. GAZ. 25 : 439-442. 


190 BOTANICAL GAZETTE [MARCH 


spirit and rulings of their own Rochester Code, especially when, 
as we were informed in 1895 by one of the Check List commit- 
tee, ‘the committee .~. 45.5 would still be grateful..... 
for useful suggestions on these matters, and that all communi- 
cations of this kind would receive fair hearing and sober judg- 
ment.’’** There is, furthermore, no possible reason why the 
author of the Catalogue which suggested this discussion, should 
have been, in 1900, uninformed of the publications on the sub- 
ject. In fact, perhaps unconscious of the thoroughly inconsis- 
tent course he was taking, he has followed one third of the 
suggestions made and has adopted for the conventional &ry- 
stmum of authors the name Cheziranthus; but he still clings to the 
names Stsymbrium and Roripa for genera to which they cannot 
be applied by conscientious followers of strict priority principles 
dating from 1753. 

Another point in regard to generic names pointed out in one 
of the articles cited*S is in the case of Cerastium and Stellana. 
It was there clearly shown that when the first part of the Lin- 
naean Séel/aria was transferred by the reformers to Alsine (a 
course not entirely free from question), one species was still left 
in Stellaria, namely, S. cerastioides L. This plant is treated by 
modern authors as a Cerastium, and in the Botanical Club Check 
List, the Illustrated Flora and in Mr. Heller’s new Catalogue it 
appears as C. cerastioides (L.) Britton. But in the Species Plan- 
tarum of Linnaeus Stel/aria preceded Cerastium, and therefore 
the portion of Stellaria (S. cerastioides) left when the remainder 
was transferred to Adsine should, according to the strict priority 
principle, become the type of Sved/aria, and the succeeding genus 
Cerastium should be absorbed by it. Why, then, after this mat- 
ter was clearly pointed out in June 1898, does the author of the 
Catalogue, who does not hesitate to launch a lot of new combina- 
tions based upon plants of whose status he is much less certain, 
still keep up the name Cerastium in its traditional sense ? 

The familiar vine known to most of us as Wisteria is listed in the 

4 COVILLE, F. V.: Bot. Gaz. 20: 164. 

*5 ROBINSON, B. L.: Bor, Gaz. 25 : 444, 445. 


ON ener SSS Nuesnesnstmmemmerenereerere———— . 


1901] NOMENCLATORIAL PRINCIPLES Igl 


Catalogue as Kraunhia frutescens (L.) Greene (though that name 
was first published by Rafinesque in 1808); but the recently 
described Apios Priceana Robinson is given without change of 
name. The author of the Catalogue must admit that he knew of 
the publication of the latter plant, else how could he include it 
in his Catalogue. But will he inform us how it happens that he 
has ignored the facts ‘presented in the original discussion,” of 
that species? Was it not shown as clearly as could be desired 
by anyone that the names Agios Moench (1794) and Kraunhia 
Rafinesque (1808) were both antedated by Bradlea Adanson 
(1763)—a name applied to two Linnaean species of Glycine, G. 
Apios (Apios tuberosa Moench), and G. frutescens ( Wisteria frutes- 
cens Poir.) now referred by the reformers to Kraunhia? And 
was it not made clear that by those who would follow the Roch- 
ester Code the name Brad/ea must be taken up for Aptos? How 
does the author of the Catalogue, who lists Apios Priceana, explain 
his failure to stand by the principles he claims to follow ? 
There are many other generic names accepted by the reform 
botanists and now adopted in this Catalogue, which, according to 
the rules to which they have committed themselves, have no 
better status than those pointed out. But the few cases already 
explained in the past and here again emphasized are sufficient 
to show him who cares to examine the original references that 
the member of the Check List Committee, who, in 1895, wrote 
that ‘all communications of this kind would receive fair hearing 
and sober judgment” could not have been speaking for all the 
members of the committee, nor indeed for many whose prolific 
writings have done more than anything else to stultify the rules 
of which they claim to be true advocates. That such absolute 
recklessness in the application of these rules is not satisfactory 
to all members of the Check List Committee is occasionally 
made apparent. Professor Underwood’s position in regard to 
fern names has been remarked; and another of the committee 


has thus expressed himself: ‘‘Why are some of us so openly at 


war with our own rules? Certainly no rule relating to the 


*©Rosinson, B. L.: Bot. Gaz. 25: 452. 


192 BOTANICAL GAZETTE [MARCH 


observance of priority has been more generally recognized and 
deferred to than this, that a genus, as to its name at least, stands 
or falls with its type species; no rule is more indispensably 
necessary; and nothing but endless change and confusion can 
come of the neglect of it.” 77 

Numerous inconsistencies as to the treatment of species as well 
as genera have been publicly pointed out; yet here, as in case of 
some other perfectly just and logical criticisms, the effort seems 
to have been wasted upon those who are bringing us ‘‘the day 
of ‘law.’” In areview*® of the first edition of the Catalogue 
attention was called to some of these specific names. Anoda 
Javaterioides Medic., for instance, as there intimated, has a Lin- 
naean synonym in Sida cristata, while Arenaria sajanensis Willd. 
is the same as the Linnaean Svellaria biflora ( Arenaria biftora (L.) 
Watson, which is the name accepted by Dr. Britton). Yet in 
spite of these very clear cases which have been emphasized in 
print, the second edition of the Catalogue follows the first in giv- 
ing Anoda lavateroides | lavaterioides| and Arenaria Sajanensis. 
By what “law” are these names reconciled with the Rochester 
Code, and why is the public criticism of their use by the so- 
called reformers so openly ignored ? 

Another point emphasized by the same reviewer, whose 
words apparently bore too much of ‘‘authority” to influence 
the author of the book criticized, was the abundance of ‘‘ perfect 
and confessed synonyms” in the Catalogue, thus swelling its 
bulk, but decreasing by inverse proportion the confidence we can 
feel in it as the product of careful work. Several cases were 
cited (Silene Cucubalus and S. vulgaris, for example); but, as we 
have now learned to expect, the same misleading and unjustified 
duplication of names occurs in the new edition. When, how- 
ever, the same species appears under different genera, as in case 
of Aster nemoralis Ait., we must confess the least bit of surprise. 
Professor Greene, in splitting the genus Aster, revived for part of 
it the Nuttallian genius Eucephalus. Among other species which 

7 GREENE, E. L.: Pittonia 3: 129. 

ROBINSON, B. L.: Am. Nat. 32: 460. 


Igor] NOMENCLATORIAL PRINCIPLES Tg3 


he proposed was Eucephalus nemoralis, based upon Aster nemo- 
ralis Ait., though in Aster he left the closely related and often 
indistinguishable A. acuminatus Michx. In the new Catalogue 
we find under Aster, A. nemoralis Ait., listed and numbered, while 
under Eucephalus we have E. nemoralis Greene, treated in the same 
handsome manner. The troublesome Aster nemoralis var. Blakei 
Porter, however, a plant which so mingles the characters of 
Lucephalus nemoralis Greene and Aster acuminatus Michx. as to 
embarrass even its own author, is wisely left with Aster nemoralis. 
Why, then, if Zucephalus nemoralis is identical with Aster nemo- 
rats, does the author of the Catalogue list the variety of the latter 
only under Aster, when the species is treated as belonging to 
both genera ? 

Many of us were brought up to speak of Adsma Plantago L. 
and Veronica Anagallis L., but during the past decade the fol- 
lowers of the Rochester Code have adopted the fad of calling 
these plants Alisma Plantago-aquatica and Veronica Anagalls- 
aquatica, The use of such names has indeed afforded an inter- 
esting diversion and has kept us constantly tingling with expect- 
ant excitement as we have waited to see what other familiar 
names would appear in new and fantastic garb; but it must be 
confessed that a careful search in the volumes of Species Plan- 
tarum, where these names are said to occur, has failed to reveal 
them. Instead this is what is found: Veronica Anagall.y and 
Alisma Plantago A. Thus it seems that Linnaeus did not write 
even Anagallis in full; and we should like to be informed on 
what authority (in the Species Plantarum) we know that A and Vv 
are both mysterious ways of writing aguatica? And if a triangle 
is said to mean aguatica why do not the reformers append that 
adjective to their Rorifa Nasturtium, for Linnaeus wrote Sisym- 
brium ib sen Siaeata’ or does the position of the triangle in relation 
to the name give it a new meaning? Here is a great oppor- 
tunity for someone to hunt up all the triangles in the Species 
Plantarum and thus give us a new lot of specific names. But, 
Seriously, we may ask why, in the new edition of the Catalogue 
{as in the old), this modern fad was followed ? 


194 BOTANICAL GAZETTE [MARCH 


The triangles, like some other symbols occasionally used by 
Linnaeus, presumably descended from earlier authors, and if 
they are taken to mean aguatica, and that adjective is written as 
a portion of the plantname, we are simply reverting to the pre- 
Linnaean method of polynomial (or at least trinomial) names, 
and the whole system of binomials is weakened. The date 1753 
has been generally accepted as the limit back of which we are 
not to go for names; and if Linnaeus himself did not use the 
name Alisma Plantago-aquatica or Veronica Anagallis-aquatica, are 
we justified in going back to some earlier author for such 
names? Right here is a very dangerous tendency in the usage 
of the reformers. If they will thus admit an occasional pre- 
Linnaean name which was not used in the first edition of the 
Species Plantarum, what assurance do they give us that their strict 
priority rule with a time-limit definitely set at 1753 may not at 
any time be made elastic enough to protect any whimsical excep- 
tion its advocates choose to set up? 

One of the members of the Check List Committee, speaking 
of the citation of the original author of a combination, has 
informed us that “it is no longer a question of credit, but a 
question of practical utility.”* Surely this is the ideal for 
botanical nomenclature which every serious student will com- 
mend; and we may well put to ourselves the question, is ‘“ prac- 
tical utility’’ in view or does it seem very near actual attainment, 
when we find the members of the committee which set out to 
give us a uniform system of names “at war with” their own 
rules? Has the “day of ‘law’” really begun when those with 
whom a great trust has been placed juggle with it as with a toy, 
now following this principle, now that, and ignoring at their own 
wills such candid criticisms of their methods as show the incon- 
sistencies in their work? Is the * day of ‘authority’ as such” 
indeed ended when, after one of their own associates on the com- 
mittee has publicly reprimanded them and has pointed out the 
only course for one who would live up to the principles he has 
espoused, the supporters of the Rochester Code continue to 

*9 WARD, L. F.: Bull. Torr. Bot. Club 22: 325. 


1901 | NOMENCLATORIAL PRINCIPLES 195 


employ names inconsistent with the principles there empha- 
sized? 

A member of the committee already quoted has said, in 
defending the principles of the Rochester Code (principles which 
as abstract principles need little defense), ‘‘if matters are to be 
left to the individual judgment of publishing botanists, there will 
be no comparing the confusion that is in store for us with that 
which we have had in the past.” Where in the past (as 
embodied in the recent editions of Gray’s Manual, the book 
selected by this author for his comparisons and generaliza- 
tions), will he find 25 per cent. of the names changed, as has 
occurred within four years in the case of our ferns, and that after 
the names were said to be established on strict priority principles? 

The same author in speaking of the Rochester Code has writ- 
ten further: ‘‘Those who oppose this movement, if there be any 
(and I have no doubt there are) who really see that it might be the 
last time that serious changes would have to be made in botani- 
cal names, would seem to do so purely from a personal disincli- 
nation to incur the annoyance of accustoming themselves to a 
new set of names. It must be admitted that “is motive ts not as 
high as we might hope botanists generally to be actuated by | italics 
ours].”7* An associate of this writer on the committee has 
expressed ‘‘the hope that Dr. Robinson and the few who think 
with him on this subject will day aside personal prejudices and join 
the remaining nine tenths of our botanists . . . . ina nomenclature 
based on scientific needs and a scientific method | italics ours ].”’** 
Both of these authors wrote in 1895, when the Check List was a 
comparatively new topic for discussion. Can it be that now, in 
view of the facts here presented, they still believe that the Check 
List really represented “the last time that serious changes would 
have to be made in botanical names,” or that the loose and 
undiscriminating methods employed by many who are now 
active exponents of the Rochester Code are bringing us any 
nearer that “last time?” 

7° WaRD, L, F.: idid., 316. 

** WaRD, L. F,: tid., 319. 22 COVILLE, F. V.: Bor. GAZ., 20: 167. 


196 BOTANICAL GAZETTE [MARCH 


In formulating a system of nomenclature we should keep 
constantly in view the “question of practical utility.” If this 
fundamental point is neglected, what woeful confusion must be 
encountered by everyone who attempts to use plant names! 
Already matters have reached such a state, that few followers of 
the Rochester Code can say offhand what many common plants 
should be called. The well-known species described by Vente- 
nat as Dalea purpurea, then by Michaux as Petalostemum viola- 
ceum, but generally known of late as Petalostemon violaceus, has 
been treated as follows during the past decade. Otto Kuntze, 
in his Revisio Generum Plantarum, called it Kuhniastera violacea, 
ascribing the name to Aiton, who, however, wrote Awhnistera 
violacea. This latter name is taken up by Kellerman and Werner 
who ascribe it to Otto Kuntze although (according to Steudel’s 
Nomenclator, ed. 2, 1:851, a well-known work) the name origi- 
nated with Aiton. In the Metaspermae of the Minnesota Valley the 
plant is called Kuhnistera purpurea (Vent.) MacMillan ; but it has 
recently been published as Petalostemon purpureum (Vent.) Ryd- 
berg, and in Mr. Heller’s new Casalogue it is listed essentially 
under this name (as P. purpureus). After these Jekyll-and-Hyde- 
like changes it is certainly reassuring to see Dr. Jekyll getting 
the upper hand, and to find in the latest writings of some of the 
reformers the long established name Petalostemum (on) reappear- 
ing. But do these names used by various reformers represent 
uniformity? Even if an occasional systematic botanist can keep 
track of the changes in names, how about the morphologist, the 
histologist, the physiologist, the pathologist, the paleontologist, 
the ecologist, to say nothing of the horticulturist, the pharma- 
cist, and the everyday student of plants? Should not all these 
followers of pure or applied botany be considered in our inter- 
pretation of the ‘question of practical utility?’’? And what-can 
they hope forina system of names which shows no more stability 
than the one under discussion? 

To the student whose work is in other fields than systematic 
botany, the present lack of uniformity in plant names is neces- 
sarily most perplexing. But to the systematist, who sees more 


| 


Igor] NOMENCLATORIAL PRINCIPLES 197 


closely the constant haggling over names, the situation is quite 
as puzzling. The Rochester Code was formulated ostensibly to 
establish uniformity in our names. Its followers have worked 
vigorously to comply with its rulings. From time to time 
their attention has been publicly called to fundamentally weak 
spots in its wording. Again they have been asked to explain 
certain of their names seemingly inconsistent with their prin- 
ciples. Yet these criticisms have generally been ignored. 
Instead of strengthening the weak spots in their rules and cor- 
recting self-evident mistakes in their names, the reformers have 
faithfully clung to the discredited gods they had already set up. 
These statements are not extravagant nor vague generalizations. 
They are simple conclusions drawn from the facts presented in 
this discussion, and from others very apparent upon many recent 
pages. Is this the best the Rochester Code can do? Is this 
what we are to call “uniformity?” 

If we are really desirous of obtaining stability in our nomen- 
clature, and if at the same time the “question of practical 
utility’ is to be considered, our clearest course cannot be by the 
Rochester Code, especially as followed by its originators. We 
shall, however, find a comparatively clear and practical method 
by adopting in our selection of generic names the Berlin rule; 
and in our selection of specific names, the so-called Kew rule of 
retaining the first specific name used under the accepted genus. 
In this way we are able to retain a very large proportion of the 
long-established and best-known combinations, without the 
necessity of wading (often blindly) through the mazes of 
obscure and poorly indexed literature. And, what is better, 
after comparatively slight alteration of the long-established 
names, we can feel that in only very rare cases must we abandon 
those known to practically all botanists. If, like Professor 
Ward, we all feel that “it is no longer a question of credit, but 
a question of practical utility,” is not this simpler course worth 
testing ? 


Gray HERBARIUM, HARVARD UNIVERSITY. 


BRIEFER ARTICLES. 


OBSERVATIONS UPON THE FEEDING PLASMODIA OF 
FULIGO SEPTICA. 


(WITH ONE FIGURE) 


ALTHOUGH considerable attention has been paid to the plasmodia 
of the Mycetozoa, especially by the German botanists (De Bary, Zopf, 
Sachs), little work has been done upon the feeding habits of these 
interesting protoplasmic masses. In a valuable contribution to the life 
history of these organisms, Lister’ sets forth in a painstaking way the 
manner in which the plasmodium of Badhamia utricularis behaves 
when actively feeding. Various substances were tried by way of exper- 
iment. Pieces of Agaricus campestris, A. melleus, A. rubescens, A. fas- 
cicularis, Boletus flavus, and Cortictum puteanum were used, but none of 
these fungi seemed so desirable a food as Stereum hirsutum, which was 
devoured without leaving anyresidue. Agaricus fascicularis was found 
in these experiments to be a particularly unwholesome morsel. The 
digestion by the active plasmodium of the fungi above mentioned pre- 
supposes the presence of a nitrogenous ferment, namely a proteo- 
hydrolytic one. As far as Lister’s observations show, starch seems to 
be refused by the moving plasmodium, contradicting the idea of the 
presence of a diastatic ferment. The following observations upon the 
plasmodium of Fudigo septica is in part a contribution to the life his- 
tory of plasmodia in general. 

While searching for Mycetozoa in the wooded valley incorporated 
as part of Woodlands cemetery, West Philadelphia, a luxuriant growth 
of Pleurotus sapidus was found upon some partially decayed logs, which 
had been piled up in a loose manner preparatory to burning. In 
removing several large pieces of this fungus, small patches of yellow 
plasmodium were found upon the lamellar surface of the fully expanded 
pilei. These protoplasmic masses had moved out from the rotten log 
where they were seen in the crevices, and had invaded the gill surface 

* Notes on the plasmodium of Badhamia utricularis and Brefeldia maxima. Ann. 
Bot. 2 : 1-23. 1888. : 

198, [MARCH 


tgor | BRIEFER ARTICLES 199 


of Pleurotus sapidus. The appearance of the larger plasmodium at 
this time may be described as follows: The gills which were still 
rigid and in natural position were connected in the invaded portions 
of the lamellar surface by bridges of slimy yellow protoplasm. ‘The 
basidial layers were covered by the more delicate portions of the plas- 
modial reticulum. The larger, more cord-like streams of protoplasm 
stretched from gill to gill, connecting as main cables the outlying 
pseudopodial fingers of protoplasm. The plasmodia growing upon 
several separate pieces of fungus were removed at 2 P. M. Friday, 
November 2, carried to the botanical laboratory of the University of 
Pennsylvania, and covered by two bell jars provided with dampened 
filter paper. By 6 p. M. of the same day the larger plasmodium had 
increased to such an extent as to cover completely the fungal pieces 
under one of the bell jars, and the gills showed signs of collapse. At 
9 A. M., Saturday, November 3, the gills were found to be in a total 
state of collapse, Fuligo by this time having taken complete posses- 
sion. Under the other bell jar the plasmodium, which was originally 
about the size of a silver dollar in superficial extent, had increased 
until it had spread to the outer circumferential margin of the lamellar 
area. In their attack upon the edible portions of Pleurotus, masses of 
protoplasm heaped themselves up into rounded: knobs, or protuber- 
ances formed by condensations of the myxomycete substance. These 
would disappear, to be finally replaced by. others of similar size and 
form. These observations were made on Saturday morning. The 
invasion and destruction of the gill surface was complete by Monday. 
An examination of the growth under the bell jars showed a most 
remarkable development of the larger plasmodium. It not only covered 
the fungus, but also the inner sides of the bell jar in the form of a 
beautiful yellow reticulum. The wet filter paper plastered upon the 
top of the bell jar was completely covered by a dense mass of anasto- 
mosing protoplasm. Upon the main currents of plasmodial move- 
ment were beads of protoplasm of larger and smaller size. Where 
these hung, as pendent drops on the moist filter paper, they had grown 
until the protoplasm hung, as yellow stalactites, dangling from the 
dome-like roof of the inner side of the bell jar. The dome of the bell 
jar on Monday was almost entirely covered by the yellow plasmodium. 
A strip of filter paper with the actively streaming plasmodium of 
Fuligo was removed from the moist chamber and placed in a dry situ- 
ation in the bright sunlight. As the filter paper dried, the protoplasm 


200 BOTANICAL GAZETTE [MARCH 


rapidly streamed to the wettest portion, and then began to aggregate 
into an extended aethalium. The drying, however, took place so 
rapidly that the entire plasmodium had not time to withdraw itself 
from the filter paper, and therefore it dried zx se¢u, leaving a character- 
istic network of dry anastomosing threads. The reproduced photo- 
graph was taken by Mr. W. H. 
Walmsley just before the moving 
plasmodium was placed in the sun- 
light. 

Beneath the bell jars, as the dis- 
integration of the fungi proceeded 
with the production of a watery fluid 
and a gelatinous substance of a ropy 
consistency, the more active proto- 
plasmic masses confined themselves 
to the tougher portions (the stipe and 
main substance of the pileus), heap- 
ing up on these portions in the con- 
centrated effort to digest them. With 
the drying of the interior of the bell 
jars, condensation of the reticulum 
took place, so that the meshes became 
smaller and the anastomosing streams 
more closely aggregated. At this 


Fic. 1.—Moving plasmodium of 
Huligo septica on moist filter paper, begun by feeding Fuligo various sub- 
showing the varicose condition of the stances. 
recticulum while actively streaming. At 1:30 P.M. Tuesday, November 

6, pieces of fresh young Pleurotus 

were placed upon the living plasmodium. At 2:15 p.m. the small 
pieces used were invaded and the plasmodium had spread over about 
half of the superficial surface of the fungal food. Pieces of young 
pilei of Coprinus comatus were placed on different spots of the same 
plasmodium, and by 2:15 P.M. invasion had well advanced, but the 
rapidity of forward movement was much less when Pleurotus was used 
as a food. Two hours after the two food substances had been placed 
within reach of the streaming protoplasm, complete covering of the 
specimens had occurred. 

Pieces of a partially dried toadstool, Hypholoma perplexum, were also 


1901] BRIEFER ARTICLES 201 


placed within contact of another plasmodium of Fuligo at 1:30 P.M, 
November 6, but at five o’clock not a single pseudopodial branch of 
the plasmodium had moved up upon them. On the contrary, when at 
four o’clock several pieces of the pileus and stem of Coprinus atra- 
mentarius were laid upon the yellow protoplasmic mass, inside of three 
quarters of an hour the trophotropic action of the food substance 
began to manifest itself by the plasmodial invasion of the newly pro- 
vided nitrogenous food substance. The incorporation of the nutritive 
matter had well proceeded up over the edges of the young pilei of 
Coprinus atramentarius by five o’clock, November 6. An examination 
on the morning of November 7 showed the inky Coprinus almost 
entirely digested, and a black mass of spores in a gelatinous matrix 
indicated a total collapse of the fungus. Aypholoma perplexum was also 
covered by a network of the mycelium, which had spread not only over 
the stipe, but also over the gills and upper surface of the pileus. By 
evening, this agaric had collapsed, and by the next morning, Novem- 
ber 8, nothing remained but a soft gelatinous mass of substance. 

Raw beefsteak was applied to the surface of the plasmodium at 
II: 30 A.M., November 7, and by noon a single strand of protoplasm 
had advabcel upon the meat. At 1:30 P.M. one third of the surface 
of the meat, and by 5: 00 p.m. the entire surface, was covered. Diges- 
tion must have been rapid during the night, because upon returning 
to the laboratory in the morning of November 8 not a trace of the 
beefsteak was to be found. 

Pieces of the gleba and stipe of Phallus impudicus were also applied 
at the same time. During the afternoon of Wednesday, November 7, 
the pieces of gleba were well covered by the moving plasmodium, the 
stipe portions being left untouched. By the next morning the glebal 
pieces had almost entirely disappeared, and cuts from the stipe still 
remained untouched. 

Beefsteak was again supplied to the plasmodium at 10:00 A.M., 
Thursday, November 8, and by 10: 45 a.M. a few arms of the plasmo- 
dium had extended themselves over the free edges of the meat. 

The purpose was next to extend the series of observations by feed- 
ing to the active plasmodium a variety of nitrogenous and fatty 
materials. Cheese, boiled white of egg, boiled yolk of egg, and butter 
were chosen. Pieces of these substances were applied to the surface 
of the reticulum on Thursday morning, November 8. The plasmodium 
seemed at first to refuse them, but by Friday morning the fragments 


202 BOTANICAL GAZETTE [MARCH 


of boiled white of egg were found to be partially covered by the creep- 
ing Fuligo. On Saturday morning, November 1o, the hardened, 
coagulated egg albumen was completely covered and well-nigh digested. 
The yolk was but slightly affected by the plasmodium, even after 
exposure to the digestive action for two whole days. The butter was 
left untouched. 

The presence of several ferments is naturally inferred from the 
digestive action accomplished by the plasmodium. According to De 
Bary, diastase can be extracted from the plasmodium of A‘thalium 
(Fuligo).? | 

In his book on ferments Green? states: “One of the earliest known 
of these is the ferment which Krukenberg found to be procurable from 
the plasmodium of A‘thalium, one of the Myxomycetes. A glycerine 
extract of the plasmodium was found to have very marked proteolytic 
powers in the presence of lactic or hydrochloric acid. Krukenberg’s 
statement has been confirmed by Miss Greenwood, who has stated that 
the plasmodium of another member of the same group yielded to 0.4 per 
cent. hydrochloric acid an extract which showed marked solvent action 
on fibrin.” Negative results were obtained when I removed some of 
the partially digested fungus with plasmodium upon it, and treated 
the mass with glycerine, according to the directions given above. To 
the glycerine extract, which had a slightly yellowish color, a few drops 
of 35 per cent. hydrochloric acid was added, and a small frayed piece 
of raw beefsteak. After two days of trial the beefsteak was found 
unchanged, although left in the glycerine extract for that length of time. 

The plasmodium brought into the laboratory on Friday, November 
2, was still in a streaming condition on Saturday, November 10, when 
observation upon it ceased. The original fungus, with the exception 
of the more fibrous stipe, had in this time been reduced to a fibrous 
gelatinous mass, upon which the plasmodium still streamed, finding 
apparently enough remaining food to feed upon, although by this time 
the common mold had invaded it. This mold appeared for the first 
time on Wednesday, November 7, but was then brushed off to prevent 
fruiting. The plasmodium, while actively streaming and feeding, kept 
the substratum remarkably sweet and clean, and it was not until the 
original food substance had been destroyed as food that foreign 
organisms, such as the mold, had any chance for development. This 

?This statement is somewhat at variance with the observations of Lister, /oc. cit. 


3 The soluble ferments and fermentation 215. 1899. 


1901] BRIEFER ARTICLES 203 


was certainly one of the most instructive facts brought out during the 
course of my observations..— JOHN W. HARSHBERGER, University of 
Pennsylvania. 


SWARM SPORE FORMATION IN HYDRODICTYON 
UTRICULATUM Ror: 
1. The methods of fixing were by means of Merkel’s fluid and a 
mixture of iridium chlorid and acetic acid according to one of the 
following formulae: 


(1) Eisen. Iridium chlorid Se 5 BER cent. austere oh a a - 100° 
lacial acetic a - ee 

(2) Iridium chlorid (1 per she oars solution) - - - 100% 
ce 


Glacial acetic acid - - - 

The best results were obtained with the stronger iridium chlorid 
mixture. 

2. There is no differentiated chromatophore in the cell. The 
pyrenoids and nuclei are scattered irregularly throughout the cyto- 
plasm and the chlorophyll is contained in the whole cytoplasmic body. 
The nuclei in both the resting and dividing stages show the structure 
typical of higher plants and are not to be taken as types of primitive 
nuclei. 

3. Cleavage takes place by means of surface constriction of the 
plasma membrane on the outside and the vacuolar membrane on the 
inside of the protoplasmic Jayer. The process is a progressive one, 
the cleavage furrows cutting out first large irregular multinucleated 
masses of protoplasm, which are in turn divided into smaller masses, 
until each contains a single nucleus, the entire protoplast thus being divi- 
ded into spores. The swarm spores are uninucleated biciliated cells. 
At the base of the pair of cilia there is a clearly defined basal body. 

A ‘detailed description of the processes outlined above will be pub- 
lished soon in a more complete form.— H. G. TIMBERLAKE, Cniversity 
of Wisconsin. 

4 One of the best methods of procuring material for microscopic study is to remove 


the protoplasm by scraping, and then to place portions of it on slides fitted into the 
bottom of Petri dishes provided wit mei filter paper. In an hour or two these 


mounds of protoplasm will have spre to permit of their 
examination, At my suggestion, Dr. Mazijck Ravenel, Sy cecrlalogiet of the Pennsyl- 


vania Live Stock Sanitary Board, tried to grow the plasmodium upon ‘eben agar, 
and upon filter paper saturated with bouillon, but failure resulted in both cas 

5Rés of results presented at the me of the Western Naturalists at the 
Hull inka Laboratory, December 27, 1 


GURREN LITERATURE. 
BOOK REVIEWS. 
Goebel’s Organography.* 

THE second volume of this notable work has appeared recently. It deals 
with the gametophyte and sporophyte of pteridophytes, and with the sporo- 
phyte of spermatophytes. The gametophyte of the pteridophytes is discussed 
under two heads, namely, (1) structure and development of the sex organs, 
and (2) the form of the prothallia. In discussing the development of anthe- 
ridia the author advances views which are at variance not only with those of 
Belajeff and others, but also with his own previous accounts. In Isoetes, after 
the cutting off of the small cell which Belajeff has called the rhizoidal cell, 
two oblique walls divide the main body of the spore into two flat cells anda 
larger cell which is triangular in optical section. This last cell alone the 
author regards as the antheridium initial. It divides by a periclinal wall into 
an outer cover cell, and an inner cell, the latter of which by further division 
gives rise to four cells in each of which a spermatozoid is organized. Accord- 
ing to this interpretation there is within the spore wall a prothallium consist- 
ing of three sterile cells and one antheridium, and only the cover cell can be 
regarded as belonging to the antheridium wall. 

In presenting the development of both the antheridia and the archegonia 
the transition from free to imbedded organs is described in some detail. The 
prothallia of Ophioglossum, Botrychium, aud Lycopodium receive particular 
attention on account of their biological importance. The vegetative multipli- 
cation of prothallia is another interesting subject which is somewhat fully 
treated. 

In the second part of the book, which is devoted to the sporophyte of 
pteridophytes and spermatophytes, the various organs are discussed in great 
detail. The account of the embryo is particularly helpful. Among other 
interesting subjects are the transition between leaf and shoot, leaf formation 
the relation between venation and leaf devolopment, transformed leaves, 
branching, etc. 

The treatment throughout is dominated by what may be called experi- 
mental morphology, and the book cannot fail to have a good influence in 

*GOEBEL, K.: Organographie der Pflanzen insbesondere der Archegoniaten und 
Samenpflanzen. Zweiter Teil. Specielle Organographie. 2 Heft: Pteridophytem 
und Samenpflanzen. Erster Teil. 8vo., pp. xiii-xvi+ 385-648. 173 illustrations- 
Jena: Gustav Fischer. 1900. M47. 

204 [MARCH 


{9o1] CURRENT LITERATURE 205 


relaxing the too rigid notions of morphology which are still prevalent. While 
constantly calling attention to the variation which occurs in nature and which 
may also be brought about artificially, the author also recognizes that environ- 
ment is not the only factor in plant development, but that heredity is equally 
important. Representing as it does the work which is being carried on in the 
author’s laboratory, the book has the freshness of research, and is full of 
suggestions to those engaged in morphological investigation. 

Those who have read the first part will be glad to learn that the present 
volume is not so difficult, An early English translation is announced.— 
‘CHARLES J. CHAMBERLAIN. 


MINOR NOTICES. 

THE SIXTH FASCICLE of Wildeman and Durand’s ///ustrations de la flore 
du Congo has appeared recently, containing plates 61 to 72 inclusive. This 
elaborate work, with its very handsome plates, needs no further commenda- 
tion than has been given already in this journal.— J. M. C 

THE SECOND FASCICLE of Schumann's Bliihende Kakteen (Iconographia 
Cactacearum) has appeared recently. It contains descriptions and beautiful 
colored illustrations of Mamit/aria Wissmannit Hildmann, M. raphidacantha 
Lem., Echinocactus horripilus Lem.,and E. Mathssonii Berge. The publisher 
is J. Neumann in Neudamm, and the price is four marks.—J. M. C 

THE SECOND VOLUME of Primitiae Florae Costaricensis, under the editor- 
ship of H. Pittier, bearing date 1898—1g00, has appeared in seven fascicles. 
The collaborators are J. Donnell Smith (Polypetalae and Gamopetalae), 
‘Casimir DeCandolle (Piperaceae), G. Lindau (Acanthaceae), F, Pax (Euphor- 
biaceae), and A, Engler (Araceae). Numerous new species are described, 
but the only new genus published is Ko/odohilus (Acanthaceae).—J. M. C. 

THREE FASCICLEs of the first volume of Engler and Prantl’s Vat. Pflan- 
zenfamilien have appeared recently, as follows: 204 contains the conclusion 
of fossil Filicales and the Sphenophyllales by H. Potonié, and the beginning 
of the living Equisetales by R. Sadebeck ; 205 contains the conclusion of the 
living Equisetales by R. Sadebeck, the fossil forms by H. Potonié, and the 
beginning of the Lycopodiaceae by E. Pritzel ; 206 contains the conclusion 
of the Lycopodiaceae and the Psilotaceae by E. Pritzel, the Psilotaceae 2 
H., Potonié, and the beginning of Selaginellaceae by G. Hieronymus. —J. 

A. M. Fercuson (Twelfth Ann. Rep. Mo. Bot. Gard. 33-73. hi 


and varieties are recognized, all of which are illustrated. The conservative 
Spirit of the work is indicated by the fact that in a genus of great possibilities 
only one new species is described, and five forms are made new varieties. 


206 BOTANICAL GALETTE [MARCH 


It is certainly true that the author’s conception of a species differs from that 
which is rapidly becoming current, and that he still has some belief in its_ 
power to vary.—J. M.C. 

Mr. HERMANN VON SCHRENK (Twelfth Ann. Rep. Mo. Bot. Gard. 21-23. 


some heartwood has been formed in the larger branches, and from this time 
on the mycelium may be found in the heartwood. Access is obtained through 
wounds, so that a proper treatment of wounds will ward off the disease. 
Curiously enough the fungus, although growing in what is regarded as dead 
tissues, does not grow in the wood after it is cut from the living tree, and 
hence diseased wood when used for posts does not continue to rot. hether 
such a fungus is to be regarded as a parasite or a saprophyte, therefore, 
becomes an interesting question.— J. M. C 

THE FIRST THREE PARTS of Engler’s Pflanzenreich have appeared. The 
general purpose and method of this great work were announced in this. 
journal for last December (30: 432. 1900), so that it only remains to note the 
contents of the parts as they appear. The families of spermatophytes to be 
presented are 280 in number. Tart 1 (Jf 2.40) is by K. Schumann, and con- 
tains the Musaceae, the forty-fifth family, the six genera including eighty 
species. Part 2 (J/ 2) contains the eighth and tenth families, Typhaceae 
and Sparganiaceae, and is by P. Graebner. Each family is represented by a 
single genus, Typha containing nine species and Sparganium fifteen. Part 3 
(7 5.60) contains the ninth family, Pandanaceae, and is by O. Warburg. 
He recognizes 21g species, 156 of which belongs to Pandanus. The publisher 
is Wilhelm Engelmann of Leipzig.— J. M. C 

BOTANICAL ACTIVITY in Vermont is indicated by the series of ‘‘ Contribu- 
tions to the Botany of Vermont,” which has now reached its eighth number. 
The titles of the eight contributions are as follows: A list of the mosses of 
Vermont, with analytical keys to the genera and species, by A. J. GROUT 
(March 15, 1898); A partial list of the parasitic fungi of Vermont, by W. A- 
ORTON (September 1898); A preliminary list of the Hepaticae of Vermont, 
by CxiirTton D. Howe (January 1899); Supplement to the list of mosses 
growing in the state of Vermont, by A. J. Grout (January 1899); The trees 
of Vermont, by ANNA M. CLARK, with notes on the trees of Burlington and 
vicinity, by L. R. Jones (December 1899); A second partial list of the para- 
sitic fungi of Vermont, by W. A. ORTON (December 1899); and Flora of 
Vermont, a list of the fern and seed plants growing without cultivation, by 
Ezra BRAINERD, L. R. JoNES, and W. W. EGGLestTon (December 15, 1900)- 
The last publication enumerates 1330 indigenous species.— J. M. C 

A FLORA of the German possessions (Kaiser Wilhelm’s Land) in New 
Guinea (or Papua) and the adjacent islands has been published by Drs. Karl 


1901] CURRENT LITERATURE 207 


Schumann and Karl Lauterbach.? Descriptions are given of numerous new 
and critical species, but those better known are merely named and their 
distribution given. The book begins with the Myxomycetes and follows the 
sequence of Engler and Prantl. This bringing together of a great amount 
of scattered material makes the book an admirable compendium of informa- 
tion concerning the plants of the ‘South Seas.” The new genera described 
are as follows: Dammera (Palmaceae), Scleromelum (Santalaceae), Lauter- 
bachia (Monimiaceae), Macropsychanthus (Leguminosae), Syndyophyllum 
(Euphorbiaceae), Gertrudia (Flacourtiaceae), Xenodendron (Sonneratiaceae), 
Bamlera (Melastomaceae), Kentrochrosia (Apocynaceae), and Airosperma 
(Rubiaceae).—J. M. C 

A VALUABLE CONTRIBUTION to the literature of special diseases of 
plants is the recent bulletin of the Division of Vegetable Physiology and 
Pathology on Peach leaf curl3 The bulletin is divided into eleven chapters. 
The first is introductory and treats of the distribution and origin of the dis- 
ease and of the losses caused by it, which are estimated to reach at least 
$3,000,000 annually. Next is taken up the nature of the disease itself, which 
is caused by the fungus EZxoascus deformans. It is shown that the perennial 
mycelium of this fungus is responsible for only about 2 per cent. of the 
infections each spring, the others being due to spores which have remained 
over winter in the crevices of the bark and between the bud scales. In the 
next five chapters the history of the various methods of treatment is dis- 
cussed, and the plans of the experiments for the prevention of the disease 
and the results in saving of foliage and fruit are given in great detail. It is 
shown that from 95 to 98 per cent. of the injury to the foliage can be pre- 
vented by treating the trees, while still dormant, with various sprays, the best 
being a Bordeaux mixture containing five pounds each of copper sulfate and 
lime, and forty-five gallons of water. One chapter is devoted to a discussion 
of the preparation, use, and character of the different sprays used, another 
to the methods of applying the sprays, and still another to a description of 
the various substances used in preparing them. The sprays are shown to be 
successful only when applied while the tree is dormant, preferably one to 
three weeks before the flower buds open. The final chapter is devoted to a 
comparison of the susceptibility of different peach varieties to the disease, 
and toa discussion of the treatment of nursery stock. A short summary con- 
cludes the bulletin Ernst A. BESSEY. 


? Die Flora der deutschen Schutzgebiete in der Siidsee. Small 4to, pp. xvi + 613. 
with map and 22 plates. Leipzig: Gebriider Borntraeger. 1901. M 40 unbound, 
Mf 45 bound. 

3PIERCE, NEWTON B.: Peach leaf curl: its nature and treatment. Bulletin 20. 
Division a Veosintie Physiology and Pathology, U. S. Department of Agriculture, 
8vo, pp. 204. figs. 70, pls. 30. Washington: Government Printing Office. 1900. 


208 BOTANICAL GAZETTE [MARCH 


NOTES FOR STUDENTS. 


EBERHARDT‘ has performed a series of experiments with a view to find- 
ing the influence of dry and humid air on plant structures. These experi- 
ments in general confirm the work of Kohl and others. Humid air causes an 
increase in the length of the stem and the size of the leaf surface, while there 
is a decrease in the stem diameter, the amount of chlorophyll, and the root 
development. Dry air increases the thickness of the cuticle, the number of 
stomata, the woody tissue, the sclerenchyma, and the palisades.— H. C. 
COWLEs. 


E INTERESTING STUDIES have been made by Nestler5 upon the well- 


by testing the effect upon him The hairs of Primula Sinensts act in a 
similar w t the poisonous effects are much less marked.— 
COWL 


R. G. KLess published last year® the third paper of a series on the 
aaa of reproduction in fungi in which he brings together the previ- 
ous investigations with some hitherto unpublished researches, and seeks 
to present general considerations on the whole subject. The paper is full of 
suggestions and too valuable to mutilate by an attempt to summarize it. One 
general criticism lies against Klebs’ work and his conclusions, namely, that 
he does not take into account sufficiently the effect of changes in osmotic 
pressure to which his experimental plants are subjected with the changing 
composition. It remains to be seen whether the conclusions are not vitiated 
by this untested factor.—C. R. B 

ARNEY? discusses the Lower Austral element in the southern Appa- 
lachians. The mountains have representatives from all of Merriam’s zones 
from the Lower Austral to the Hudsonian, though the Transition zone is 
most fully represented. Austro riparian colonies are found up to 1200 feet 
along the eastern boundary of Tennessee. Kearney divides the Austral 
mountain plants into those which are probably of neotropical origin and have 
come in since the ice age, and those which have probably descended from 
the Tertiary floras of northern regions. The plants of the first group are 
chiefly xerophytic, while those of the second group are mainly ligneous 
tropophytes. Lower Austral forms must have left the mountains during the 
4Compt. Rend. 131: 193-196, 513-515. 1900. 
5 Ber. deut. bot. Gesell. 18 : 189-202, 327-331. 1900. 
Jahrb. f. Wiss. Bot. 35 : 80-203. 1900. 7 Science N. S. 12 : 830-842. 1900. 


a he li aie 


1901] CURRENT LITERATURE 209 


ai pe Interesting hypotheses are given on the origin of the Austral 
form WLES 
LLE. RODRIGUE® has made a painstaking study of the anatomy of 

variegated leaves with a view to ascertaining the relation between color and 
structure. The author gives an excellent summary of the literature of her 
subject, and makes a detailed study of thirty-three species. The white effect 
is due in most cases to the absence of chlorophyll, although a similar appear- 
ance is given by certain dissolved pigments, and by the reflection of light in 
some special instances. The modifications in the latter cases are slight and 
are confined to the epidermis, Where chlorophyll is absent, the leaf may be 
regarded as diseased, and the tissues are different from normal leaves, having 
no palisade development, and consequently a great reduction in thickness, 
In other words, the hile tissues remain pitta where chlorophyll is 
absent.—H. C. Cov 

ALB, NILSSON? has made some interesting studies on the dynamics of 
some Swedish plant societies, especially of cliffs and moors. He finds three 
types of cliffs, those which are forested either with conifers or deciduous 
trees, and those without trees. On all cliffs the first plants are crustaceous 
lichens. On the conifer cliffs there follow in succession fruticose lichens, 
herbs, heath plants, conifers. Cliffs with deciduous trees have no fruticose 
lichen or heath stages, the author attributing the absence of the fruticose 
lichens to wind. On the third type of cliff the lichens remain longer and 
foliose lichens and mosses are added to the stages after crustaceous lichens. 
Dying lakes pass into sedge moors, then into cotton-grass moors, finally into 
shrub moors and forest moors with pines or birches. Retrogressive phases 
are common on the moors, lichens growing over the peat moss and shrubs; 
again the water collects and the lichens pass away. The peat moss appears 
again and we have what Nilsson calls a secondary moor.—H. C. COWLES. 

THE STYLIDIACEAE (Candolleaceae in Engler and Prantl’s Nat. Pflanzen- 
Jam), a small family almost confined to Australia, New Zealand, and farther 
India, has been recently studied by G. P. Burns. The greater part of the 
paper is devoted to a morphological study of the various tissues. Before 
fertilization the structures of the embryo sac present no unusual features, . 
but immediately after the entrance of the pollen tube the micropylar portion 
of the sac grows out into an enormous haustorium much larger than the 
remainder of the sac. The endosperm forms rapidly and fills the sac with 
tissue before the first division of the egg takes place. Shortly after fertili- 
zation the antipodals disintegrate, and the posterior portion of the sac also 


8 Mém. l’Herb, Boiss. 17: 1900. 9 Bot. Not. 1899: 89-101; 123-135. 


* Beitrage zur Kentniss der Stylidiaceen. Flora 87: 313-354- pls. 13, 14 (and 45 
text figures). 1900. 


210 BOTANICAL GAZETTE [MARCH 


forms an haustorium. Finally, the protoplasmic contents of both haustoria 
become transformed into a network of cellulose threads which in case of the 
upper haustorium form a plug eeeetially closing the micropyle. The sac is 
surrounded by a jacket or “‘tapetum’”’ which is even more conspicuous than 
in the Compositae.— CHARLES J. CHAMBERLAIN. 

THE FUNCTION of latex, so often in past years a motive for investigation, 
has again been made a subject for study. Gaucher” gives a historical sum- 
mary of the two chief views, excretory and nutritive, from the time of Trécul 
to the present. The author gives no new theories, but presents a large num- 
ber of facts which favor the nutritional function, very much as presented by 
Haberlandt. The substances contained in latex, the connection between the 
latex tubes. and the palisade, and the reciprocal relations between latex tubes 
and conductive parenchyma are all studied, and Gaucher in these cases con- 
firms and extends Haberlandt’s observations. In one instance he finds a ring 
or festoon of chlorophyll cells arranged about a latex tube. 

Parkin * has studied the latex in rubber plants of Ceylon, and holds a some- 
what intermediate view. While he regards the proteids of latex as probably 
nutritive, he does not so regard the starch, unless perhaps the latter aids in 
the nutrition of the latex tubes themselves. The author finds that the latex 
flows far less abundantly at the first tapping than subsequently, showing an 
apparent adaptation. Mane vepards the chief function of latex to be water 
storage.—H. C. Cow. 

SOME VALUABLE CONTRIBUTIONS to the literature of forest distribution 
have been made recently by the United States Geological Survey. This 
report is under the supervision of Henry Gannett, chief of the division, and 
is a companion volume to a similar one published last year. It contains 
special considerations of the Pike’s peak, Plum creek, and South Platte 
reserves by John G. Jack; White river plateau timber land reserve by George 
B. Sudworth; the Flathead forest reserve by H. B. Ayres; and the Bitter- 
root forest reserve by John B. Leiberg, Topographic features, soil condi- 
tions, climate and rainfall, forest conditions, fires, and lumbering are some 
of the topics treated in these reports. A large number of plates, including 
both maps and reproductions from photographs, are incorporated in the 
volume, and a portfolio containing topographic eg showing distribution of 
timber areas presents the subject in a graphic w 

If the department would but i incorporate in its eweaiiee: geological and 
physiographical atlases an additional topographic map showing the distribu- 
tion of forest and other floral areas, including descriptions of the edaphic 
and climatic conditions, it would add much to their educational, economic, 


Ann, Sci. Nat. Bot. VIII. 12: 241-260. 1900. Ann. Bot. 14 : 193-214. 1900. 


3Twentieth Annual Report, U.S. G. S., Part V. Forest reserves, pp. xviii + 498. 
pls. 159. 1898-9. 


1901 | CURRENT LITERATURE 211 


and scientific value. Some such careful study of a large number of floral 
areas is an absolute necessity to a correct understanding of the complex 
climatic and ecologic factors governing the distribution of trees. Indeed, in 
the last atlas" issued by the department an approach has been made to such 
a realization. This atlas contains a brief summary of the vegetal and cli- 
matic features, a map of the floral features, and three maps showing precipi- 
tation, evaporation, and types of rainfall—H. N. WHITFOR 

THE STUDIES of Brenner™ on succulent plants must prove of great inter- 
est to all physiologists and ecologists. His work was experimental and for 
the most part on the Crassulaceae and Mesembryanthemum. After a dis- 
cussion of the normal anatomy, he describes the effects produced on succu- 
lent plants by moist air. The most striking effect in Sedum is pronounced 
internodal elongation, which the author refers to the increased turgor inci- 
dent to lessened transpiration. At first the leaves are fleshy, but later leaves 
are larger and thinner and placed like normal fleshy leaves in dry air in the 
form of a rosette. There is thus a striking correlation between stem elonga- 
tion and leaf form. Another effect of moist air on the leaves is epinasty, so 
that the new leaves place themselves at right angles to the stem; when these 
plants are placed in a dry chamber hyponasty is shown. Notable changes in 
anatomy are also induced. In two plants the normally straight side walls o 
the epidermis become wavy, and doubtless give greater mechanical strength 
to the otherwise weakened leaf. The tangential increase of the epidermal 
cells as against the radial is very noticeable, though Brenner is at a loss to 
find a physical explanation therefor. The stomata at first are the same in 
number as on normal leaves, though of course they are farther apart, since 
the leaf is larger. On later leaves the stomata are more numerous though 
the number per unit area may be much as in normal leaves. There is a 
decrease of the storage tissue and an increase of the chlorophyll tissue, 
though the cells in the latter tissue are more nearly isodiametric than i in dry 
air. The vascular system and air spaces are decreased in moist air; the 
reduction in the bundles is rather in number of cells and pi rise thas 
in cell size. The author finds the dry weight and ash and also the acid con- 
tent to be less than in normal plants. At first moist air increases the size of 
the chloroplasts, though they decrease in size later, pointing to an apparent 
readjustment to the new conditions. 

Various comparative physiological experiments were made on plants 
grown in dry and moist air. In Mesembryanthemum nutation movements 
were noticed in the latter but not in the former. Normal leaves in normal 
air transpire the same per unit area as do moist-air leaves in moist air, 
though the leaf form is very different. This observation is very instructive 


* HILL, ROBERT T.: Topographic atlas of the Texas region, pp. 12. p/s. 77. 1900. 


5 Flora 87 : 387-439. 1900. 


212 BOTANICAL GAZETTE [MARCH 


as it shows strong powers of readjustment in such highly specialized forms as 
succulent plants. The author concludes by saying that the air and not the 
soil relations are determinative for the above changes. This is in harmony 
with Kohl’s results on Tropaeolum. Brenner thinks that the phenomena 
which he observed are in a high degree purposeful, and that ene, physical 
explanations are very difficult at many points.— H. C. Cov 

ITEMS OF TAXONOMIC INTEREST are as follows: ARTHUR MINKs (Mém. 
Herb. Boiss. 22: 1-74. 1900) has published a full discussion and synopsis 
of the genus Umbilicaria.— WILLIAM R. Maxon (Proc. Biol. Soc. Washing- 
ington 13: 199, 200. 1900) has described a new Polypodium (P. hesperium), 
which is “the common form of the whole mountain region of the western 


described a new Dryofteris from Alaska.— P. A. RYDBERG (Bull Torr. Bot. 
Club 27: 614-636. 1900), in continuing his ‘Studies on the Rocky mountain 
flora,” has published an account of some of the smaller genera of Compositae. 
Those considered are Stenotus, formerly a section of Aplopappus, containing 
seven species, of which two are new; Stenotopsis, a new genus established on 
Aplopappus (Stenotus) linearifolius, and including also Aplopappus Sepanee 
interior ; Macronema, containing seven species, of which one is new; 

ranthus, a genus revived to include species formerly under Aplopappus, a 
more lately under Eriocarpum, and which is recognized as containing seven 
species, three of which are new; Pyrrocoma, with sixteen species, five of 
which are new; Ba/samorrhiza, with nine species, two of which are new; 
Thelesperma, with seven species, two of which are new; Hymenopappus, with 
eight species, four of which are new.—EpwarpD L. GREENE (Pittonia 
4: 159-226. 1900) has recently made some important contributions as fol- 
lows: A fascicle of new forms of Arnica contains twenty-four species; 
Gentianaceae are enriched by three new species of Gentiana, three of Swertia, 
and three of Frasera; the third of the ‘Studies in the Cruciferae” discusses 
certain species of Avadzs, describing seventeen as new, describes new species 
in Cheiranthus, Sophia, Thelypodium, Thysanocarpus, Draba, and Cardamine 
(4 spp.), expresses his conclusion as to the type of the genus Draéda, and estab- 
lishes a new genus (Aédra) upon what is known as Draba brachycarpa,; the 
second of the papers on “ Neglected generic types’’ brings to us FHlaler- 
pestes as a new genus established to include Ranunculus Cymbalaria Pursh, 
R. salsuginosus Pallas, and R. tridentatus HBK., Peritoma DC. to include 
certain species of Cleome (serru/ata, inornata lutea), Celome founded on 
Cleome platycarpa Torr., Carsonia founded on Cleome sparsifolia, and Alde- 
nella founded on .Polanisia tenuifolia T. & G.; eighteen new species are 
added to the genus Aséer, all but one of which are from the Rocky mountains; 
among the “ Corrections in nomenclature’ Oveostemma is substituted for the 
untenable Oveastrum Greene (containing certain species formerly referred 


ac ce 


1901] CURRENT LITERATURE 213 


to Aster), WVerisyrenta is substituted for the untenable Parrasia Greene 
(Greggia), and Evemosemium is substituted for Grayda of the western deserts 
on account of a prior use of the name. The same author (dem 227-241. 
1901) has begun the segregation of Zavaxacum in North America by 
describing eight new species; and has described new species under 7ha/tc- 
trum, Rumex (2), Lappula, Allocarya (2), Solidago, Coleosanthus (5), Coreop- 
sts, Parthenium, Picradenia, and Zygadenus (2).— M. L. FERNALD (Rhodora 
230-233. P/. 27. 1900) has described two new northeastern species of 
Thalictrum, and two new varieties of Scirpus maritimus (idem 241. 1900), 
and has presented (zdem 3: 13-16. 1901) Monarda fistu/losa and its allies.— 
B. L. ROBINSON (Rhodora 2: 235-238. 1900) has discussed and reorganized 
the nomenclature of the New England representatives of Agrimonia, has 
presented (¢dem 3:11-13. 1901) the results of his search for the type of the 
Linnean Guaphalium plantagintfolium which proves to be 4. planiaginea as 
interpreted by Fernald, and has discovered (édem 16-17. 1901) that Sésym- 
rium Niagarense Fourn. should be transferred as a doubtful synonym under 
S. officinale L. to Brassica nigra Koch.—G. E. DAVENPORT (Rhodora 3: 1-2. 
Bl. 22. 1901) has deecetiaet: a new plumose variety of Asplenium ebeneum 
from Vermont.— J. M. GREENMAN (Rhodora 3 : 3-7. 1901) has set forth the 
genus Senecio as it exists in New England, describing two new varieties of 
S. Balsamitae— SPENCER LE MOORE (Jour. Bot. 38: 457~469. Ai. 476. 1900) 
has described two new genera of Compositae from Africa (De/amerea and 
Nicolasia), both belonging to the Inuloideae —A. B. RENDLE (Jour. Bot. 
39: 12-22. Igor) has described eleven new species of /fomoea from Africa.— 
F. LAMSON-SCRIBNER and ELMER D. MERRILL (U. S. Dept. of Agric., Div. 
of Agrost. Bull. 24: 1-54. 1901) have published new species of 77ifsacum, 
Andropogon (3), Paspalum (2), Panicum (3), Muhlenbergia, Agrostis, Tris- 
tachya, Leptochloa, Aristida (2), and Elymus (5); and have given the results 
of a study of the types of Panicum nitidum, P. pubescens, and P. scoparium.— 
J. M. C. 


IN A RECENT PAPER Hans Fitting * has given the results of his investiga- 
tions on the mode of origin of the megaspores, and the development of their 
Coats in Isoetes and Selaginella. His work was done chiefly with living spores 
examined in a physiological salt solution, and in water. Microtome sec- 
tions were used to trace the phases of karyokinesis in the spore mother cells, 
and for a check on the conclusions drawn from the living materia 

He agrees with Smith” in his account of the origin of the sporangium of 
Isoetes. It will be remembered that the latter author differed from Goebel, 
Bower, and Campbell, in asserting that “the rudiment of the sporangium is 

**Bau und Entwickelungsgeschichte der Makrosporen von Isoetes und Selagi- 
nella, etc. Bot. Zeit. 58: 107-164. pls. 5-6 

7 Bor. Gaz. 29 : 225-258, 323-346. pls. 22-20. 1900. 


214 BOTANICAL GAZETTE [MARCH 


a transverse row of superficial cells below the ligule.” Also, as regards 
the formation of the trabeculae and tapetum, Fitting’s account is identical 
with Smith's. 

The spore mother cell is distinguished by its finely granular protoplasm, 
large nucleus, and nucleolus. At one side of the nucleus lies a dense mass 
of coarse-grained protoplasm, in which are imbedded many small starch 
grains. Preceding the first division of the mother cell, radiations appear in 
the protoplasm, extending in all directions to the wall, but from no common 
center. The mass of mingled protoplasm and starch divides into two nearly 
equal parts, and new radiations appear between them as they separate. 
They finally take the positions of two foci of an ellipse, the spore mother 
cell being nearly of that shape. During this process the nucleus has shifted 
from the center to the periphery of the cell, and at its conclusion has returned 
to its original position. The two daughter masses (Zochter-Klumpen) 
elongate and lie in planes at right angles to each other and to the long axis 
of the cell. The starch grains arrange themselves in straight lines in each 
mass. Those near the middle slip toward either end and reunite in two 
groups, surrounded by the dense protoplasm. This process results in four 
masses of starch surrounded by the coarse granular protoplasm, arranged 
tetrahedrally. These changes the author followed in living material, observ- 
ing the spore through the sporangium wall and the several layers of sterile 
cells, tapetum, etc. The nucleus then divides by two rapid successive divi- 
sions, the spindle lying in such fashion that each of the four daughter nuclei 
lies by one of the Zochter-Kiumpen. New fibers arise from the surrounding 
protoplasm and extend themselves between the nuclei, thus forming a sex- 
tuple spindle. Equatorial cell-plates cross these spindles, cutting completely 
through the protoplasm in six planes from the center of the cell to its wall. 
Partition walls develop in these plates. It is evident that four of these 
walls have no connection with the spindles concerned in the division of the 
nucleus. 

The origin of the four megaspore membranes is worked out with great 
detail. The main points are as follows: While the four “* special mother” 
cells are still lying in the form of a tetrad enclosed by the mother cell mem- 
brane, each surrounds itself with a separate membrane called the «special 
mother cell”. membrane. The author did not determine whether this was 
formed by the mother cell membrane or by the protoplasm of the special 
mother cell. This thickens rapidly and divides into three lamellae which taken 
together constitute the erosfore. Between the exospore and the protoplasm 
of the cell (now called megaspore) developing from the latter, appears the 
mesospore. The outer layer of the exospore becomes roughened with spines 
or reticulations, and following their exact contour is laid down an incrusta- 
tion with much silica, which Fitting Styles the Perisfore. Some species of 


1901] CURRENT LITERATURE 215 


Isoetes lack this coat. All of these membranes grow by intussusception, and 
the author lays stress on the fact that the perispore and exospore are of 
quite different chemical nature, and yet are both growing simultaneously by 
intussusception. Finally, between the mesospore and the protoplasm con- 
tent appears a thin film of cellulose, the endospore. The nourishment 
needed for the growth of these membranes is derived from the sporangium 
wall and trabeculae, not from the tapetal cells. Until the walls are formed 
the spore content is relatively very small. 

The author was less successful in his work in Selaginella. Owing to the 
smallness of the megaspore and to imperfect technique (he never succeeded 
in avoiding shrinkage) he failed, like all his predecessors, to make out the 
stages of the development of the megaspore. The megaspore mother cell is 
easily recognized, but how it divides into spores is not known. 

Heinsen’s account and Fitting’s disagree in almost every particular as 
regards the interpretation of the spore contents and the origin of the several 
coats. The “nucleus” (according to Heinsen) is the entire protoplasmic 
content. Heinsen’s “nucleolus” Fitting interprets as the nucleus. The 
several small “corpuscles,” whose nature Heinsen could not explain, are, 
according to this author, the nucleoli. The sequence of events as regards 
the development of the coats is much like that of Isoetes, making an addi- 
tional reason why [soetes and Selaginella should not be separated in any 
system of classification. The author thinks that the extremely smal] amount 
of protoplasm in the spore can have nothing to do with the nourishment of 
the spore coats, which soon far exceed it in thickness and bulk. Between 
the tapetum and the four megaspores is a sort of slimy matter which Bower 
interpreted as the remains of the disorganized sterile mother cells. Fitting 
Says that these cells do not disorganize, and that the slime is a secretion from 
the tapetum, which acts like a gland. This material nourishes the spores up 
to their maturity, when they fill the entire sporangial cavity. Like those of 
Isoetes, the spore walls develop by intussusception. A very significant fact 
is that the greatest growth of the spore walls takes place when they are not 
in contact with the plasma body within. Four walls are found, exospore, 
mesospore, endospore, and perispore (the latter sometimes lacking in certain 
species). The increase in size of the plasma body without corresponding 
increase in the amount of matter of which it consists, followed by cell divi- 
sion and the formation of the prothallium, were not followed in detail. The 
author says, however, that in some species this occurs before the spores are 
shed (S. Martensii, S. Galeottiz), and in others ‘a long time afterward,”’— 
FLORENCE May Lyon. 


NEWS. 


On MARCH 30 the Imperial Zoological-botanical Society of Vienna will 
celebrate its fiftieth anniversary by a jubilee session in which related acade- 
mies, societies, associations, and institutes are invited to participate. 

A NEW monthly botanical journal, 7orreya, has been begun by the Torrey 
Botanical Club of New York City, under the editorship of Marshall A. Howe. 
The journal is intended for the shorter notes and items which have been lately 
rather crowded out of the Budletin. 

THE YALE SUMMER ScHootr of Forestry, under the direction of Henry 5S. 
Graves and James W. Toumey, will hold its sessions at Grey Towers, the 
estate of Mr. James W. Pinchot, near Milford, Pa. The school begins July 
8, I901, and continues eight weeks. 

IN A RECENT “minor notice” (30:418. 1900) in this journal, certain 
publications being issued by the botanists of Japan were confused. A letter 
from Professor Y. Yabe makes the following statement: Three series are 
being issued, whose titles may lead to confusion. Jcones Florae Japonicae is 
a university (Imperial Univ., Tokyo) publication, and but one part has 
appeared, having been issued last April. It is in the editorial charge of Mr. 
T. Makino. Phanerogamae et Pteridophytae Japonicae Icon. Iilustr. and 
Cryptogamae Japonicae Icon. Ilustr. are the private publications of a few 
botanists. 

Tue Soctety for Plant Morphology and Physiology met at Johns Hop- 
kins Medical School, Baltimore, Md., Dec. 27 and 28, rgoo, under the presi- 
dency of D. P. Penhallow. The following officers were elected for the 
ensuing year: President, ERWIN F. SMITH; vice presidents, F.C. NEWCOMBE 
and L. M. UNDERWOOD; secretary, W. F. GANONG. The following new 
members were elected: M. A. CARLETON, F. D. CHESTER, E. B. COPELAND, 
T. H. KEARNEY, J. W. ToumMEy. The most important business of general 
interest was the presentation in printed form of the report of the committee 
on reviews of botanical literature, which has already been referred to in this 
journal (p. 143). A committee (Messrs. Ganong, Lloyd, and Atkinson) was 
appointed to take into consideration the subject of a standard college 
entrance option in botany, to be made as widely acceptable as possible. 
The address of the retiring president, 4 Decade of North American Paleo- 
botany, has been published in full in Science, together with abstracts of the 
papers presented, 


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GOLD MEDAL, PARIS, 1900 W E R F % | 


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TRADE-MARK. 


WEBER W 
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Established 1780. DORCHESTER, MASS. {81 Tremont Street, Boston. 


WAREROOMS : 


Vol. XXXI APRIL, 1901 No. 4 (*/ 
“4 THE; See ae 
-— BOTANICAL GAZETTE © 


4 


ual 
bagi 


i 
i 


i 
aie 
ai 


age “ih i ARE ' j 


die 
Witetitarts 


j 
| 
t 
¥ 
: 
; 


Potanical Gazette 


A Monthly Fournal Embracing all Departments of Botanical Pasig 
Subscription per year, $4.00 a umbers, 40 Cents 
The subscription ea must be paid in advance. No b after . expiration 

of the time paid for. No reduction is made to dealers or afin 


FOREIGN AGENTS: 


Great Britain— Wm. WEsLEY & Son, 28 Essex Germany — GEBRUDER BORNTRAEGER, Berlin 
St., Strand, London. 18 Shillings. SW. 46, acted 17a, 18 Marks 
Vol. XXXI, No, 4 Issued April 15, 1905 


CONTENTS 


NEW OR LITTLE KNOWN NORTH AMERICAN TREES. III. Charles S. Sargen 217 
THE EFFECT OF Bibs ACID GAS UPON GRAINS AND OTHER ace 
(WITH SIX FIGURES). C. O. Townsend 241 
THE TUBER-LIKE ROOTLETS OF CY CAS REVOLUTA. CONTRIBUTIONS FROM THE 
HuLL BoranicaAL LABORATORY. XXVI (WITH TEN FIGURES). A.C. Life - - 265 
BRIEFER ARTICLES. 
WALNUT BACTERIOSIS. Mewton B. Pierce - - - - - - - - ory | 
CORRENT LITERATURE. 
BOOK REVIEWS - - - - - - - “ - in TE 
VEGETABLE PHYSIOLOGY. THE PROBLEMS OF LIFE. 
MINOR NOTICES - - - - - - - - a paar’ 2 A 
NOTES FOR STUDENTS - - - - - - - - oe 298 
OPEN LETTERS. 
THE ROCHESTER Cove. Charles Louis Pollard - - : - - - - 285 
NEWS ‘ a F Ae erp cies ¢ = i si = a? 298 


Separates, if desired, must be ordered in advance of publication. Not less than 50 separates of lead- 

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wihok: casera will also be fea dare ed at cost. The table below shows the approximate cost of separates 


consisting of plain text or with line engravings. The actual cost may vary from the figures given, 

and will depen sada upon the a nf abe ork in re-makin se: eae into forms, press work, paper, binding, 
ne Separates containing half- tones may be expected to cost somewhat more than the rates given, the 
increase Nesehebas g upon the number of cuts and the amount of sack required upon them. 

Number of copies 50 100 150 200 

Letter-press, for 4 pagesorless. . .. . $1.60 $2.00 $2.25 $2.50 

etter-press, for 8 pages or less aoe 2.25 2.75 3-15 -. 
Letter-press, for 16 pages or = eed gg 4.00 5.00 80 pip 
Single plates (1 double = 2 single) . 1.00 1.35 1.70 . 
Covers, with title (paper like GAZETTE cover). 1.50 2.00 2.50 3.00 


Manuscripts. — Contributors are requested to write scientific and proper names with ppp’ care, 
and in Sp Aa: to follow the form shown in the pages of the GAZETTE. Manuscripts shoul sent to 
he Botanical 


Books and Pamphlets for via os hould be sent to the same ad 
sen 
will be replaced free only when claim is aude within thirty days after receipt of the 
oe aoe dee d bl he order of The University of Chicago. 
Pte ss should be made payable to the order o e Uni 

espondence regarding subscriptions, advertisements, and bills rendered, should be addressed to 

The Daren of ee cat — ey. mee 
ll cond-cl il ] 


Every Botanist 


Should be familiar with 
the prominent works of 


GEBRUDER BORNTRAEGER 
Publishers. 


Die Glykoside. Chemische Monographie der Pflanzengly- 
koside nebst syst tischer Darstellung der kinstlichen 
Glykoside von Dr. I. I. L. von Ryn, Director der Reichsver- 
suchsstation in Mastricht. 8vo. Cloth, $2.50. 


Das Werk giebt—wie es bisher noch nirgends geschehen— eine eingehende 
chemische Behandlung der Glykoside—nicht nur eine kurzgefasste Zusammen- 
stellung der chemischen Eigenschaften dieser Kirperklasse, sondern die Darstet- 
lungsmethode, die Griinde, welche zur Aufstellung der Constitutionsformeln 
gefithrt haben etc., so dass das Buch in chemisch-pharmaceutischen wie pharma- 
hologischen Kreisen sowie unter den Studirenden und sonstigen Freunden der 
phytochemischen Forschung sicher mit grosser Freude begriisst werden wird. 


Die Harze und die Harzbehalter. Historisch- 
kritische und experimentelle, in Gemeinschaft mit zahlreichen 
Mitarbeitern ausgefiihrte Untersuchungen von Proressor DR. 
A. Tscuircn, Director des pharmaceutischen Institutes der Uni- 
versitat Bern. Mit 6 Tafeln. 8vo. Half calf, $5.00. 

Das Werk stellt zum ersten Mal das gesammte Material dieser wichtigen 
Gruppe von Pflanzenproducten kritisch durchgearbettet dar. Die streng Wwissen- 
schaftlichen Untersuchungen werden auch fiir die Praktiker, besonders fir adie, 
die sich mit Harzen und Harzproducten beschiftigen, von Interesse sein, da jede 
rationelle Technik ja auf wissenschaftlicher Grundlage ruht. 


Write for free Catalogue; postpaid. Address: 
Gebruder Borntraeger, Publishers, 


BERLIN, S. W. 46. 
SCHONEBERGERSTRASSE 17a 


Big Four Route 


FROM 
CHICAGO 
TO 


Indianapolis, Cincinnati, Louisville, 
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THE SCENIC LINE TO 
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CINCINNATI, O. 
J. €. TUCK C. TUCKER, G.N.A. 234 Clark St., CHICAGO 


Portraits 
of Botanists 


HE FERN BULLETIN is publishing a 
series of portraits of prominent 
botanists that is likely to interest all 
connected with the science. Another 
valuable feature is the complete set of 
the species and forms of Aguzsetum in 
North America being issued free to sub- 
There is a series of articles for 


scribers. 
beginners, and special attention is paid 
to the description of new species. 
Awarded Grand Prize at Paris. 
Sample free. Price of current num- 
ber, 15c. 


Address 


Ghe Fern Bulletin 


BINGHAMTON, N.Y. 


SoS See 


[JUST PUBLISHED] 


Being 
No, 137-138 of 


PISS SSS 


28 Essex Street, Strand, - 


Transactions of Scientific Societies 


Send 12 cents, U. S. stamps, for 


- Wm. Wesley & Son’s Botanical Catalogue, 1901 J 


CONTENTS: 
Periodicals 


THE f Bibliography History ~ oe = Portraits 
NATURAL f nee o Early Botanical Science Linnaeus 
eatery 7 Microscopy Morphology aad Physiology 
BOOK f Encyclopaedic works Classification | Nomenclature 
CIRCULAR 4 Cryptogams Phanerogams Fossil Plants 
Natural distribution of Plants (Floras) 
BOTANY J Agriculture and Horticulture to the end of the J8th century 
More than 3300 works, J Gardening Landscape Gardening 
classified under 42 f The Flower and Ornamental Garden 
headings Husbandry Tropical Agriculture Commercial Plants 
dt deo Medical Botany Forestry Diseases of Plants 


William Wesley 8& Son, Booksellers, 


: London, England. 


|S CORO CERN GN TE ora Ean oe 


2 RSIS © RR Oy 


Pe UNIVERSITY OF CHICAGO 


FOUNDED BY JOHN D. ROCKEFELLER 


THE SUMMER QUARTER 
IQOI 


The Circular of Information 


For the Summer Quarter of 1901 at The University of Chicago 
presents a program full of interest for those who desire to 
spend some part or all of the summer months in study. 


Change of Dates 


During the coming summer, in response to many suggestions, 
the arrangement of the terms will be materially changed. AY e 
quarter will- begin on Wednesday, June 19, the two terms being 
divided as follows: first term, Wednesday, June 19, to Friday, 
July 26; second term, Saturday, July 27, to Saturday, August 31 


Departments Offering Courses 


Instruction will be given in twenty-eight distinct departments 
in the University, and also in the Divinity School. In these 
various departments a large number of distinct courses are 
announced, thus affording a very wide range of subjects. 


The Staff 


A large number of teachers and lecturers will be in residence, 
including not only one hundred and twenty-five of the regular 
faculty of thé University, but also representatives of a number 
of other institutions in America and Europe. 


University Open Lectures 


The program of The University Open Lectures, to which per 
sons who are not regularly matriculated in the University wil 
be admitted on moderate terms, will be sent on application. 


THE UNIVERSITY OF CHICAGO 


CHICAGO, ILLINOIS 


PICTURESQUE 
RO i 


UT 
TO PAN. AMERICAN 


NEA. MEETING] || SERS DON 


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The Wabash, the “Niagara Falls 
JULY 8 to JULY 12 Short Line,” has made special ar- 
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AND THE en 
during the Exposition. New equip- 
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Pan-Am erican Expositi on be provided. The Wabash is the 
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MAY to NOVEMBER, 1901 stop-over at Niagara Falls, 
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Che Elementary School Record 


A SERIES OF MONOGRAPHS ON THE EXPERIMENTAL SCHOOL OF 
THE PEDAGOGICAL DEPARTMENT OF THE UNIVERSITY OF CHICAGO 


Edited by JoHN DEweEy and LAuRA L. RUNYON 


HE object of the Elementary School Record is to make possible for use in 

other schools the details of subject-matter and method in the application 

of modern psy chology in education, as demonstrated by the University Ele- 

mentary School. The plan includes a series of nine monographs, each number 

containing a record of work done by a group in the school, and also an article 

concerning the work of one department in all grades. The complete series is 
d 


now ready. 
EACH NUMBER OF Rlnag SERIES IS a ee he hasan IN PAPER. 
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The Journals of the University of Chicago 
BEING THE DESCRIPTIVE LIST OF ONE WEEKLY, FOUR 
MONTHLY, ONE BI-MONTHLY, THREE QUARTERLY, 
AND ONE SEMI-QUARTERLY PUBLICATIONS & &# # 


THE BIBLICAL WORLD 


eager by President W te Harper, A popular rar eto 
nthly magazine. Sub scription price ¢ iit the Uni iolpiig 
$2.00 = fore foreign, $2.50; single copies, 20 cen 
The Biblical eden is devoted sia i to bibli- 
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he greatest aid to the busy cler sree ie pro 
ca ssive Sunday- bras teacher, and the aakiue 
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THE SCHOOL REVIEW 
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ar edikation in the Un via ee It is 
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THE BOTANICAL GAZETTE 
Edited by Joun M. Coutrer. Published monthly, with illus- 
ations, Sabscxiption Lise $4.00 a F gen = the United 
States; foreign, $4.50; single copie 
Th tanical Gazette is 7 illus trated 5 seatkly 
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more than tw enty _— it has be een the representa- 
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Europe. 


THE BS JOURNAL OF SOCIOLOGY 
Edited by ALB Smatt, Published pel Beegt aah with 
oo oy Subscription price, $2.00 a year in the United 
States; foreign, $2.50; single nts, 
The special aim of the American Journal of Soci- 
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many problems and one problem. fe has already 


copies, 35 Cae 


made itself apa tear to A re 
trying to keep informed about the venievat tenden- 
cies in the rapidly changing field of sociology. 


THE AMERICAN JOURNAL OF SEMITIC 
LANGUAGES AND LITERATURES 
Edited by Fagg a8 Wittiam R, Harper, of The Uni niversity of 
Chicago. ished quarterly, ome price, $3.00 
a yea ear; fore et hg $3.25; single copies, 75 ce 
The object of this journal is to encourage the 
study of the Semitic languages and literatures, to 
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THE JOURNAL OF GEOLOGY 


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ublished semi- Peery 5! bite 


Devoted to the interests of geology seas - allied 
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The ony journa al in eth world so catholic in its 

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The University of Chicago. It co tains 


aE PROSPECTS 


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By WILLIAM R. HARPER 


ECAST from an address deliv- 
ered before the National Edu- 
ional Association, at Charles- 


a 
ton, South Carolina, July 10, 1900. 
The substance of the book was also 


as a course of lectures at The 


if 
University ~ Chicago, during the 
past summe 


Now Ready, J2mo, paper, 25c (postpaid), 


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VOLUME XXXI . [) NUMBER 4 


DOTANICAL: G4AZETIE 
APRIL, rgor 


NEW OR LITTLE KNOWN NORTH AMERICAN 
‘LREIGES, <1: 
CHARLES S.-SARGEN'T. 

’ Crataegus acutifolia, n. sp.—Glabrous. Leaves oval to 
oblong-obovate, acute or acuminate, or rarely rounded at the 
apex, cuneate at the base, finely crenulate-serrate with gland- 
tipped teeth except toward the base, or occasionally only above 
the middle, on vigorous leading shoots sometimes irregularly 
divided toward the apex into two or three pairs of short acute 
lobes; nearly fully grown when the flowers open and then thin 
and lustrous above and at maturity thin but firm in texture, dark 
green and very lustrous on the upper surface, pale yellow- 
green on the lower surface, about 1% in. long and I in. wide, 
and on leading shoots sometimes 2% to 3 inches long and 
nearly 2 in. wide, with slender light yellow midribs compara- 
tively deeply impressed above, and four or five pairs of thin pri- 
mary veins without the parenchyma; petioles slender, winged 
above by the decurrent bases of the leaf-blades, deeply grooved, 
glandular with minute dark caducous glands, from % to ¥ in, in 
length ; stipules linear, elongated, dark red, caducous. Flowers 
% in. in diameter on slender pedicels, in compound many-flowered 
compact corymbs; bracts and bractlets linear, glandular-serrate, 
caducous; calyx-tube narrowly obconic, the lobes lanceolate, 
acuminate, entire or occasionally obscurely glandular-serrate ; | 
stamens 10; filaments slender, elongated; anthers small; styles 

if 217 


218 BOTANICAL GAZETTE [APRIL 


2 or 3. Fruit in drooping few-fruited clusters, oblong, full 
and rounded at the ends, bright scarlet, marked by few large 
dark lenticels, about % in. long; calyx-tube prominent with 
a broad deep cavity, the lobes closely appressed, often decidu- 
ous before the maturity of the fruit; flesh thin, yellow, dry and 
mealy; nutlets 2 or 3, broad, prominently ridged on the back 
with broad rounded ridges, about ,%, in. long. 

A tree 25 to 30 ft. in height with a trunk 8 or Io in. in 
diameter, and wide-spreading branches forming a broad sym- 
metrical rather flat-topped head and slender nearly straight 
branchlets marked by oblong pale lenticels, dark chestnut-brown 
or orange-brown and lustrous during their first year, becoming 
dull gray-brown during their second season, and slightly armed 
with few slender straight chestnut-brown spines from I to nearly 
2 in. in length. 

Flowers early in May. Fruit ripens toward the end of 
September. 

Open oak woods near the bank of the Mississippi river at 
Carondelet in South St. Louis, Missouri. First collected in May 
1887 by H. Eggert, and in September 1900 by Eggert, Norton, and 
Sargent. 

This is one of several forms which have been usually confounded with 
Crataegus Crus-galli of Linnaeus, but which can probably best be separated 
from that species. From Crataegus Crus-galli as limited by Linnaeus and 
Aiton, and now universally recognized, it differs in its thin usually acute or 
acuminate leaves with comparatively prominent veins, in its smaller flowers, 
much smaller early-ripening scarlet fruit, and in its more slender branchlets 
only sparingly armed with much more slender spines. 

/ Crataegus erecta, n. sp.—Glabrous. Leaves broadly oval to 
obovate, or on leading shoots nearly orbicular, acute and short- 
pointed at the apex, cuneate and entire at the base, finely crenu- 
late-serrate above, or on vigorous shoots coarsely dentate with 
broad nearly straight gland-tipped teeth, thin, dull green and 
nearly fully grown when the flowers open, and at maturity thin 
but firm in texture, dark dull green on the upper surface, pale on 
the lower surface, 1% to 2 in. long, 1 to 1% in. wide, and on 
leading shoots sometimes 3 in. long and 2% in.. wide, with 


~ — e— 


igor} NORTH AMERICAN TREES 219 


slender midribs and thin prominent primary veins; petioles 
slender, deeply grooved, wing-margined above particularly on 
vigorous shoots, glandular with minute dark glands, often dark 
red after midsummer, from 4% to ¥% in. long; stipules linear, 
glandular-serrate, fading red, % in. long, caducous. Flowers 
% to % in. in diameter, on slender elongated pedicels, in broad 
loose many-flowered very thin-branched compound corymbs; 
bracts and bractlets linear, glandular-serrate, caducous ; calyx- 
tube narrowly obconic, the lobes narrow, acuminate, elongated, 
entire or occasionally obscurely dentate; stamens usually 10, 
occasionally 11 to 13; filaments slender; anthers small, pale 
yellow; styles 3 or 4, surrounded at the base by a narrow ring 
of short pale hairs. Fruit in few-fruited drooping clusters, sub- 
globose, usually a little longer than broad, full but flattened at 
the ends, dark dull crimson, marked by occasional large dark 
lenticels, from \% to ly in. in length; calyx-tube very short 
with a broad shallow cavity, the lobes gradually narrowed from 
broad bases, closely appressed, usually persistent on the mature 
fruit ; flesh thin, yellow, dry and mealy; nutlets 3 or 4, broad, 
prominently doubly ridged on the back, about 3% in. long. 

A tree usually 25 to 30 ft. in height with a trunk a foot in 
diameter covered with dark gray-brown or nearly black bark 
broken irregularly into thick plate-like scales, ascending branches 
forming a broad open erect head and slender slightly zigzag 
branchlets marked by numerous large oblong pale lenticels, 
green more or less tinged with red when they first appear, 
orange or reddish-brown during their first season and gray or 
gray-brown during their second year, and armed with numerous 
straight slender chestnut-brown spines from I to 2 in. in length. 

Flowers about May. fo. Fruit ripens toward the end of 
September. The leaves before falling turn dull orange-color. 

Rich bottom-lands of the Mississippi river in Illinois opposite 
St. Louis. First noticed October 5, 1899, in a vacant lot in the 
city of East St. Louis by Eggert, Norton, and Sargent, and on 
May 12, 1900, collected in flower by J. B. S. Norton, and in 
October 1g00 with ripe fruit by Eggert, Norton, and Sargent. 


220 BOTANICAL GAZETTE [APRIL 


This is another of the Crus-galii group, differing from the species of 
Linnaeus by its thin pointed or suborbicular leaves with prominent veins, which 
even on vigorous shoots show no tendency to lobing, by its pale yellow not 
rose-colored anthers, small nearly globose bright scarlet fruit, by its ascend- 
ing not wide-spreading branches, and by its less numerous spines standing 
out from all sides of the branchlets, as they do usually in Crataegus with the 
exception of Crataegus Crus-galli. This species in its typical form can always 
be recognized by the direction of the spines which point downward from the 
branch in two ranks. 

Growing in a field in the Mississippi bottom near Fish lake in the village 
of Cahokia about four miles below East St. Louis there is a specimen of 
Crataegus erecta forty feet in height with a trunk three feet in diameter three 
feet above the surface of the ground and a broad head of numerous large 
upright branches. 

/ Crataegus Lettermani, n. sp—Leaves obovate to broadly 
oval on leading shoots, acute, acuminate or rounded and short- 
pointed at the apex, gradually narrowed from near the middle 
to the base and decurrent on the stout elongated glandular 
tomentose but ultimately nearly glabrous petioles, often slightly 
and irregularly divided above into three or four pairs of short 
acute lobes, coarsely doubly serrate, often nearly to the base, 
with glandular incurved or straight teeth; as they unfold 
strongly plicate, tomentose above and below like the young 
branchlets, with a thick coat of hoary tomenttim, and when 
the flowers open covered on the upper surface with short soft 
hairs and villose on the lower surface, and at maturity thin but 
firm in texture, scabrate and dark green above, pale below, 
from 2 to 2% in. long and from 1 to 1% in. wide, with slender 
midribs and primary veins impressed on the upper surface and 
puberulous below, and conspicuous forked secondary veins and 
reticulate veinlets. Flowers about 34 in. in diameter in compact 
many-flowered thick-branched tomentose cymes; bracts and 
bractlets linear, glandular-serrate, caducous ; calyx-tube narrowly 
obconic, coated with thick hoary tomentum, the lobes narrow, 
acuminate, villose, finely glandular-serrate, reflexed after anthe- 
sis; stamens 10; filaments slender, elongated; anthers small; 
styles 5, surrounded at the base by a broad ring of white 
tomentum. Fruit nearly globose and somewhat flattened at the 


— 


1901] NORTH AMERICAN TREES 221 


ends, dark red, marked by a few large pale lenticels, 54 in. in 
diameter; calyx-cavity broad and shallow, the lobes serrate, 
villose, spreading and closely appressed; flesh thin and yellow ; 
nutlets 5, thick, very prominently doubly ridged on the back, 
about ¥% in. long. 

A tree from 15 to 18 ft. in height, with a trunk 6 or 8 in. 
in diameter, and stout wide-spreading branches forming a sym- 
metrical round-topped head and slender branchlets dull red and 
puberulous during their first season and dark gray-brown during 
their second year, and ‘armed with stout straight spines from 
1% to 2 in. in length. 

Flowers in May. Fruit ripens late in October. 

Rich woods, Allenton and Pacific, Missouri, George W. Letter- 
man, May and October 1882. 

This species, which was referred by Engelmann to Crataegus tomentosa 
Linnaeus, and to Crataegus punctata Jacquin, is still very imperfectly known. 
Its relationship appears to be with Crataegus collina Chapman, from which it 
differs in its more tomentose young leaves and branchlets, in its short thick- 
branched and more tomentose corymbs, ten stamens, rather larger fruit, and 
in its much later flowers. 

v Crataegus Arnoldiana, n. sp.— Leaves broadly ovate or rarely 
oval, acute at the apex, rounded, truncate or occasionally broadly 
cuneate at the base, irregularly divided above the middle into 
numerous short acute lobes, coarsely doubly glandular-serrate 
except at the base; in early spring densely villose above and 
below, and at maturity from 2 to 3 in. long and broad, membra- 
naceous, smooth, dark green and lustrous on the upper surface, 
paler on the lower surface, slightly villose on the under side 
of the slender midribs and thin remote primary veins running 
to the points of the lobes and faintly impressed above; petioles 
Slender, only slightly grooved, densely villose, ultimately 
puberulous, from 34 to 1% in. long; stipules linear, coarsely 
glandular-serrate, often I in. long, caducous. Flowers % in. 
in diameter on slender pedicels in broad loose compound 
many-flowered thin-branched tomentose cymes; bracts and 
bractlets lanceolate to oblanceolate, coarsely glandular-serrate ; 


me 
7 
‘ : 
é 


222 BOTANICAL GALETTE [ APRIL 


calyx-tube broadly obconic, densely tomentose, the lobes nar- 
row, elongated, acuminate, glandular-serrate, villose on both 
surfaces, reflexed after anthesis; stamens 10; filaments slender ; 
anthers large, pale yellow; styles 3-5, usually 3 or 4, sur- 
rounded at the base by a broad ring of thick white tomentum. 
Fruit on slender pedicels in erect, spreading or sometimes droop- 
ing few-fruited slightly villose clusters, subglobose but rather 
longer than broad, bright crimson, marked by numerous large 
pale lenticels, villose particularly toward the ends with long 
white scattered hairs, 34 in. long; calyx-cavity broad and 
shallow, the lobes coarsely glandular-serrate, villose, wide- 
spreading, often deciduous before the falling of the fruit; flesh 
thick, bright yellow, subacid; nutlets usually 3 or 4, thick, light- 
colored, prominently ridged on the back with high rounded 
ridges, about % in. long. 

A tree 15 to 20 ft. in height with a short trunk 10 or [2 in. 
in diameter, stout ascending branches forming a broad open 
irregular head, and slender strongly zigzag branchlets marked 
by large oblong pale lenticels, coated when they first appear 
with long matted hairs, becoming dark orange-brown and lus- 
trous before midsummer and glabrous or puberulous during 
their first winter, light orange-brown and lustrous during their 
second season, and finally ashy-gray, and armed with very 
numerous stout straight or slightly curved bright chestnut-brown — 
lustrous spines from 2% to 3 in. long and brilliant for four or five 
years. Winter buds oblong, gradually narrowed to the obtuse 
apex, bright red and lustrous, about .3, in. long. 

Flowers during the last week of May. Fruit ripens by the 
middle of August and falls before the 1st of September. 

Known in a wild state only in a small group of plants 
growing on a wooded bank in the Arnold Arboretum, but now 
frequently cultivated in the neighborhood of Boston, and in cul- 
tivation forming a tall trunk and promising to attain a large size. 

When it is covered with its brilliant and abundant fruit Crataegus A rnol- 
diana is one of the most beautiful of the thorns which ripen their fruit in 
summer or early autumn. From Crataegus submollis (Sargent, Bor. GAZ. 


1901] NORTH AMERICAN TREES 223 


31:7. 1901) of the same region it differs in its broader darker green more 
villose leaves which are usually rounded, not cuneate, at the base, in its 
smaller flowers, subglobose, not oblong or pear-shaped, crimson fruit with 
smaller spreading calyx-lobes, borne on shorter peduncles and ripening two 
or three weeks earlier, and by its much more zigzag and more spiny branches 
which make this tree particularly noticeable in winter when it may be readily 
recognized from all other thorn trees. From Crataegus Champlainensts 
(Sargent, Rhodora 3: 20), another of the eastern species of the mo//is group 
with ro stamens, it differs in its larger thinner leaves which are yellow-green, 
not blue-green, in its smaller subglobose, not oblong or obovate, early- 
ripening fruit, the fruit of Crataegus Champlainensis beginning to ripen early 
in September and remaining on the branches until midwinter. It differs also 
from this northern species in the form of the fruiting calyx and in its much 
more zigzag branchlets. 

V Crataegus Arkansana, n. sp.— Leaves oval to oblong-ovate, 
acute at the apex, broadly cuneate, or on vigorous leading shoots 
occasionally rounded at the base, usually divided above the 
middle into three or four pairs of short broad acute lobes, 
sharply serrate, sometimes to the very base, with spreading 
gland-tipped teeth; in early spring coated with short soft pale 
hairs, particularly on the under surface of the midribs and veins, 
and at maturity thick and leathery, dark dull green and glabrous 
on the upper surface, pale yellow-green on the lower surface, 
from 2 to 3 in. long, from 134 to 2 in. wide, or on vigorous 
shoots often 4 in. long and 3 in. broad, with stout light yellow 
midribs and primary veins deeply impressed above and slightly 
villose below with scattered pale hairs, conspicuous secondary 
veins and reticulate veinlets; petioles stout, deeply grooved, 
more or less winged above, glandular with minute usually decid- 
uous dark glands, tomentose, ultimately glabrous or puberulous, 
usually dull red in the autumn, from I to 1% in. long; stipules 
glandular-serrate, villose, linear-lanceolate to linear-obovate, 
about % in. in length. Flowers 1 in. in diameter in broad com- 
pound many-flowered thin-branched villose corymbs ; bracts and 
bractlets oblong-ovate, acuminate, finely glandular- serrate ; 
calyx-tube narrowly obconic, coated with long matted pale hairs, 
the lobes short, acute, very coarsely glandular-serrate, glabrous 
or slightly villose; stamens 20; filaments slender ; anthers large, 


224 BOTANICAL GAZETTE [APRIL 


pale yellow; styles 5. Fruit in few-fruited clusters, oblong or 
rarely obovate, full and rounded at the ends, bright crimson, 
lustrous, marked by few large dark lenticels, slightly tomentose 
at the ends, particularly at the apex, from 34 to I in. long, 34 
in. thick; calyx-cavity deep but comparatively narrow, the lobes 
small, linear-lanceoiate, coarsely glandular-serrate, red at the 
base on the upper surface, erect and persistent ; flesh yellow, 
thick, dry, subacid; nutlets small in comparison with the size of 
the fruit, thin, rounded or slightly and irregularly ridged on the 
back, % in. long. 

A tree from I5 to 20 ft. in height with a tall straight stem 
covered with pale scaly bark, thick ascending branches and stout 
slightly zigzag branchlets marked by large oblong pale lenticels, 
dark green and covered with long scattered pale hairs when they 
first appear, light orange-brown and very lustrous during their 
first winter, becoming ashy-gray during their second year, and 
unarmed or armed with occasional straight bright chestnut- 
brown spines gradually narrowed from broad bases and usually 
from % to % in. long. Winter-buds about % in. in length, 
nearly as broad as long, dark red, puberulous along the margins 
of the scales. 

Flowers at the Arnold Arboretum about the middle of May. 
Fruit ripens at the end of October and remains on the branches 
for several weeks longer, falling gradually. Late in October or 
early in November the leaves turn bright clear yellow. 

Apparently common in southern Arkansas (B. F. Bush 953); 
but first distinguished from plants in the Arnold Arboretum 
raised from seeds collected in 1883 by George W. Letterman at 
Newport, Ark. 

From Crataegus mollis Scheele, with which it has been confounded, Cra- 
taegus Arkansana differs in the form and particularly in the leathery texture 
of the leaves which when young are villose, not tomentose, in its villose 
corymbs, its oblong late-ripening fruit, the fruit of Crataegus mollis falling 
from the middle of August to the middle of September, and in its unarmed 
or only slightly armed branchlets. Perfectly hardy in the Arnold Arboretum, 
Crataegus Arkansana is unsurpassed late in the autumn in the beauty of its 
large brilliant and abundant fruits which make it one of the most desirable 
garden plants of the genus. : 


1901 | NORTH AMERICAN TREES 225 


Crataegus Texana Buckley, Proc. Phil. Acad. 1861: 454.— 
Leaves broadly ovate, acute or rarely rounded at the apex, 
cuneate or on leading shoots truncate or slightly cordate at the 
base, usually divided above the middle into four or five pairs of 
broad acute lobes, very coarsely and doubly glandular-serrate 
except toward the base ; when they first unfold covered above with 
short soft pale hairs and below with a thick coat of hoary tomen- 
tum, and at maturity thick and firm in texture, dark green and 
lustrous on the upper surface, pale and pubescent or tomentose 
on the lower surface particularly along the stout light-colored 
midribs and primary veins, and on the conspicuous secondary 
veins and reticulate veinlets, from 3 to 4 in. long, from 2% to 3 
in. wide; petioles stout, more or less winged above, deeply 
grooved, tomentose, ultimately nearly glabrous, from % to % 
in. long; stipules lunate, apiculate, often stipitate, coarsely ser- 
rate, from 4 to % in. long. Flowers 34 in. in diameter on 
| elongated slender pedicels in broad loose many-flowered tomen- 
} tose cymes; bracts and bractlets oblong to oblong-ovate, broad, 

acuminate, villose, sometimes % in. long, very conspicuous ; 
calyx-tube broadly obconic, coated with thick pale tomentum, 
the lobes gradually rounded from broad bases, acuminate, very 
coarsely glandular-serrate, villose with long matted hairs, reflexed 
after anthesis ; stamens 20; filaments'slender ; anthers large, dark 
red; styles 5. Fruit in broad drooping many-fruited tomentose 
ultimately nearly glabrous clusters, short-oblong or somewhat 
obovate, full and rounded at the ends, tomentose until nearly 
fully grown, at maturity bright scarlet, lustrous, marked by 
occasional large pale lenticels, puberulous toward the apex, from 
3% to 1 in. in length; calyx-cavity broad and deep, the lobes 
much enlarged, glandular-serrate, villose, dark red at the base 
on the upper side, usually erect, often deciduous ; flesh thick, 
yellow, sweet and edible; nutlets 5, thick, slightly grooved on 
, the back, from % to ¥% in. long. 
A tree often 30 ft. in height with a tall trunk sometimes a 
foot in diameter covered with dark closely appressed scales, 
Stout spreading branches forming a symmetrical round-topped 


226 BOTANICAL GAZETTE | APRIL 


head, or on young trees upright and forming an open irregular 
crown, and comparatively slender slightly zigzag branchlets, 
dark bronze-green and villose when they first appear, soon 
becoming dull reddish-brown, lighter reddish-brown in_ their 
second season, and ultimately pale ashy-gray, and often unarmed, 
or armed with occasional slender nearly straight bright chestnut- 
brown spines usually about 2 in. in length. 

Flowers the middle of March. Fruit ripens toward the end 
of October. 

Rich bottom-lands, central and western Texas. 

Long confounded with Crataegus mollis Scheele (see Gray, Proc. Phil. 
Acad. 1867: 163), it can be distinguished from that species by the shape of 
the thinner leaves which are nearly always cuneate and only rarely cordate 
at the base even on the most vigorous shoots, and are usually less deeply 
lobed and much more coarsely serrate, by the smaller flowers in fewer- 
flowered more tomentose cymes, by the late-ripening fruit, and by the color 
of the branchlets and their more numerous spines. Crataegus mollis appar- 
ently does not extend south of central Missouri and middle Tennessee, being 
replaced in southern Missouri and Arkansas by several forms of the Mollis 
group which are still imperfectly known. 

g Crataegus pedicellata, n. sp.— Leaves broadly ovate, oval, or 
occasionally obvate or rhomboidal, acute or acuminate, broadly 
cuneate or rounded, and on vigorous leading shoots occasionally 
truncate or slightly cordate at the base, divided above the middle 
into four or five pairs of short acuminate lobes, coarsely and 
often doubly serrate, except toward the base, with spreading 
glandular teeth; in early spring roughened above by short rigid 
pale hairs and at maturity membranaceous, dark rich green and 
scabrate on the upper surface, pale and glaucous below, from 3 
to 4 in. long, from 2 to 3 in wide, with slender midribs slightly 
impressed above and thin remote primary veins arching to the 
points of the lobes; petioles slender, only slightly grooved, vil- 
lose but ultimately glabrous, obscurely glandular with minute scat- 
tered dark glands, from 1 ¥% to 2in. in length; stipules of leading 
shoots strongly falcate, stipitate, coarsely glandular-serrate, y 
in. long. Flowers % in. in diameter on slender-elongated 
pedicels, in loose lax rather few-flowered slender-branched 


1901] NORTH AMERICAN TREES 227 


slightly villose corymbs ; bracts and bractlets laciniate, glandular- 
serrate, caducous; calyx-tube narrowly obconic, glabrous, the 
lobes broad, acute, very coarsely glandular-serrate reflexed 
after anthesis ; stamens 10; filaments slender, elongated ; anthers 
rose color; styles 5, surrounded at the base by a conspicuous 
ring of pale tomentum. Fruit in few-fruited drooping glabrous 
clusters, oblong, full and rounded at the ends, bright scarlet, 
lustrous, marked by numerous small dark lenticels 34 in. long, 
from to % in. thick; calyx-cavity broad and deep, the lobes 
much enlarged, coarsely serrate, usually erect and incurved ; 
flesh thin and pale; nutlets 5, rounded and deeply grooved on 
the back, % in. long. 

A tree from 18 to 20 ft. in height with a stout trunk a foot 
in diameter and short ascending branches forming a broad open 
shapely head, and rather slender slightly zigzag branchlets 
marked by numerous small pale lenticels, dark chestnut-brown and 
slightly villose when they first appear, becoming bright chest- 
nut-brown and very lustrous during their first season and ashy- 
gray during their second year, and armed with few stout straight 
or slightly curved chestnut-brown lustrous spines from 1% to 2 
in. long. Winter-buds nearly globose, bright red, very lustrous, 
¥% in. in diameter. 

Flowers during the last week of May. Fruit ripens and 
mostly falls during the second half of September. 

Rochester, N. Y., C. C. Laney and John Dunbar, 1899 and 
1900. 

This handsome tree is most conveniently placed in the F/ade//atae group 
with Crataegus Holmesiana Ashe. From that species it differs in its larger 
darker green and more scabrous mature leaves, in its more lax villose 
corymbs, larger flowers on longer pedicels, with coarsely glandular-serrate 
calyx-lobes, more numerous stamens and styles, and in its larger and later- 
ripening fruit. 

~ Crataegus lucorum, n. sp.— Leaves broadly ovate to obovate 
or rarely oval, acute or acuminate, gradually narrowed and full and 
rounded or broadly cuneate at the base, deeply divided above the 
middle into three or four pairs of broad acute or acuminate lobes, 


228 BOTANICAL GAZETTE [APRIL 


dentate, except at the very base, with straight glandular teeth; 
when the flowers open not more than one third grown, bronze 
color, covered above with short soft pale hairs, glabrous below, 
and at maturity membranaceous, smooth dull dark green and 
glabrous on the upper surface, pale yellow-green on the lower 
surface, from 1% to 2 in. long and broad, with slender yellow 
midribs slightly impressed above and thin primary veins run- 
ning obliquely to the points of the lobes; petioles very slender, 
often short-winged above, nearly terete below, glandular, from 
I to 1% in. in length; stipules lanceolate to oblanceolate, finely 
glandular-serrate, from 4 to % in. in length, caducous. Flowers 
34 in. in diameter on slender pedicels in narrow compact few- 
flowered thin -branched slightly villose corymbs ; bracts and bract- 
lets linear-lanceolate, glandular, caducous; calyx-tube broadly 
obconic, glabrous, the lobes narrow, acuminate, conspicuously 
glandular-serrate, villose on the -upper surface, reflexed after 
anthesis; stamens 20; filaments slender; anthers small, dark 
purple; styles 4 or 5. Fruit in erect few-fruited slightly vil- 
lose clusters, pear-shaped until nearly fully grown, but at matu- 
rity oblong or somewhat obovate, full and rounded at the ends, 
crimson, lustrous, marked by small pale lenticels, from y% to 5% 
in. long; calyx-cavity deep but narrow, the lobes elongated, 
glandular-serrate, villose above, spreading and closely appressed, 
often deciduous before the maturity of the fruit; flesh thick, 
yellow, dry, and mealy; nutlets 4 or 5, thin, rounded, and 
obscurely or not all ridged on the back from ¥% to 5% in. in length. 

A tree from 20 to 25 ft. in height, with a tall trunk from 6 
to 8 in. in diameter covered with close dark red-brown bark, 
slender erect branches forming a narrow head and thin slightly 
zigzag branchlets marked by numerous oblong pale lenticels, 
dark green and slightly villose when they first appear, dull 
orange-brown in their first season, ultimately dark gray-brown, 
and armed with occasional straight or slightly curved bright red- 
brown spines from 1 to 1% in. in length. 

Flowers during the first week of May. Fruit ripens about 
the middle of September and soon falls. 


: 
Se ee eee 


See 


1901] NORTH AMERICAN TREES 229 


Margins of oak groves in rich moist soil along the banks of 
sloughs near Barrington, Illinois, £. 7. AHid/, May and June 1899; 
fill and Sargent, September 1900. 

Distinguished from the other species in the 7enudfoliae group now recog- 
nized by it more numerous stamens. 

“ Crataegus Columbiana, n. sp.— Leaves oval to obovate, or on 
vigorous shoots often broadly ovate to oblong-oval, usually 
acute, or occasionally rounded at the apex, full and rounded or 


_ gradually narrowed or abruptly cuneate at the base, irregularly 


doubly glandular-serrate, except toward the base, with broad 
spreading teeth; on leading shoots sometimes slightly divided 
above the middle into three or four pairs of short acute lobes; 
when they unfold conspicuously plicate, covered above with long 
soft white hairs and below with a thick coat of snow-white 
tomentum, and at maturity thin but firm in texture, dark green, 
lustrous and scabrate on the upper surface, paler and pubescent or 
tomentose on the lower surface, from 2 to 2% in. long and broad, 
and on leading shoots often from 2 to 4 in. long and broad, with 
slender midribs, few thin primary veins and conspicuous reticulate 
veinlets; petioles stout, tomentose, about % in. long; stipules 
lunate, often apiculate, more or less stipitate, often coarsely glan- 
dular serrate, villose, from Y% to % in. long. Flower 3 in. in 
diameter on_ slender elongated pedicels covered with hoary 
tomentum like the slender branches of the broad lax many- 
flowered compound corymbs; bractsand bractlets oblong-obovate, 
acute or rounded and apiculate at the apex, finely glandular- 
serrate, slightly villose; calyx-tube narrow, obconic, coated with 
thick hoary tomentum, the lobes short, acute, coarsely glandular- 
serrate, tomentose on both surfaces; stamens 20, filament 
slender, elongated; anthers small, dark red; styles 5, sur- 
rounded at the base by long tufts of snow white hairs. Fruit in 
few-fruited spreading tomentose clusters, subglobose, often rather 
longer than broad, full and rounded at the ends, tomentose until 
nearly grown, dark red, marked by numerous large pale dots, 
glabrous at maturity, from 3 to nearly % in. in diameter; 
calyx-tube prominent, with a broad deep cavity, the lobes short, 


230 BOTANICAL GAZETTE [APRIL 


spreading, usually deciduous before the ripening of the fruit ; 
flesh thin, light-colored, hard, and dry, shrivelling on the 
branches; nutlets 5, rounded and usually ridged on the back, 
about ¥ in. long. 

A tree 25 ft. in height, with a tall trunk from 6 to 8 in. in 
diameter covered with pale closely appressed scales becoming 
dark brown near the base of old individuals, and numerous 
upright branches often forming a broad symmetrical head, and 
slender slightly zigzag branchlets coated when they first appear 
with hoary deciduous tomentum, light reddish-brown and more 
or less villose during their first season, becoming rather darker 
during their second year and ultimately pale ashy-gray, and 
armed with numerous stout straight or slightly curved chestnut- 
brown lustrous spines usually from 1 to 1% in. in length. 

Flowers toward the end of March. Fruit ripens after the 
middle of October and sometimes does not entirely fall until the 
following spring. 

Sandy bottom-lands of the Brazos river, usually in open 
forests of live oaks at Columbia, Texas, 5. F. Bush, November 
1899; Canby, Bush, and Sargent, March 1900; Bush, April and 
October 1goo. 

~ Crataegus Berlandieri, n. sp—-Leaves oblong-obovate to 
oval, acute or acuminate at the apex, gradually narrowed below 
from near the middle and cuneate and entire at the base, irregu- 
larly divided into numerous acute or acuminate or, on vigorous 
shoots, rounded lobes, coarsely and often doubly serrate with 
broad spreading orincurved gland-tipped teeth; at the flowering 
time coated above with short pale soft caducous hairs and below 
with thick hoary tomentum, and at maturity rather thin but firm 
in texture, glabrous, dark green and very lustrous on the upper 
surface, pale and pubescent below, usually about 3 in. long and 2 
in. wide, and on leading shoots often 4 or 5 in. long and from 
2% to 2 in. wide, with slender midribs and remote primary veins 
slightly impressed above and conspicuous secondary veins and 
reticulate veinlets; petioles stout, usually more or less winged 
above, tomentose, ultimately pubescent, from %4 to % in. long; 


1901] NORTH AMERICAN TREES 231 


stipules villose, falcate or on vigorous shoots lunate, coarsely 
glandular-serrate and frequently % in. in length. Flowers 34 
in, in diameter on stout elongated pedicels coated with hoary 
tomentum like the stout branches of the broad compound many- 
flowered cymes becoming lax after anthesis ; bracts and bractlets 
oblong-obovate to lanceolate, finely glandular-serrate, conspicu- 
ous; calyx-tube broadly obconic, tomentose, the lobes broad, 
acute, very coarsely glandular-serrate, villose on both surfaces ; 
stamens 20, filaments slender, elongated; anthers small; styles 
5, slender, surrounded at the base by small tufts of snowy white 
hairs. Fruit in lax drooping clusters, short-oblong to sub- 
globose, scarlet, about ¥y% in.long; calyx-cavity broad and deep, 
the lobes enlarged, coarsely serrate ; flesh thin, yellow, dry, and 
hard; nutlets 5, rounded but not ridged, and occasionally 
obscurely grooved on the back, about ¥ in. long. 

A tree 15 to 20 ft. in height, with spreading branches 
forming a broad open head, and slender slightly zigzag branch- 
lets coated when they first appear with hoary tomentum, becom- 
ing puberulous and dull reddish-brown during the summer and 
pale gray-brown and glabrous during their second year, and 
nearly unarmed or furnished with occasional straight spines I in. 
in length. 

Flowers from the middle to the end of March. Fruit ripens 
after the middle of October. 

“De Bejur a Austin, Avril 1828,” and “Villa d’Austin frio 
de los Brazos, Maio 1828,” /. L. Berlandier, nos. 356 and 267 in 
Herb. Gray; near the banks of the Brazos river at Columbia, 
Texas, B F. Bush, October 1899, March, April, and October 
1900. 

’ Crataegus nitida, n. sp. (Crataegus viridis var. nitida Britton 
& Brown, U7. Fl. 2: 242. 1897).—Glabrous with the exception 
of a few scattered pale caducous hairs on the upper side of the 
midribs of the unfolding leaves. Leaves lanceolate to oblong- 
obovate, acuminate at the apex, abruptly or gradually narrowed 
and cuneate at the entire base, coarsely glandular-serrate above 
with straight or incurved glandular teeth, more or less divided, 


232 BOTANICAL GAZETTE [APRIL 


particularly on leading shoots, into two or three pairs of broad 
acute lobes; when they first unfold membranaceous, dark red, 
soon becoming green and lustrous, almost fully grown when the 
flowers open, and at maturity thick and coriaceous, dark green 
and very lustrous on the upper surface, pale and dull on the 
lower surface, from 2 to 3 in. long, from I to. 1% in. wide, and 
‘on vigorous shoots often from 4 to 5 in. long and from 2 to 2% 
in. wide, with prominent midribs usually red on the lower side 
and few slender prominent primary veins slightly impressed 
above and usually running to the points of the lobes; petioles 
stout, grooved, more or less winged above by the decurrent 
bases of the bladés, glandular particularly on vigorous shoots, 
from % to 3% in. long; stipules on vigorous shoots lunate, stipi- 
tate, coarsely glandular-serrate, occasionally % in. in length. 
Flowers 34 in. in diameter on slender elongated pedicels in 
broad compound very thin-branched many-flowered corymbs ; 
calyx-tube narrowly obconic, the lobes narrow, acuminate, 
elongated, entire or sparingly and irregularly glandular-serrate, 
reflexed after anthesis; stamens 15 to 20; filaments slender; 
anthers small, pale yellow; styles 2 to 5. Fruit in many-fruited 
drooping clusters, oblong, dull brick red, pruinose with a slight 
glaucous bloom, marked by small dark lenticels, from % to % 
in. long, % in. thick; calyx-cavity deep and narrow, the lobes 
little enlarged, dark red near the base on the upper side, usually 
erect, often deciduous before the maturity of the fruit; flesh 
yellow, thick, dry, sweet, and mealy; nutlets 2 to 5, rounded 
and ridged on the back with a low broad rounded ridge, light- 
colored, about ¥ in. long. 

A tree often 30 ft. in height with a tall trunk from 12.to 18 
in. in diameter covered with close dark bark broken into thick 
plate-like scales, spreading lower and erect upper branches 
forming a broad open rather irregular head, and slender nearly 
straight branchlets, bright orange-brown and lustrous during 
their first and second seasons, becoming pale reddish-brown 
during their third year, and ultimately ashy-gray, and unarmed 
or armed with few straight slender bright chestnut-brown lus- 
trous spines from I to 1% in. in length. 


Too! | NORTH AMERICAN TREES ~ 838 


Flowers early in May. Fruit ripens toward the end of 
October. 

Rich woods on the drier parts of the bottom-lands of the 
Mississippi river opposite St. Louis. East St. Louis, Illinois, 
G. W. Letterman, June 10, 1881, and H. Eggert, 1882. Banks of 
Mississippi river, near Oquawka, Illinois, H. WN. Patterson. In 
November 1882, Mr. Letterman collected specimens of a Cratae- 
gus at Prescott, Arkansas, which may possibly belong to this 
species but the specimens are too fragmentary to make the 
determination satisfactory. 

In the Arnold Arboretum the flowers of Crataegus nitida open during the 
first week in June and the fruit ripens towards the end of October and falls 
gradually. At this season of the year it is a handsome object, the large 
leaves of the long vigorous shoots having gradually turned to a rich orange- 
yellow color through shades of bronze and orange-red, while the leaves on 
the shoots of lateral branchlets are still green and very lustrous and make a 
beautiful contrast with the abundant but rather dull-colored fruit. 

This species, which was distinguished by Dr. Engelmann on its dis- 
covery as Crataegus nitens in herb. was not published by him. It has 
been variously considered a variety of Crataegus viridis L., and as a natural 
hybrid of that species and of Crataegus Crus-galli L. The supposition of a 
hybrid can probably be safely dismissed. The plants are too numerous and 
were formerly too generally distributed over the Mississippi bottoms near St. 
Louis to make such a supposition probable; and the seedlings of this tree 
raised at the Arnold Arboretum which flower and fruit freely every year show 
none of the variation found in the descendants of hybrids when these are 
fertile. From C rataegus viridis it differs in its larger and much thicker and 
more lustrous leaves, larger flowers, much larger oblong pruinose fruit, and 
in its dark close bark, the bark of Crataegus viridis being pale or often nearly 
white and covered with thin loose scales. 

” Crataegus Brazoria, n. sp.— Leaves oval to obovate, acute or 
acuminate at the apex, sharply wedge-shaped, or on leading 
shoots occasionally oblong and usually broadly cuneate or some- 
times rounded at the base, coarsely and irregularly glandular- 
Serrate above the middle with spreading teeth, mostly entire 
below; when the flowers unfold covered on both surfaces with 
short soft pale hairs, particularly on the lower side of the thin 
midribs and primary veins, and at maturity thin and firm, gla- 
brous, dark green and lustrous on the upper surface, paler on the 


234 BOTANICAL GAZETTE [APRIL 


lower surface, from 2 to 2% in. long, from 14% to 1% in. wide, 
and on leading shoots often from 3 to 4 in. long and 3 in. wide; 
petioles slender, more or less winged above, tomentose, ulti- 
mately glabrous or puberulous, from ¥% to 34 in. long; stipules 
foliaceous, slightly falcate, acuminate, coarsely serrate, villose, 
long-stalked, from % to ¥% in. long, often % in. wide, or on 
vigorous shoots lunate and usually entire. Flowers about 34 in. 
in diameter in loose broad many-flowered compound thin- 
branched villose corymbs; calyx-tube narrowly obconic, villose 
with long matted white hairs, the lobes lanceolate, acuminate, 
obscurely glandular-serrate or nearly entire, villose on both sur- 
faces, reflexed after anthesis; stamens 20; filaments slender, 
elongated ; anthers small, dark red ; styles 5, surrounded at the 
base by a thin ring of hoary tomentum. Fruit in spreading or 
drooping few-fruited glabrous clusters, subglobose or often rather 
longer than broad, bright canary-yellow, marked by a few large 
dark lenticels about % in. long; calyx-tube elongated, with a 
broad deep cavity, the lobes deciduous before the maturity of 
the fruit; flesh thin, light-colored, dry and mealy; nutlets 5, 
rounded and ridged on the back, about ¥ in. long. 

Flowers from the middle to the end of March. Fruit ripens 
after the 1st of October. 

A tree 20 ft. in height with a trunk 6 in. in diameter and 8 ft. 
tall with ashy gray bark covered with small closely appressed 
scales, numerous upright branches forming a handsome symmet- 
trical round-topped head and slender slightly zigzag branchlets 
marked by small oblong pale lenticels, coated when they first 
appear with hoary tomentum, soon becoming glabrous and light 
reddish-brown, and ashy-gray during their second year, and 
apparently unarmed. 

Still known only from a single individual growing by the 
roadside in the bottoms of the Brazos river in the town of Bra- 
zoria, Texas, where it was found on March 25, 1900, by W. M. 
Canby, B. F. Bush, and C. S. Sargent, and subsequently visited 
by Mr. Bush in April and October, 1900. 


i. 
i 
| 
1 


—— 0 


1901] NORTH AMERICAN TREES 235 


This very distinct and interesting species can perhaps be best grouped 
with Crataegus viridis Linn. From that species it differs in its larger 
flowers, in the bright canary-colored fruits, unlike in color those produced by 
any of the North American species now known, in the form and texture of 
the leaves and in the color of the bark of the trunk and branchlets. 

~ Crataegus glabriuscula, n. sp. Glabrous with the exception 
of a few soft caducous hairs on the under surface of the large 
leaves of vigorous shoots and on the upper side of the calyx- 
lobes. Leaves oblong-ovate to semi-orbicular, or to broadly 
ovate on vigorous shoots, rounded, acute, or short-pointed at the 
apex, cuneate from below the middle and decurrent on the slen- 
der often glandular petioles, coarsely doubly serrate except 
toward the base, occasionally more or less deeply lobed above 
the middle, particularly on vigorous shoots, with two or three 
pairs of short broad acute lobes, membranaceous at the flower- 
ing time but at maturity subcoriaceous, hard and firm, dark green 
and lustrous on the upper surface, pale on the lower surface, 
from 1% to 2 in. long, from 3% to % in. wide, with slender 
pale yellow midribs ‘and primary veins running obliquely 
toward the apex of the leaf, conspicuous secondary veins and 
reticulate veinlets; stipules lunate to semiorbicular, coarsely 
glandular serrate, from % to % in. wide, Flowers % in. in 
diameter on slender elongated pedicels, in few-flowered rather 
compact compound thin branched corymbs; bracts and bractlets 
linear, obscurely glandular-serrate, minute, caducous ; calyx-tube 
broadly obconic, the lobes short, acute, entire or slightly and 
irregularly glandular-serrate, reflexed after anthesis; stamens 
20; filaments slender, elongated; anthers comparatively large ; 
Styles 5. Fruit drooping on slender stems, oblong to obovate, 
dull orange-color, marked by minute dark lenticels, 4 in. long; 
calyx-tube elongated with a deep broad cavity, the lobes but 
little enlarged, dull red on the upper surface toward the base, 
spreading or appressed, often deciduous; flesh very thin, yellow, 
dry and hard; nutlets 5, rounded or obscurely grooved on the 
back, about 3, in. long. 

A tree 20 to 25 ft. in height with a slender trunk often 12 


| 


236 BOTANICAL GAZETTE [APRIL 


in. in diameter, covered with brown scaly bark, ascending 
branches, forming a narrow head, and thin branchlets marked 
by many small pale lenticels, dark red-brown when they first 
appear, becoming bright chestnut-brown and very lustrous dur- 
ing their first summer and ashy-gray during their second year, 
and unarmed or armed with very slender straight chestnut-brown 
lustrous spines usually from 3{ to 1 in. in length. 

Flowers about the middle of April. Fruit ripens in August. 
High dry bottom lands of the Trinity river and its branches at 
Dallas, Texas, in forests of Ulmus crassifolia and Celtis Mississip- 
prensis, F. Reverchon, July 1899, B. F. Bush, April 1900. 

Betula Alaskana, n. sp.—Leaves rhomboidal to broadly ovate 
and truncate or rounded at the base, acuminate, very coarsely den- 
tate above the middle with glandular teeth, entire below, more or 
less covered with resinous glands while young, from 1% to 3 in. 
long, from I to 1% in. wide, dark green on the upper surface, 
yellow-green on the lower surface, the slender midribs and 
remote veins puberulous below or ultimately glabrous. Stami- 
nate catkins clustered, sessile, about 1 in. long, % in. thick, their 
scales ovate, acuminate, puberulous, light red, yellow on the 
margins. Pistillate catkins slender, cylindrical, pedunculate, 
about I in. long and % in. thick. Fruiting catkins from 1 to 
1% in. long, from ¥% to % in. thick, their scales about as long 
as broad, ciliate on the margins of the lobes, the central lobe 
acute or acuminate, the lateral lobes erect and acute or spread- 
ing and rounded. 

A tree with close light red bark, usually from 35 to 40 ft. in 
height with a trunk 6 or 8 in in diameter, and occasionally 50 
ft. in height with a trunk a foot in diameter, spreading and pen- 
dulous branches, slender red-brown branchlets more or less ver- 
rucose with conspicuous resinous glands, and obovate obtuse 
winter buds ¥ in. in length. 

Saskatchewan, &. Bourgeau, 18 58 (in Herb. Gray); near 
Prince Albert in latitude 53, July 1876, Yohn Macoun; north- 
westward, reaching the Alaskan coast on the shores of the Lynn 
Canal (Muir, Canby, and Sargent, August 1897); and westward. 


1901 | NORTH AMERICAN TREES 237 


This is the ‘‘canoe birch”’ of all travelers in Alaska, and it is the 
common birch tree of the Yukon valley, where it grows spar- 
ingly near the banks of that river in coniferous forests and abun- 
dantly on sunny slopes and hillsides, and extends down the river 
at least as far as the Russian mission, two hundred and thirty- 
three miles above its mouth (M. W. Gorman 7x Uitt.). 

From Betula papyrifera Marsh, which is common in one of its forms in 
southern Alberta, northern Idaho, and Montana, and in the Blue mountain 
region of eastern Washington and Oregon, Betula Alaskana may be distin- 
guished by the close reddish bark of the trunk which is less flexible and does 
not separate as easily into layers as the bark of the eastern canoe birch, by the 
much more verrucose branches, obtuse winter buds, rhomboidal long-pointed 
leaves, stouter fruiting catkins, and by the more habitually acute or acuminate 
central lobe of theirscales. The specimen in Herb. Gray collected by Bourgeau 
in flower on the Saskatchewan was referred by Regel (Bu//. Mosc. 18: 398; 
DC. Prodr. 16 : 164) to his Betula alba, subspecies verrucosa 5 resinifera, 
the Asiatic types of this variety being from Udskoi in eastern Siberia and 
from Transbaical. Specimens of the Alaska tree sent by me to the Herbarium 
of the Imperial Botanic Garden at St. Petersburg were pronounced, however, 
unlike any Asiatic species, and with the slight knowledge of the northern 
Asiatic species of Betula which we possess in this country it does not seem 
safe to follow Regel in uniting North American with Asiatic species. To Mr. 
Gorman I am indebted for very complete specimens of Betula Alaskana 
collected at different points on the Yukon in 1899, and for the first reliable 
information I have been able to obtain in regard to the trees of the Yukon 
valley and their distribution. 

BETULA OCCIDENTALIS Hooker. There are three arborescent 
birches in the region between the eastern base of the Rocky 
mountains in the United States and the shores of Puget sound 
and British Columbia. 

First, Betula papyrifera Marsh, in one of its forms which 
enters this region from the east and is not rare in northwestern 
Nebraska, northern Dakota, Idaho, and Montana, and reaches 
the mountains of eastern Oregon. 

Second, the large tree which grows on the lower Fraser 
river, on the shores and islands of Puget sound, and on Van- 
couver island. This tree has reddish bark which is particu- 
larly noticeable on young plants, pubescent branchlets, acute 
winter-buds, leaves pubescent on the lower surface, and fruiting 


238 BOTANICAL GAZETTE [APRIL 


catkins about 14% in. long and ¥% in. thick, with scales longer 
than broad, the middle lobe being acuminate and much elon- 
gated. Specimens of this tree, which is perhaps the largest 
of all birch-trees, were first gathered on the shores of the straits 
of Fuca by Dr. John Scouler during his visit to the northwest 
coast in 1825-1827. These specimens were described by Hooker 
in his Flora Boreali-Americana as Betula occidentalis, although 
with them he united a specimen collected by Douglas in the 
interior but west of the Rocky mountains. The tree from the 
straits of Fuca appeared first in the description of Betula occiden- 
talis which was evidently drawn principally from the specimen 
of that tree, and must be considered the type of Hooker’s 
species, while the second specimen included in this description, _ 
collected by Douglas, is the Rocky mountain form of Betula 
papyrifera. In the shape of the leaves this species resembles 
some of the forms of Betula papyrifera. The bark, however, is 
very different from that of the eastern tree, and it is probably 
best to consider it a species. 

Third, the half-shrubby dark-barked species with spreading 
gracefully drooping stems which ranges as far south as Colo- 
rado, Utah, and northern California. This plant was collected 
by Nuttall on the Sweetwater, one of the branches of the Platte, 
and was first described and figured by him as Betula occidentalis 
(Sylva 1: 23. pl. 7). Torrey in the Botany of Fremont’s Expe- 
dition repeats this error. This same species was also described 
and figured in King’s Rep. (5 : 323. pl. 35) as Betula occidentalis 
by Watson who repeated his error in the Botany of California, and 
it is this plant which is described and figured as Betula occiden- 
fais in my ninth volume of The Silva of North America, where 
an allusion only is made to the true Betula occidentalis of the 
coast in a note under Betula papyrifera. Nuttall found another 
small birch in the Rocky mountain region and on the plains of 
the Columbia which he described and figured as Betula rhombi- 
folia in the first volume of his Sylva published in 1842. This | 
plant, judging by one of Nuttall’s original specimens in the Gray 
Herbarium, is the narrow-coned form of the plant described by 


1901 | NORTH AMERICAN TREES : 239 


Nuttall as Betula occidentalis, which is common in eastern Oregon 
and Washington and eastward into Montana and Idaho. I¢ the 
two forms, which seem to vary only in the thickness of the 
cones, are considered to belong to one species, this would have 
to bear Nuttall’s name of Betula rhombifolia, if Tausch four years 
before had not used that name for an European species. Some 
of the specimens of this third species bear a strong resemblance 
to a fragmentary specimen of Betula microphylla Bunge, as 
pointed out to me by Mr. M. L. Fernald, but this evidence of 
the identity of the Rocky mountain and the Altai plants would 
hardly seem to warrant the adoption of Bunge’s name for our 
tree, for which I now propose the name of Betula fontinalis. 

“ Cupressus pygmaea, n. sp. (Cupressus Goveniana var. pygmaea 
Lemmon, Handbook West-American  Cone-bearers 27. ASOR, 
Cupressus Goveniana Sargent, Silva, N. Am. 10: 107 in part (not 
Gordon). 1896.—The Cupressus of the coast region of Mendo- 
cino county, California, can be readily distinguished from the 
other North American species by its thin black seeds not more 
than % in. long which show no tendency to vary to the thick 
light red seeds of Cupressus Goveniana which are fully % in. in 
length. This character and the isolation of the region which it 
inhabits remote from that occupied by other species make it pos- 
sible and convenient to separate this northern tree from the Cupres- 
sus Goveniana of central and southern California, to which it was 
doubtfully referred by Englemann in herb. who, like myself 
when the tenth volume of Zhe Silva of North America was pub- 
lished in 1896, was unacquainted with the seeds. From Cupres- 
sus Goveniana the northern tree differs also in its rather stouter 
branchlets with deeper green never glaucous foliage, usually 
sessile often oblong cones with less prominent bosses on their 
scales which vary from six to ten in number, while the cones of 
Cupressus Goveniana are usually composed of six scales. In a 
genus like Cupressus where individuals vary greatly within cer- 
tain limits and good specific characters are so difficult to find, 
these peculiarities would hardly justify the separation of the 
northern tree from Cupressus Goveniana were it not for the 


240 BOTANICAL GAZETTE [APRIL 


character found in the seeds which make this the easiest of 
our species to recognize. 

Cupressus pygmaea inhabits the high barren region on the 
coast of Mendocino county, extending from Ten Mile run on 
the north to the Navarro on the south, and, beginning about 
three quarters of a mile from the ocean, does not extend inland 
more than four miles. The soil of these barrens is yellow clay 
covered with deposits of sea sand and a thin layer of peat. On 
this poor soil the plants begin to bear cones when only a foot or 
two high, but on the borders of the barrens and of the deep 
gullies which penetrate them where trees occasionally escape for 
several years the fires which constantly sweep over the region 
they grow in better soil to a height of 30 or 4o feet, but from 
overcrowding rarely develop the spreading branches peculiar to 
all species of Cupressus growing in abundant space. 

The name pygmaea used by Lemmon to distinguish the 
dwarf plant stunted by overcrowding and insufficient nourish- 
ment is unfortunate as a specific name, for there is no difference 
between the smallest and the largest plants except in size; and 
it is probable that individuals of this species on the borders of 
the barrens, if they could be protected from fire, would in time 
grow to a large size, for the oldest plants now standing show no 
signs of maturity and none of them are believed to be much 
more than fifty years old (C. Purdy im Ui#.). 


ARNOLD ARBORETUM. 


| ; 
: 
. 


% THE EFFECT OF HYDROCYANIC ACID GAS UPON 
GRAINS AND OTHER SEEDS. 
C. O. TOWNSEND. 
(WITH SIX FIGURES) 
INTRODUCTION. 

THE experiments which form the basis of this paper were 
undertaken more than two years ago for the purpose of deter- 
mining the effect of hydrocyanic acid gas upon the germination 

of seeds. The necessity for the work arose from the rapidly 
increasing use of this gas in the destruction of insect pests 
| infesting stored grains and other seeds. Since these experi- 
ments were begun a number of the largest flour mills in this 
country and Canada have been fumigated for the destruction of 
insects without removing the thousands of bushels of grain stored 
' in the mills, and likewise numerous barns and granaries filled 
with grain have been fumigated for the same purpose. Since 
) the grain thus fumigated may be used either for seed or for food, 
it is important to know whether or not the germinating quality 
of the grain has been injured and whether grain thus treated 
would be injurious for food. A preliminary account of the 
experiment in regard to germination of seeds after treatment 
with hydrocyanic acid gas was read before the Botanical Section 
| of the A. A. A. S. at the Columbus meeting and subsequently 
| published. 

The use of hydrocyanic acid gas for fumigating purposes is 
of comparatively recent origin, having been introduced by the 
2. S. Department of Agriculture, Division of Entomology, in 
1886, for the fumigation of orangetrees. It was first used for the 
fumigation of greenhouses by Dr. A. F. Woods,’ Director of the 

i *TOWNSEND: “The effect of hydrocyanic acid gas upon the germination of 
| seeds.” Sci. Amer. Suppl., no. 1248, 2 D. 1899. 
| *Woops: The variable effects of hydrocrani acid gas on plants and animals. 
Rep. Soc. Plant Morph. and Physiol., Am. Nat. 1899. 
Igor] 241 


242 BOTANICAL GAZETTE [ APRIL 


Division of Vegetable Physiqlogy and Pathology, and even 
before it was used for fumigating purposes its effect in solution 
was determined by Dr. Loew.3 

This gas has now come into universal use not only for the 
fumigation of trees and grains as above indicated, but also for 
the fumigation of greenhouses,‘ nursery stock, and even orchard 
trees of all kinds.5 

The object of the writer in performing these experiments, 
the results of which are herein recorded, has been simply to 
determine whether or not hydrocyanic acid gas has any injurious 
effect upon grains or other seeds; and, if so, under what condi- 
tions and to what extent the injury is produced. Hence this 
paper is merely preliminary to the more minute study of the 
subject, since it deals only with the cause and effect, while the 
nature of the action of the poison upon the several parts of the 
seed still remains for further investigation. 

In general the methods employed in these experiments have 
consisted simply in placing the seeds to be tested in air tight. 
chambers in which hydrocyanic acid gas is generated. In the 
first experiments the chambers consisted of air tight boxes of 
several cubic feet capacity, used primarily for the fumigation of 
nursery stock. These boxes were open on one side, the open 
side being placed next the ground, which was first made per- 
fectly smooth and level. In experimenting with these boxes, 
they were raised slightly on one edge, the material to be fumi- 
gated was placed under them, and at the moment the hydro- 
cyanic acid gas was generated the boxes were lowered and sand 
quickly pressed around the edges next the ground, making them 
practically gas-tight. In these as in the subsequent experiments 
the gas was generated by placing cyanid of potassium (KCN) in 
dilute sulfuric acid (H,SO,), the result of the chemical action 
being the evolution of hydrocyanic acid gas (HCN) according 

3LoEW: Natur. System der pecinims Bull. Imp. Univ. Coll. Agr. 1+: 34- 
1893. The poisonous action of dicyan 

4Woops and Dors : The use ae acid gas for fumigating green- 
houses and cold ra ‘tes no. 37. Ser. 2, U. S. Dept. Agr., Div. 
SJoHNSON: Reportonthe SanJoséscale. Bull. 57, Md. Agr. Exper. Station. 1897- 


a Se eee 


—_— eS re: 


De a a 


1go1] EFFECT OF HYDROCYANIC ACID GAS 243 


to the following chemical equation: 2KCN + H,SO, = 2HCN 
+K,SO,. Inthe later experiments bell jars were used having 
a capacity of approximately one third of a cubic foot each (fig. 
z). The bell jars were provided with ground glass stoppers, 
and were placed on ground glass plates, the edges being sealed 
with oil to prevent the escape of the gas. Having prepared the 
seeds in the manner desired, they were placed on the ground 
glass plate, and in a small beaker directly in the center of the 
plate was the dilute sulfuric acid, so that when the bell jar was 
placed in position the beaker containing the sulfuric acid was 
exactly below the glass stopper in the bell jar. The potassium 
cyanid was carefully weighed and wrapped in fine tissue paper. 
A piece of twine, just equal in length to the distance from the 


. lower surface of the glass stopper in the bell jar to the bottom 


of the beaker containing the sulfuric acid, was tied to a pack- 
age containing the desired weight of potassium cyanid, and the 
other end of the string was fastened by means of soft paraffin to 
the lower surface of the glass stopper of ‘the bell jar. When 
everything was ready to begin the experiment the package of 
potassium cyanid attached to the string was lowered through 
the opening in the top of the bell jar, and just at the instant 
when this package came into contact with the dilute sulfuric 
acid the ground glass stopper fell into place and thus hermeti- 
cally sealed the jar. By this means it was possible to keep 
the seeds for any required length of time in an atmosphere of 
hydrocyanic acid gas of desired strength. When the bell 
jars were opened, even after several months, the characteristic 
odor resembling peach kernels could be detected, showing that 
the gas had neither escaped nor been absorbed, although in at 
least some of the experiments a portion of the gas was probably 
absorbed, since it has a strong affinity for moisture. It should 
be borne in mind that both the potassium cyanid and the hydro- 
cyanic acid gas are deadly poisons and great care should be 
exercised in using them. Potassium cyanid is a white solid and 
is on the market in several forms. It should be purchased for 
use in the lump form usually known as 98 per cent. cyanid of 


Fic. 1.—Bell jars in which the effect,of hydrocyanic acid gas upon the germination of seeds is being determined. 


ALLAZVID TRIINVLIOT 


Thad 


EO ae net nt 


Igor | EFFECT OF HYDROCYANIC ACID GAS 245 


potassium, and should be kept in air tight cans or jars until ready 
for use. 
4 If the seeds were to be treated in the dry state, no moisture 
was added except what was used to dilute the acid, and this 
consisted of only a few cubic centimeters of water. On the 
other hand, if damp seeds were to be treated with hydrocyanic 
acid gas, they were first soaked for the desired length of time in 
water, and then placed on moist filter paper under the bell jar, 
which was lined with damp filter paper, thus providing for the 
seeds a moist atmosphere favorable for their germination. All 
the chambers, whether damp or dry, were kept in the general 
laboratory where a pretty constant temperature of about 18° C. 
was maintained. Further details in regard to methods can best 
| be made clear by referring to the accompanying tables and their 
descriptions. The materials used in these experiments con- 
| sisted for the most part of corn, wheat, beans, and clover seed. 
: Occasionally other seeds were used in addition to those men- 
| tioned, but these four were carried through all the experiments 
and may be considered as fairly representing the seeds of mono- 
cotyledonous and dicotyledonous plants. Under certain condi- 
tions some of the seeds used seemed to be more sensitive to the 
treatment to which they were subjected than the others, but in 
general the behavior of the seeds under the conditions used was 
| similar for all seeds under identical conditions, the difference 
being simply a matter of degree. In other words, if the effect 
of the treatment was to accelerate germination all the seeds ger- 
minated more quickly than under normal conditions, whereas if 
the effect of the treatment was to retard germination then all 
germinated less readily than under normal conditions, although 
| all seeds of the same kind did not germinate at the same 
| moment. 

The strength of hydrocyanic acid gas used in these experi- 
ments is stated in terms of the weight per cubic feet of space 
to be fumigated of the cyanid of potassium used in generating 
the gas. In fumigating nursery stock for the destruction of 
insect pests, as well as for fumigating mills, barns, etc., 0.25%" of 


246 BOTANICAL GAZETTE [APRIL 


cyanid of potassium is used for each cubic foot of space to be 
fumigated.® 

In these experiments the strength of the gas varied from 
0.0038" per cubic foot to 1.458" per cubic foot. While in the 
first experiments the object was simply to determine whether or 
not the ordinary strength of the gas used in fumigation would 
be harmful to the grain or other seeds, in the later experiments 
an effort was made to determine the maximum and minimum 
strength of gas that the seeds could resist under the several 
conditions employed. It should be noted that time is an impor- 
tant factor in these experiments. Nursery stock may be fumi- 
gated in thirty minutes, so that all insect life will be destroyed ; 
but in fumigating buildings in which large bulks of grain are - 
stored time must be allowed for the gas to penetrate the mass 
of grain and thus reach the insects that may be scattered 
through it. While time is being given for the gas to penetrate 
the bulk of grain, the grain on the surface of the bulk is exposed 
to the influence of the gas from the time it is generated. Hence 
an effort was made to determine the maximum and minimum time 
required for the grains to respond to the influence of the various 
strengths of gas used under the several conditions employed, 
and with this point in view the exposures varied from one hour 
to one year. 

EXPERIMENTS WITH DRY SEEDS. 

Trial experiments.—In the first experiments dry seeds were 
placed in shallow dishes and exposed for one hour to charges of 
gas varying in strength from 0.258" to 1.452" of potassium cyanid 
per cubic foot. Only a thin layer of seeds was placed in each 
dish, so that the gas came directly into contact with each seed. 
At the end of one hour the seeds were taken from the fumigat- 
ing chamber and placed in hydrant water for twenty-four hours. 
At the expiration of this time they were spread upon moist filter 
paper in damp chambers and kept at ordinary room temperature 
of about 18° C. In this, as in all the other tests, control experi- 
ments were used in which the seeds were treated in precisely 

SJOHNSON, Joc. cit. 


om 


Igo1 | EFFECT OF HYDROCYANIC ACID GAS 247 


the same manner and kept under exactly the same conditions as 
in the subsequent experiments, except that they were not sub- 
jected to the influence of the gas. 

The first line in each table expresses the result of the con- 
trol experiments for that particular set of experiments, so that 
in whatever manner the seeds under examination behaved it was 
clear that the variation in behavior from the seeds in the control 
experiments resulted entirely from the effect of the hydrocyanic 
acid gas. 

In all cases it was found that the seeds that had been sub- 
jected to the gas for one hour germinated just as readily as if 
they had not been in the gas at all. Hence it may be con- 
cluded that perfectly dry seeds may be subjected for a short 
time to the influence of gas six times as strong as that ordinarily 
used in fumigation without any appreciable injury to the ger- 
minating quality of the seed. 

TABLE I. 


Dry seeds subjected to the influence of hydrocyanic acid gas, then soaked and placed 
under conditions favorable for germination; temperature 18° C. 


oo of — Time of expo- | Time of soaking | a, ime required Percentage of a pr 
cubic sure to gas rt germination | germination ceadies 
a 
C00 irate: I hour 24 hours 24 hours 100 normal 
0.25 I ““ 24 6c 24 ‘6 100 “ 
0.45 I ‘“ 24 se 24 $6 100 6 
0.60 i 24 ue 24 ee 100 " 
0.75 ee 24 ‘“ 24 #6 100 i 
1.00 I ‘“ 24 ‘ 24 bad 100 & 
1.25 : Peed 24 es 24 hs 100 . 
1.45 sai of ae ee 100 


Subsequent experiments.—In the trial experiments the seeds 
were subjected to the influence of hydrocyanic acid gas for one 
hour only. In order to determine the influence of this gas upon 
seeds exposed for a long time, subsequent experiments with dry 
seeds were undertaken. The seeds were placed in shallow dishes 
which were covered by bell jars, made air tight, and the gas 
generated in the manner described. Control experiments were 
Started at the same time, in which the seeds were covered by 


248 BOTANICAL GAZETTE [APRIL 


bell jars under the same conditions as in the experiments, with 
the exception that no KCN was placed in the dilute sulfuric acid. 
-The results are best expressed in the following table: 


TABLE II. 


Dry seeds treated with peas bocs acid gas for ae periods of time: temperature 
oughout about 18° 


ee ee ee fe | Pee ohn 
foot to gas ecient germination germination seedlings 

G,000))5 ca, fo) 24 hours 24 hours 100 normal 

OeF8 So as 15 days ro Seah fe 100 accelerated 

THO oo. gi aq St eae 100 wie 

[2 We ly ee es Bee 60:5." op Sa 3 aes 100 celer. 

POCO Cs 60.°2" 24°" ve 100 a aipeane 

U333 60 rss Ba oats aq.“ 75 slightly coene 

DOO cee Gr aS ss ae Bieca, 60 greatly retarde 

eet edo 2. heres ys ee 50 retarded half 

HOO ons 24000 0, ey 96..." 10 very slight 

0.333 S05 0s 2a a 40 20 none 

T00022005- 0 S6g ns 2400" no germination te) 


From the foregoing table it will be seen that the seeds were 
constantly under the influence of the gas for from fifteen days 


to one year. The strength of the gas was constant in each jar, 


z. e., it did not vary except possibly by the absorption of a small 
quantity of the gas into the seeds and into the few cubic centi- 
meters of dilute acid used in generating the gas. It is surprising 
that the seeds should germinate more readily after they had 
been exposed to the gas for a few days, although the same 
phenomenon was observed in the action of ether upon the germi- 
nation of seeds.’ 

At the expiration of sixty days the acceleration of germina- 
tion had passed its maximum, and shortly after began to be 
retarded, until at the end of two hundred and forty days germi- 
nation practically ceased for those seeds that were exposed to 
the gas from one gram of KCN per cubic foot (fig. 2,¢). The 
seeds that were in the weaker charge of gas continued to 


7 TOWNSEND: The influence of ether upon the germination of seeds and spores. 
Bot. Gaz. 27: 458. 1899. 


Tgor | EFFECT OF HYDROCYANIC ACID GAS 249 


germinate about four months longer, but that was evidently the 
limit, as only a few of the seeds germinated very feebly at the 
expiration of that time (jig. 3,4). Not only was the germination 
hastened, but the subsequent growth was likewise accelerated for 
a limited time. Although the subsequent growth of seedlings 


a b é 


G, 2.—a, control; 4, from seeds that were eight months in des from 0.333%" 
KCN per cubic foot; c, seeds that were eight months in gas from 18" KCN per cubic 


was oe some instances very marked, being almost double that of 
the normal, this acceleration was of short duration, rarely lasting 
more than one week, and then gradually resuming the normal 
rate of growth. Hence the acceleration is probably not of 
sufficient duration to be of any commercial value. 

It appears from these experiments that dry seeds may be 
treated with hydrocyanic acid gas continuously for several weeks 
Without reducing the percentage of germination or interfering 


* 


250 BOTANICAL GAZETTE [APRIL 


seriously with the subsequent growth of the seedlings. It is 
clear, however, that the gas has a marked influence upon dry 
seeds. The extent of the influence depends upon the amount 
of gas per cubic foot and upon the time the seeds are subjected 
to its influence. After sixty days the percentage of germination 


a é 


Fic. 3.—a, control; 4, seeds that were one year in gas from 0.3332" KCN per 
cubic foot; ¢, seeds that were one ead in gas from 18™ KCN per cubic foot. 


was gradually reduced, and the subsequent growth of seedlings 
was likewise retarded. The greater reduction in the percentage 
of germination, as well as in retardation of growth, depended 
upon the length of time of exposure to the gas and upon the 
amount of gas present. At the end of eight months the seeds 
‘in the gas from one gram of KCN had practically lost their ability 
to germinate, since only one in ten was capable of even slight 
germination, and those were too feeble for subsequent growth 
(fig. 2,¢).. lt the gas was generated from only one third of a 
gram of KCN per cubic foot, the seeds retained their ey to 


1901 | LFFECT OF HYDROCYANIC ACID GAS 251 


germinate four months longer, or one year from the beginning of 
the experiment (jig. 3,4). In general it may be stated that the 
length of time that dry seeds retain their ability to germinate 
varies inversely with the strength of gas to which the seeds are 
subjected. Likewise the converse of this is true, viz., that the 
strength of gas that will prevent seeds from germinating varies 
inversely with the time that the seeds are subjected to the influ- 
ence of the gas. These rules are stated only for the seeds that 
are subjected to the gas under the foregoing conditions, and will 
not apply with certainty to seeds underother conditions. It should 
be stated in this connection that the seeds in the control jar did 
not germinate quite normally either in percentage or point of 
time at the end of twelve months (fg. 3,@). To what this 
slight variation was due has not yet been determined, but it does 
not seem probable that it was sufficient to affect the results in 
the jars containing the gas. Seeds from the original packages 
not inclosed in a jar germinated normally, z. ¢., 100 per cent. in 
twenty-four hours. 


EXPERIMENTS WITH MOIST SEEDS. 


Since it may sometimes be necessary to fumigate grain or 
other seeds while in a moist condition, experiments were 
undertaken for the purpose of determining what effect hydro- 
cyanic acid gas would have upon seeds that had been soaked 
for a longer or shorter time in water. After soaking the seeds 
the following conditions were observed, viz., the damp seeds 
were left to germinate in the gas which was of different 
strengths in different jars; they were then placed for a time 
in the gas of different strengths and then removed to a gas 
free atmosphere. 

Damp seeds allowed to remain in hydrocyanic acid gas.— 
By referring to tables III, IV, and V it will be seen that 
the seeds were soaked for different lengths of time, and 
that they were subjected to atmospheres of gas of different 
Strengths. 


| APRIL 


G. 4.—Corn and wheat seedlings. The larger one in each case is normal, 
while pe smaller ones were from seeds treated eight months with gas from 0.333 
KCN per cubic foot. 


1901] EFFECT OF HYDROCYANIC ACID GAS 253 


TABLE IIL 
Germination of damp seeds and subsequent growth of resulting seedlings in hydro- 
cyanic acid gas; temperature throughout about 18° C. 


Amount of KCN per cubic foot Time of —  e pre 5 
soaking germination germination seedlings 

ODO We ces ve a ea Seg 24 hours 24 hours 100 normal 
MOS oar oe Senge ag tine, Y 2405" 405-0" bs go very slow 
ROTO. aspene ee a Cislshin Nanas oo 248 no germination fc) sees 
OG te crating ice at os BA te no germination fo) 
PrOO0r We eee eo a Ba IES no germination fe) 
GLI Ome Relate tet aie pe hae aes 0 no germination fc) 


It will be observed in table III that the amount of KCN used 
varied from 0.003 to 0.0758" per cubic foot, but that no germina- 
tion took place if more than 0.0038" was used, and indeed in this 
Strength of gas germination was delayed for six days. Other 
quantities of KCN between 0.003 and 0.0108 were used, but the 
seeds seemed to be unable to germinate in any charge stronger 
than 0.003%" per cubic foot. The subsequent growth of the 
seedlings in this very small amount of hydrocyanic acid gas was 
very slight, and did not exceed two inches in ten days, in some 
instances the growth being only a fraction of an inch. 


TABLE IV. 
Same as table III, except that the seeds were soaked for a shorter time before placing 


| Time required Percentage | Subsequent 
Amount of KCN bic foot Salen cleo for of growth 

are mann soaking | germination germination| of seedlings 
ety ahem sale eee er me cake “ hours | 4 hours 100 . 
sce acl) nas OG Sa APE RSA aes 36 80 slight _ 
CO rt Gi ay = 62:5 40 very slight 
Se ae aise es ak eG eaten ee 20 none 
ght ce re. 12" no germination ty) 
pF Ac ale Orso ge a ane eo i Sie ° 


In the experiments recorded in table IV the seeds were 
soaked only twelve hours, but the control seeds germinated in 
practically the same time as in the preceding case where the 
seeds were soaked for twenty-four hours. It is also important to 


254 BOTANICAL GAZETTE [ APRIL 


note that the small charge of only 0.0038" of KCN retarded the 
germination but a few hours as compared with the retardation 
produced by the same charge of gas when the seeds were soaked 
for twenty-four hours. It is likewise a noteworthy fact that 
when the seeds were soaked but twelve hours they were much 
more resistant to the influence of hydrocyanic acid gas, as 
evidenced by the fact that at least a few of the seeds were able 
to germinate in as much as 0.0508" of KCN per cubic foot. The 
subsequent behavior of the seedlings was the same as, in the 
preceding case, 7. e., there was little or no growth after the 
plant escaped from the seed coats. In general it was found that 
the shorter time the seeds were soaked the more resistant they 
were to the influence of the gas. 


TABLE V. 


Same as table III, except that the seeds were soaked for a longer time before placing 


Amount of KCN per cubic foot ace of Time — ae pepeua mH 
es germination germination| of seedlings 

511,513 Aueestear cuts g Siar oes er 36 hours | 20 hours 100 normal 
eee ae tee 60 | slight 
Si 2A ENCE oN Ae 36." no germination Oo Rea 
SE cc se ae ear cio 36 e it ‘“s Oo 
GOGO: seni ee ena: 36 “ » es 
0.075 36 73 ‘“ ké a 


In these experiments it will be seen that the extra amount of 
soaking caused the seeds to germinate more readily than they 
would have done had they been soaked but twenty-four hours, 
both in the case of the control experiments and in the experi- 
ments in which but 0.0038" of KCN was used. However, the 
seeds did not germinate as readily as they did when soaked but 
twelve hours, nor was the percentage of germination so high as in 
either of the preceding cases. It is probable that the germination 
took place more quickly than in the experiments recorded in 
table III, because the seeds were more advanced before they 
were removed from the water and placed in the gas. In all 
cases the seeds were extremely sensitive and made but very 


tS ee 


Igor] EFFECT OF HYDROCYANIC ACID GAS 255 


slight growth after breaking through the seed coats. Leaving 
out of account the advance that the seeds made toward germina- 
tion while they were in the water and before placing them in the 
gas, it appears that their sensitiveness increased with the length 
of time that they were allowed to soak. This is to be expected 
since hydrocyanic acid gas has a strong affinity for moisture and 
is readily absorbed by water. 

Seeds soaked for a definite time in water, placed re a given time 
in hydrocyanic acid gas, and then removed to a gas free atmosphere.— 
When it was observed that the seeds which had been soaked and 
then placed in hydrocyanic acid gas germinated more slowly than 
under normal conditions, it was apparent that the seeds were 
being held in a dormant state for a longer or shorter time 
depending upon the length of time the seeds were soaked, upon 
the strength of gas used, and upon the length of time that the 
seeds were allowed to remain in the gas. In order to get more 
definite information in regard to this point, experiments were 
undertaken whose results are recorded in tables VI and VII. 


TABLE VL 
Germination of seeds and subsequent growth of seedlings after soaking in water, 
then placing for a definite time in hydrocyanic acid gas, and then removing to a 
gas free atmosphere ; temperature 18° C 


ount : Time of Time required Percentage | Subsequent growth 
KCN per cubic feta of exposure , ot | of 

‘oot aking to gas germination germination seedlings 
O:000) 2. 24 hours ° 24 hours 100 normal 
GGGs tS ry EARS: 168 hours | already germ. go slow 
GOR ek 24° Aes Ses 168 hours 50 very slow 
aa kag ae et ia 168“ Ce 20 sale started 
O.C60 oni, ea eee 16s." no germination ° 
0.120 24" POS os es 0 


As shown in table III, seeds that have been soaked for 
twenty-four hours and then left for seven.days or 168 hours in 
an atmosphere containing gas from 0.003 of KCN per cubic 
feet were able at the end of that time to germinate without 
removing from the gas, while seeds subjected to a stronger 
charge of gas were unable to germinate in the gas. Hence the 


256 BOTANICAL GAZETTE [ APRIL 


experiments as recorded in table VI were undertaken. In these 
experiments the seeds were treated as in the experiments recorded 
in table III, except that at the end of seven days the seeds 
exposed to the gas from 0.010% of KCN or more per cubic foot 
were removed from the gas and placed in a gas free atmosphere. 


6 ¢ 


Fic. 5.—a, control; 4, seeds were in gas from .0308™ KCN per cubic foot for 168 
hours; c, Seeds were in gas from .o10o™ KCN per cubic foot for 168 days. — Photo- 
grablied at end of 18 days. See table VI. 


At the end of seven days, after placing the seeds in ordinary 
atmosphere, 50 per cent. of those that had been exposed to the 
gas from 0.010 of KCN germinated. In other words, the seeds 
that germinated had been held in a dormant condition for seven 
days in the gas and for seven days after removing from the gas, 
a total of fourteen days. Likewise seeds that had been in the 
gas from 0.050" of KCN per cubic foot were held in a dormant 
condition for twenty days. In the preceding case only five 


1901 | EFFECT OF HYDROCYANIC ACID GAS 257 


seeds out of every ten germinated, while in the latter instance only 
two seeds out of every ten germinated. Hence it should be noted 
that much depends upon the natural vitality of the seeds, not 
only in these experiments but in all cases where seeds are sub- 
jected to similar treatment. If the seeds were subjected for 
seven days to a charge of gas greater than that produced from 
0.050®" of KCN per cubic foot, they were unable to germinate in 
a single instance. In those cases where germination did take 
place, the subsequent growth of the seedlings was always slow 
and feeble. It is true that some of the seedlings, in these as well 
as in other experiments where the subsequent growth was slow, 
were able to overcome the influence of the gas after a longer or 
shorter time and to grow at the normal rate. 

Since the seeds subjected to the gas from more than 0.050%" 
of KCN per cubic foot were unable to germinate, it was decided 
to test the effect of a comparatively strong charge of the gas fora 
short time. The results of these experiments are recorded in 


table VIT. 
TABLE VII. 


Same as table VI, except that a stronger charge of hydrocyanic acid gas was used for 
a short time. 


Amt. of KCN Time of door e! Time required for at Subsequent growth 
per cubic foot soaking to gas germination nation of seedlings 
O.000 i 2 24 hours oO 24 hours 100 normal 
O250' ee Ba oi Ss hours pgae oi 50 fi 
0.250. . SAS 6:5 no germination ) 
0.250 - i 24 “ I 2 “ it % “ re) 


In these experiments it was found that some of the seeds 
were able to resist for more than three hours the influence of the 
gas from 0.25°" of KCN per cubic foot, although after three 
hours one half the seeds were unable to germinate and the other 
half were held in check for seventy-two hours or forty-eight hours 
beyond the usual time of germination. However, the seeds that 
did germinate produced seedlings that grew at the normal rate. 

Seeds soaked, treated with gas, washed and placed in HCN-free 
atmosphere —The experiments recorded in table VIII were 


258 BOTANICAL GAZETTE [ APRIL 


undertaken in order to determine whether the hydrocyanic 
acid gas that retarded germination after the seeds were removed 
from the atmosphere of the gas remained in the moisture that 
adhered to the seeds, whether the gas was primarily absorbed 
by the seeds, or whether retardation was simply an after effect 
of the gas upon the seeds. 
TABLE VIII. 
Seeds soaked, placed for a definite time in hydrocyanic acid gas, then washed and 
placed in an atmosphere free from hydrocyanic acid gas ; temperature throughout 
experiment about 18° C. 


KEN’ od Time of exposure Subsequent ie grain - Percentage of ae 
cubic foot soaking to gas treatment germination seedlings 
0.000 . 24 hours (a) Borsa 24 hours 100 normal 
O400°: Ba 6 hours | washed ora ay 100 i 
Gil00 ss P 24ge (aie notwashed| 96 “ 50 3{ nat. size 
EGO 2a pn washed i440 100 normal 
D.250 -; 1 Sg oe not washed| no germ. fe) wane 
Oso Sea ies ade washed 168 hours 60 ¥ nat. size 
0.250 . | 24 “ 24035 washed ye eile 10 slight 
C250 48h eae ADS washed no germ. fc) 


It is apparent that more or less of the hydrocyanic acid gas 
remains in the film of water that adheres to and surrounds the 
seeds after they are removed from the gas, since they germinate 
much more readily after washing. It should be noted that the 
washing did not consist in soaking the seeds but simply rinsing 
them under the hydrant. If the seeds were not washed they 
were unable to germinate after six hours in the gas from 0.250 
of KCN per cubic foot, whereas the seeds were able to germi- 
nate after eighteen hours in the gas, providing they were washed 
immediately after removal. If they were exposed to the same 
charge of gas, viz., 0.250" of KCN per cubic feet for twenty- 
four hours or more, they were unable to germinate even when 
they were thoroughly washed after removing from the gas. It 
would seem from the experiments that more or less of this pol- 
sonous gas remains in the moisture that surrounds the seeds, 
and that the gas clings to the damp surface of exposed seeds. 
If the gas is not removed by washing it seems to have a strong 


a Sea 


Igo1] EFFECT OF HYDROCYANIC ACID GAS 259 


affinity for moisture and for the seed, and eventually penetrates 
the seed even after the treated seeds have been exposed to the 
ordinary atmosphere. However, not all of the gas remains on 
the outer surface of the seed nor in the moisture sorrounding it, 
since a marked influence of the gas could be observed in those 
seeds that were first treated with the gas and then washed. 


a b 

6.—a, seeds in gas from 0.108" KCN per cubic foot for six hours, then 

washed and placed in gas-free atmosphere ; growth practically normal; 4, seed in gas 
from 0.258" KCN for eighteen hours, then washed and placed in gas-free atmosphere ; 
¢, Seeds in gas from 0.258" KCN per cubic foot for twenty-four hours, then washed and 
placed in gas-free atmosphere.— Photographed at end of two weeks. See table VIII. 
Conclusions.— Seeds, whether in the dry or moist condition, 
are capable of absorbing hydrocyanic acid gas from the sur- 
rounding atmosphere, whether the amount of gas in the atmos- 
phere is large or small per cubic foot. The gas thus absorbed 
has a marked influence upon the germination of the seeds and 
upon the subsequent growth of the seedlings. This influence is 
exerted upon the embryo or upon the enzymes or upon the 


260 BOTANICAL GAZETTE [ APRIL 


stored food material of the endosperm, or it may be that the gas 
exerts influence directly or indirectly upon two or more of these 
seed parts. If the grain or seeds are dry the influence of the 
gas is far less marked than if they are moist, and the drier 
they are the less the influence of the gas. It would seem that 
the gas exerts its influence, therefore, through the medium of 
the moisture contained in the seeds and in the seedlings. Even 
in older plants it is the more succulent parts that are most 
readily affected by the gas. The seed coats serve more or less 
as a protection to the inner seed parts, for as soon as the seed- 
lings escape from the seed coats they are more seriously affected 
by the gas, and if the charge is sufficiently strong, the seedlings 
refuse to grow almost as soon as they leave the seed coats. Dry 
seeds are sufficiently resistant to the influence of hydrocyanic 
acid gas to be treated for several weeks with an atmosphere 
saturated with the gas without destroying their vitality. It 
would be impossible, however, to preserve even dry seeds indefi- 
nitely in any strength of the gas, since it eventually penetrates 
them and impairs and finally destroys their vitality. If the 
seeds are damp they are much more susceptible to the influence 
of the gas, and should not remain more than two or three hours 
in gas of sufficient strength to destroy animal life. If for any 
reason it becomes necessary for the damp seeds to remain for a 
longer time in the gas, the injury will be materially lessened by 
thoroughly washing the seeds in water as soon as they are 
removed. 

The fact that damp seeds are more susceptible to hydro- 
cyanic acid gas than dry seeds would indicate that the gas is 
taken up more readily by the damp seeds, and the same is true 
of other gases. 


EFFECT OF HYDROCYANIC ACID GAS ON SEEDS FOR FOOD. 
Dry seeds.— Only a few experiments have been performed 
along this line, but probably a sufficient number to determine 
the point in question, viz., whether dry seeds treated with 
hydrocyanic acid gas retain enough of the poison to make them 


1901] EFFECT OF HYDROCYANIC ACID GAS 261 


injurious to animal life. Grain was subjected to gas of different 
strengths and for longer or shorter periods of time, varying from 
one to sixty days. Grains thus treated were from time to time 
fed to mice that had been caught without injury and placed in 
glass cages so that they could be observed constantly. The 
cages were provided with an abundant supply of air and water 
and kept at ordinary normal temperature of the laboratory 
where the mice had been living previous to the beginning of the 
experiment. Occasionally the mice began eating the grains as 
soon as they were placed within reach, but as a rule several 
minutes to several hours elapsed between the time the grains 
were taken from the hydrocyanic acid gas and the time they 
were eaten by the mice, thus giving time for any gas that remained 
in contact with the seed or that had penetrated the seed coat to 
escape into the atmosphere. In one instance, for example, a 
mouse was fed one dozen kernels of corn and three dozen grains 
of wheat that had been for four and one fourth days in an 
atmosphere containing gas from one gram of potassium cyanid 
per cubic foot. The mouse began eating the grain at once, and 
at the end of twenty-four hours had eaten the whole of five 
grains of corn and had eaten the chit out of five other grains. 
It had also eaten fourteen grains of wheat and had eaten the 
chit out of eleven others with no pathological results. Several 
similar experiments were carried through with like results. 
Hence it seems safe to conclude that dry grains treated for 
several days with hydrocyanic acid gas of sufficient strength to 
destroy insect pests that may be in the grain will in no way 
affect animal life, and may therefore be used for food without 
injury. 

Damp seeds—The damp seeds were soaked for twenty-four 
hours and then treated with gas in the same manner as in the 
preceding experiments, and were kept in the gas for different 
periods of time varying from several hours to several days. 
Here as in the germination experiments we find that moisture 
has a decided influence upon the susceptibility of the grains 
to the gas. For example: after some corn and wheat had 


262 BOTANICAL GAZETTE [APRIL 


been soaked for twenty-four hours in water and then left 
for forty-eight hours in the gas obtained from one gram of 
potassium cyanid per cubic foot, a mouse in apparently good 
health was given twelve grains of the corn and thirty-six grains 
of the wheat. The mouse began eating at once and ate the chit 
out of one kernel of corn and began ona second kernel when he 
suddenly became stupid and was unable to walk without stag- 
gering. That the mouse was hungry is evidenced by the fact 
that it began eating as soon as the grain was placed in the cage, 
and from the fact that it had been given but little food on the 
preceding day for the purpose of having it hungry enough to 
begin at once on the grain as soon as it should be removed from 
the gas. Although the mouse lived for several hours, it eventu- 
ally died apparently from the effects of the small amount of 
grain eaten, as it did not eat any more of either kind of grain 
nor would it eat cheese or any other material placed before it. 
In general, it was found that if the mice ate the grain immedi- 
ately after taking it from the gas they became stupid and even- 
tually died from the effects. However, if the grain was allowed 
to remain for a time out of the gas before it was eaten, no ill 
effects seemed to be produced, although the grain did not seem 
to return to its normal condition, as it was never eaten readily 
even after it became perfectly dry. When the mice could be 
induced to eat it, as they were in several instances, it did not 
seem at all injurious. It may be concluded, therefore, that the 
fumigation of dry grains with hydrocyanic acid gas does not in 
any way injure the grain for food purpose. And even if the 
grain is damp it will not be injured for food if it is allowed to 
air for a short time after fumigating before it is prepared for use. 


SUMMARY. 


Dry seeds may be fumigated with the usual strength of 
hydrocyanic acid gas for the length of time required for the 
destruction of animal life without in any degree interfering with 
the germinating power of the seeds. 

Dry seeds may be subjected for several months to the 


1901 | EFFECT OF HYDROCYANIC ACID GAS 263 


influence of hydrocyanic acid gas at the rate of one gram or 
less of potassium cyanid per cubic foot without entirely destroy- 
ing the ability of the seeds to germinate. 

Dry seeds subjected to the influence of hydrocyanic acid gas 
derived from one gram of potassium cyanid per cubic foot will 
lose their germinating power at the expiration of eight months, 
while the same seeds subjected to the gas from one third of a 
gram of potassium cyanid per cubic foot will retain their vitality 
until the expiration of twelve months. 

The exposure of dry seeds for from fifteen to sixty days to 
the influence of hydrocyanic acid gas from one third to one 
gram of potassium cyanid per cubic foot will hasten germination 
and accelerate the growth of the resulting seedlings. Although 
the acceleration continues for several days, it does not seem to 
be of sufficient duration and degree to be of any practical value. 

Damp seeds are much more sensitive to the influence of 
hydrocyanic acid gas than dry seeds. 

Seeds soaked twenty-four hours or more will not germinate 
in gas stronger than 0.0038" of potassium cyanid per cubic foot, 
whereas, if the seeds are soaked but twelve hours, they are able 
to germinate in an atmosphere containing hydrocyanic acid gas 
from 0.050™ of potassium cyanid per cubic foot, and in much 
less time than when soaked for twenty-four hours. 

Seeds soaked thirty-six hours will germinate much more 
readily than when soaked only twenty-four hours, but will not 
germinate in a stronger atmosphere of hydrocyanic acid gas than 
0.003*" of potassium cyanid per cubic foot. 

Seeds soaked for twenty-four hours and then left for seven 
days in an atmosphere of hydrocyanic acid gas will remain 
inactive while in the gas, and from seven to twelve days after 
removal, but will eventually germinate to some extent if the 
strength of gas used does not exceed 0.050™ of potassium cya- 
nid per cubic foot, 2. ¢., hydrocyanic acid gas is capable of hold- 
ing seeds in an inactive state for two weeks or longer without 
destroying their vitality, even when the conditions are otherwise 
favorable for germination. 


264 BOTANICAL GAZETTE {APRIL 


Damp seeds are injured to a much less extent when fumi- 
gated with hydrocyanic acid gas if they are washed immedi- 
ately after removing from the poisonous atmosphere. 

The vitality of damp seeds is seriously injured if treated for 
three hours with gas from 0.25®" of potassium cyanid per cubic 
foot, provided the seeds are not washed after removing from the 
nfluence of the gas. On the other hand, damp seeds may be 
treated with the same strength of gas for six hours with no 
appreciable injury, provided they are washed and placed in an 
atmosphere of ordinary air to germinate. 

Dry seeds treated for several days with hydrocyanic acid gas 
_ of any strength will not be injured for food. 

Damp seeds treated with hydrocyanic acid gas of any strength, 
and even for short periods of time, should not be used for food 
until several hours after removing from the gas. The effect of 
the gas eventually passes off and the grain may be eaten with 
safety, although long exposure to the gas seems to render it 
unpalatable. Grains and other seeds may be fumigated with 
hydrocyanic acid gas for the destruction of insect pests without 
injury to the germinating quality of the seeds and without ren- 
dering them injurious as foods. 


AGRICULTURAL COLLEGE, 
College Park, Md. 


na cn, 


THE TUBER-LIKE ROOTLETS OF CYCAS REVOLUTA. 
CONTRIBUTIONS FROM THE HULL BOTANICAL LABORATORY. 


A, CC. LF rR. 
[WITH TEN FIGURES | 


THE coral-like outgrowths in connection with the roots of 
Cycas revoluta have long been known. In 1871 and 1872 Reinke? 
published an account of them and described the associated endo- 
phytic alga, which he referred to Anabaena (fig. 70). In 1894 
Schneider? published further details, and referred to these struc- 
tures as root tubercles. My own work was begun at Indiana 
University and continued at The University of Chicago. 

The occurrence of these structures in Cycas revoluta is by no 
means uniform in greenhouse plants, as they are abundant in 
some cases, few in others, and entirely lacking in still others. 
They are connected with the upward rising rootlets, usually 
spreading radiately from the apex and just behind it, and are 
most abundant at or very near the surface of the ground, but 
may occur several inches beneath it. They vary considerably in 
Size, but are always larger than the normal roots of the same 
age, and by branching may form clusters 2.5°™ or more in diam- 
eter (fig. 7). 

This branching is apparently dichotomous and such a claim 
has been made for these rootlets. According to Reinke? this fact 
indicates relationship with the Lycopodiales, while Schneider’ 
ee that it is a case of atavism. If by true dichotomy it is 

*Einige Bemerkungen iiber das Spitzenwachstum der Gymnospermen Wurzel. 
Gott. ‘tick ble 530. 1871. 

Parasitische Anabaena in Wurzeln der Cycadeen. Gott. Nachrichten 107. 1872. 

* Mutualistic symbiosis of algae and bacteria with Cycas revolute. Bot. Gaz. 
19: 25-32. pls. 3,4. 189 


4 3 Beitrag zur Kenntniss der Gymnospermwurzel. Just’s Bot. Jahresb. 1: 205-207. 
1673. 


Igor] 265 


266 BOTANICAL GAZETTE [ APRIL 


meant that the whole of the apical meristem passes into the 
meristem of the branches, this is not a case of true dichotomy. 
At an early stage the meristems of the two branches become 
definitely outlined, but between them, retaining a central apical 
position, there remains a portion of the original meristem. This 
true apical meristem 
gives no indication of 
continuing the axis, and 
does not even begin to 
form bundle elements, 
but soon becomes indis- 
tinguishable, and the 
later stages of the root- 
lets show what seems 
to be a real dichotomy. 
The conclusion is that 
this is a case of apparent 
dichotomy in which the 
meristem of the main 
axis ceases to function 
almost as soon as the 
meristems of the two 


Fic. 1.— Habit of tubercles; nat. size. branches are differenti- 

ated from-it. It may 

be of interest to note that some of the tubercled rootlets do not 

branch, and that the normal rootlets do not show even an appa- 
rent dichotomy. 

A transverse section of one of the tubercles (fig. 2) shows a 
central vascular cylinder like that of the normal root, and a very 
thick cortex interrupted about midway by the greenish algal 
zone (jig. 2,a). The inner mass of cortex consists of ten or 
twelve layers of cells. The outer region is more differentiated, 
consisting of elongated parenchyma cells next to the algal zone, 
a zone of roundish cells with small intercellular spaces, a layer 
of cork cambium, and two layers of corky cells. 

In longitudinal sections the algal zone is seen to extend from 


1901] TUBER-LIKE ROOTLETS OF CYCAS 267 


the base of the tubercle almost to the very apex (fig. 3). Where 
first distinguishable it consists of cells whose contents are some- 
what denser than those of the neighboring cells, and before any 
intercellular spaces appear the cells 
next to the zone on both sides become 
very distinctly differentiated into a 
layer (/) resembling a tapetum. When 
fully developed the algal zone consists 
of loosely connected cells with large 
intercellular spaces which are occupied 
by the algae (figs. 4,5). The radial 
elongation of these loosely connected 
cells induced Schneider to call them 
palisade tissue. Where large lenticels Fic. 2.— Transverse section 
occur there is a break in the algal zone, of tubercle. X15. a, algal 
and instead of the usual layers the 
round parenchyma cells extend from the phellogen to the vascu- 
lar cylinder (fig. 6). The round cells with their small inter- 
cellular spaces 
form an effective 
air-conducting tis- 
sue. 
Lenticels occur 
_- in abundance upon 
es the tubercles, and 
are found near the 
tip of a root upon 
which young tu- 
bercles are grow- 
@ ing. They occur 
also in thickened 
primary roots 
which do not bear 
tubercles. That 
they are developed 


G. 3.— Longitudinal section of tubercle. XX 40. 5 
a, rte zone; f, tapetum-like layer, with hyphae. very early in the 


N 
° 


268 BOTANICAL GAZETTE [APRIL 


growth of a tubercle is evidenced by the fact that they may 
be seen forming very near the tip. To determine whether 
these structures are really lenticels, air was forced through 
the tubercles under water by means of an aspirator. The 
streams of air which were given off indicated that the struc- 
tures are lenticels in function as well as in form. It might be 
inferred, therefore, that the so-called tubercles are used in aera- 
tion, but this would seem to be but an incidental result of their 
structure. It has been shown by Jost‘ that plants deprived of a 
sufficient supply of oxygen develop air conducting tissue and 
abnormal growths, and it is possible that this represents the con- 
dition of Cycas. 

Upon comparing the general structure of the tubercle-like 
rootlets with those which are normal, it is to be noted that the 
tubercles have round tips which do not possess a true root cap. 
Instead of the conical root cap of the normal root they have a 
sheath of several cell layers extending over the tip and also 
enveloping the entire tubercle as an outer cortex (fig. 3). No 
such cortex is developed in the normal root. 

In studying the fungi and bacteria of the tubercles, cultures 
were made on agar. From these cultures at least three bac- 
terial forms and an organism resembling the Rhizobium of 
Schneider’? were obtained (jigs. 7, 8). The bacteria included 
one very large form, a small chain-like form, and a coccus form. 
The hyphae of fungi were also observed in the cells of the 
tubercles just in advance of the algal zone (fg. 9), but they 
were not in a condition to be identified. In fact, a zone of dif- 
ferentiated cells extends from the algal zone to the meristem of 
the tip (fig. 3), the cells being shorter than the. adjoining cells 
and their contents more granular. 

The fungi and bacteria which are in the cells in advance of 
the algal zone seem to prepare the way for the algae (fig. 0), 
since their presence seems to result in very much enlarging the 
small intercellular spaces, which become the relatively large 


‘Ein Beitrag zur Kenntniss der Athmungsorgane der Pflanzen. Bot. Zeit. 45 °37— 
39. 1887. 


Igor} TUBER-LIKE ROOTLETS OF CYCAS 269 


chambers occupied by the algae. The presence of the fungi and 
bacteria within the cells seems to retard the nutritive work, so 
that the cells cannot keep pace with those adjacent. This pro- 
duces tension which results in a development of spaces, and by 


Coke 


Fics. 4-10.— a, algae of intercellular spaces; 4, bacteria; c, parenchyma; s, inter- 
cellular spaces of algal zone; #, elongated parenchyma; /, round corky cells; , cam- 
bium of lenticel ; J, hyphae of fungus; ¢, crystal of calcium oxalate. 

Fic. 4. Intercellular spaces of the algal zone. X 120.— Fic. 5. The same. X 190. 
~—Fic. 6. Part of a cross-section, showing a lenticel and a break in the algal zone 
beneath. X 120.— Fics. 7 and 8. Bacteria and rhizobia found in cultures from 
tubercles. XX 1425.— Fic. g. Part of longitudinal section showing beginning of the 
formation of interce!lular spaces of the algal zone, and cells containing hyphae of 
fungus. XX 300.— FIG. 10. Algae found in the tubercle. X 480. 


means of this tension some of the cells are broken, while others 
appear to be broken down by the organisms within them. After 
the algae have gained entrance, their growth and multiplication 
Probably result in still further increasing the size of these 


270 BOTANICAL GAZETTE [APRIL 


intercellular spaces, since they become larger the further they 
occur from the region of origin. 

In the early stages of the growth of the tubercles, before 
they contain the algae, numerous lenticular areas may be 
observed at their bases and also upon the adjacent part of the 
root which bears them. These peculiar areas have openings in 
them which appear as crevices, and as the tubercles grow older 
the whole area frequently breaks away, and it is doubtless 
through these rents that the algae effect an entrance. Together 
with the algae the fungi and bacteria also enter, the latter forms 
penetrating the cells and causing the enlargement of the inter- 
cellular spaces. 

In reference to the symbiotic relations which exist between 
these various organisms it is difficult to speak with any cer- 
tainty. The fungi and bacteria doubtless find congenial condi- 
tions of moisture and food in connection with the algae, and are 
in turn the principal agents in producing the intercellular spaces 
in which the algae thrive. It is barely possible that there may 
be some such chemical attraction between the fungi and algae and 
the cells of the differentiated layer as has been stated by Miyoshi. 
It has been demonstrated by Vines,° Frank,? MacDougal,’ 
and others that fungi growing upon the surfaces of cells or 
within them may aid in nutritive work by converting free nitro- 
gen and the simpler nitrogen compounds into the more complex 
forms used by the plant. The same function has been attributed 
to certain algae by Prantl,9 especially including the nostoc 
forms. This last observation suggests the possibility of the use 


5 Ueber Chemotropismus der Pilze. Bot. Zeit. 52 : 1-28. p/. 7. 1894. 


§On the relation of the formation of tubercles on the roots of Leguminosae and 
the presence of nitrogen in the soil. Ann. Bot. 2: 386- 389. 8. 


7 Ueber die auf Wurzelsymbiose beruhende Ernahrung gewisser Baume durch 
unterirdische Pilze. Ber. deut. bot. Gesell. 3 : 128-145. f/. zo. 188 


8 Symbiotic saprophytism. Ann. Bot. 13: 1-47. pls. 7, 2. 1899. 


9 Die Assimilation freien Stickstoffes und der Parasitismus von Nostoc. Hed- 
wigia 28: 135. 1889. 


gor] TUBER-LIKE ROOTLETS OF CYCAS 271 
of the nostoc forms within the tubercles of the cycads in assist- 
ing nitrogen assimilation. 

In conclusion, therefore, the tubercles of cycads may be 
said to have at least two functions, that of aerating, and that of 


' assisting in nitrogen assimilation. 


THE UNIVERSITY OF CHICAGO. 


BRIEFER ARTICLES. 


WALNUT BACTERIOSIS. 


I CHARACTERIZE here briefly a new micro-organism pathogenic to 
Juglans regia and related species, which has been under investiga- 
tion at this laboratory for several years. Announcements of the isola- 
tion of the organism and a demonstration of its pathogenic action on — 
walnuts were contributed several years since to the leading horticultural 
journals of the Pacific coast. Further communications relative to this 
disease are in course of preparation, which will enter more fully into 
the characterization of the organism, its pathogenic powers, its cultural 
peculiarities, and will give full details as to the nature and treatment of 
the disease which it causes. 

Pseudomonas juglandis, n. sp.—A short, rod-shaped micro-organism 
with rounded ends, actively motile, bearing a single long polar usually 
wavy flagellum. Length of organism, taken from colony in acid gela- 
tin, set directly from walnut, and stained with gentian violet, 1-24, 
according to whether the germ has just divided or has elongated but 
not yet divided. Just before separation a pair of germs will usually 
average about 2m in length. Average breadth of organism about 
0.5m. Occurs singly or in pairs, and sometimes in shorter or longer 
chains. Produces a bright chrome yellow growth on potato and many 
other media. When growing normally on potato the starch is so acted 
upon by a diastatic ferment produced by the organism, that it is altered 
throughout a wide band beyond the margin of the culture of organ- 
isms. This band of converted starch may extend o.5-1™ or more 
beyond the margin of the growth or germs, appears white to the eye, 
fails to show normal starch reaction to iodine, yields marked grape 
sugar reactions, has an exceedingly sharp and well defined limiting 
outline, often passing so sharply through a cell as to include only the 
starch grains on one side of the cell. This broad and distinct ferment 
band distinguishes this organism at once from Pseudomonas Stewartt 
and P. hyacinthi, as well as from the more nearly related P. campestris, 

‘which occasionally forms a weak but much narrower band, and from 
272 [APRIL 


1901] BRIEFER ARTICLES 273 


all other uniflagellate organisms studied. Organism prefers neutral or 
acid reaction of culture medium, a moderate degree of alkalinity 
inhibiting growth; it liquefies neutral and acid gelatin. Produces no 
gas in fermentation tubes of sugar solutions; growth confined to neck 
and bulb of tube, hence aerobic, no growth under mica plate. Colo- 
nies in malic acid potato gelatin and agar circular; at first clear but 
soon decidedly yellow, margin sharp. This organism is distinguished 
from P. campestris, the most nearly related species of the genus, aside 
from the characters already assigned, in producing an abundant and 
bright yellow pigment on the surface of extracts of the leaves of the 
following plants, while P. campestris produces little or no pigment 
upon such extracts: walnut (/uglans regia), magnolia (Magnolia macro- 
phyla), fig (Ficus Carica), castor bean (Ricinus communis), loquat 
(Eriobotrya Japonica). 

Organism strikingly pathogenic to nuts, leaves, and tender branches 
of Juglans regia. In the young walnuts the epicarp and forming shell 
and kernel are destroyed, in the older nuts the epicarp alone may be 
affected. The leaves are commonly attacked along the veins and on 
the petiole, the organism often entering the vessels. The branches 
become infected near the growing point, and all tissue systems are 
destroyed, the organism entering and wintering in the pith cavity; it 
also winters in fallen nuts. An unlimited number of infections may 
be induced— and thousands have been so induced —by spraying the 
young nuts with a pure water culture of the organism. This test shows 
the germ to be one of the most active, self-effective parasitic or patho- 
genic species of the genus yet known to infect plants. 

Pseudomonas campestris has not been found infesting the cabbage 
fields of the regions where the present disease prevails. Walnut bac- 
teriosis is known to exist over large areas in southern California and 
to a limited extent in the central and northern portions of the state.— 
NEwrton B. Pierce, U. S. Dept. Agric., Div. Veg. Phys. and Path., 
Pacific Coast Laboratory, Santa Ana, Cal. 


GCURRENT.LEPERATURE. 
BOOK REVIEWS. 
Vegetable physiology. 

THE NEED of a book of moderate compass to present in clear and suc- 
cinct form the principles of plant physiology is well met by Professor Green’s 
recent volume.!. In a handy volume of 450 pages the author has discussed 
the structure and differentiation of the body (35 pp.); the skeleton (17 pp.); 
the absorption, transport and loss of water (50 pp.); aeration and respiration 
(29 pp.); the various processes of nutrition, including absorption and syn- 
thesis of foods, translocation, storage, digestion, secretion and waste products 
(156 pp.); energy relations (17 pp.); growth (15 pp.); temperature and the 
influence of environment on plants (25 pp.); irritability and movement (39 
pp.). These are not the author’s chapter headings but fairly represent the 
various topics. He has “endeavored to present the plant as a living organ- 
ism, endowed with particular properties and powers, realizing certain needs, 
and meeting definite dangers,’ and to keep clearly before the reader the 
fundamental identity of the activities of plants and animals. 

In general the presentation leaves little to be desired. Particularly com- 
mendable is the treatment of nutrition. The author not only lays stress upon 
the true nature of foods, from which category he rightly excludes water and 
carbon dioxid, but discusses clearly photosynthesis, from which he differ- 
entiates, none too strongly, the amylogenic function, and presents, as in his 
earlier work, an excellent chapter on digestion. 

The treatment of respiration and the energy relations of plants, however, 
is not well arranged. To discuss respiration in connection with aeration and 
before nutrition seems to be a reversal of the logical sequence and that to no 
gain. It needs to be brought into intimate relation with nutrition, of which 
it is a phase, and also with the release of energy, of which it is one method. 
Aeration, though incidental to respiration, should be treated in connection 
with gas ae just as root hairs are discussed in connection with water 
absorption. 

The handling of osmosis might be much improved by a broader physical 
treatment, for there is no process whose fundamental principles are so little 
comprehended by most students. The author seems to overestimate the 
meres ‘of root pressure in the transport of water through the stems, for 

N, J. REYNOLDs: An introduction to plant oo 8vo. pp. xx-+ 
459. “rp eel London: J. and A. Churchill. 1900. ros. 
274 [APRIL 


1901] CURRENT LITERATURE 275 


it certainly has no effect beyond the cortex of the root when transpiration is 
active. On the other hand, the presentation of the probable relation of 
evaporation and osmosis in the mesophyll to water transfer is very satisfactory. 

The illustrations are remarkably unequal. Some are excellent; some are 
crude, but sufficient; some are so diagrammatic, without such remark, as to 
be misleading ; and some disfigure a book which deserves the best possible 
engravings. We suspect some of the worst of being relics of a long-deceased 
“botany,” resurrected from the publishers’ stock-room. Many of the illus- 
trations, though not original, are not credited to their sources, while others 
are; of this no explanation is given. 

No references are made to original papers or to other works; only in a 
few cases is the proponent of any particular view or theory referred to; and 
controverted points are scarcely alluded to. The danger from this is that the 
student may get the too common impression that the book deals out finally 
settled knowledge. It is difficult to know where to draw the line between 
the necessary dogmatism of the elementary work and the discussion of the 
treatise, but we believe in showing up the gaps in a subject fully. While 
this book only professes to be “an introduction,” it is nevertheless so thor- 
ough that it would have been improved by a selected bibliography following 
each of the various topics. For though we have Pfeffer’s treatise at hand, its 
Citations of literature are too detailed for the student of elementary physiol- 
ogy, and he would have profited greatly by the same judicious sifting of the 
literature as Professor Green has made of the facts.—C. R 


The problems of life. 


THE FUNDAMENTAL problem of physiology is the constitution of living 
matter. But we are so far from a knowledge of the chemistry of proteids, 
not to mention protoplasm, that the best we can do now is to attempt the 
framing of logical and consistent hypotheses, in the hope that these will sug- 
gest research in fruitful lines. If the true character of such hypotheses is 
kept clearly in view, they may prove helpful, but there is always a danger 
that the hypothesis may be held so aun that new facts are distorted to 
fit our interpretation of other facts. 

There is before us the first part of a work by Dr. Giglio-Tos, of the Uni- 
versity of Turin, which is an attempt to set forth a logical and consistent 
hypothesis of the organization of protoplasm and its fundamental functions, 
notably assimilation, reproduction, respiration, photosynthesis, cell 
division? A second part is to discuss ontogeny and its problems. 

LI0-Tos, ERMANNO: Les problémes de la vie: Essai d’une interpretation 
sro. mm des phénoménes vitaux. 1° partie: La substance vivant et la cytodiérése. 


vo. pp. viii 286. figs. 33. Turin: The author, Palazzo Carignano. 1900. Io 
francs 


276 BOTANICAL GAZETTE [APRIL 


The general thesis of the book is that vital phenomena are explicable 
without invoking hypothetical or mysterious forces and are all referable to 
relatively simple fundamental causes. Since the minutest observable organ- 
isms undergo fission, the author makes the assumption that fission occurs 
also in the molecules of living substance. This he illustrates by the behavior 
of certain chemical compounds. Methyl-ethyl-ketone, for example, on taking 
up three atoms of oxygen, cleaves into two molecules of acetic acid. The 
chemical analogy may be carried further. By treating the acetic acid mole- 
cules with phosphorus pentachlorid, and the acetyl chlorid so formed with 
zinc ethyl, there are finally produced four molecules of methyl-ethyl-ketone 
and in the course of the process several by-products. Certainly such an 
analogy, if it be nothing more, renders easily conceivable the author’s hypo- 
thesis that assimilation of atoms or atomic groups from the medium brings 
about an increase and a rearrangement of the atoms in each molecule of liv- 
ing substance. When the proper relations within the molecule are reached, 
it divides into two new ones like the original; or into two (alike) which 
through a subsequent cycle of development produce molecules like the origi- 
nal ; or into two (unlike), only one of which develops and reproduces the origi- 
nal molecule, while the other finaliy produces again unlike molecules. By 
assuming this different behavior the author explains the production of non- 
living matter by living, and the origin of genetic and somatic plasma. 

The respiratory and photosynthetic activities of these living molecules are - 
elucidated in terms of this hypothesis ; certainly what we know of them falls 
in with it easily. 

The next assumption is that many unlike biomolecules, to use the author’s 
word for living molecules, are aggregated into living particles, or biomores, 
in which they have an arrangement determined by the chemical affinity of the 
various atoms and groups, and shifting as the chemical constitution changes. 
The life of the biomore, however, does not depend on its constitution but on 
the life of the biomolecules. 

The aggregate of biomores, submerged in water (not of constitution as 
Nageli held, but of capillarity), constitutes the bioplasm. Any view as to 
the morphological structure of the bioplasm, whether the filar structure of 
Flemming, the reticular of Frommann, the alveolar of Biitschli, or the granu- 
lar of Maggi and Altmann, is equally consonant with this theory. 

The bioplasm is not of indefinite extension but consists of small masses, 
each a symbiotic system of unlike biomores, constituting a biomonad. A 
very complex biomonad, especially when biomores constituting a nucleus are 
present, is a cell. 

ell division is based upon the same fundamental principle as the fission 
of the biomolecules. In them it is not mere addition of atoms which suffices 
to bring about division, but their mutual relations and orientation. In the 


1901} CURRENT LITERATURE 277 


biomores division again depends on the nature, number, and arrangement of 
the biomolecules. In the biomonad, division is possible (1) because the bio- 
mores are mobile, being in liquid ; (2) because they exercise a mutual attrac- 
tion (shown by their union into masses giving parts of the cell their definite 
characters), and consequently assume a determinate arrangement. But 
division of the cell is completely independent of its structure, though the 
figures accompanying division, such as the spindle, the equatorial plate, etc., 
are dependent upon structure. The efficient cause of division is indeed 
always the same, whether in biomolecule or cell, namely, the mutual orienta- 
tion of the parts; whatever the number, nature, or structure may be, division 
is inevitable when the proper arrangement is attained. Assimilation is indis- 
pensable to division, but not always sufficient. The discussion of cell division 
is elaborate and is illustrated by diagrams which make the presentation very 
clear and the process fascinatingly simple — would that it were so. 

Such is a very brief outline of the hypothesis. Of course, it adds nothing 
to our knowledge, and it would be easy to revile it as an unstable pile of 
assumptions. But the real questions are, Does it take account of the present 
facts and coordinate them? Will it suggest fruitful research? That it takes 
account of all the facts is improbable; that it does fairly coordinate the more 
important ones (alas, too few) seems evident. That it will stimulate inves- 

tigation remains to be seen. At any rate it deserves attention for what it 
purports to be, namely, an attempt to give a scientific interpretation of vital 
phenomena.—C. R. B. 

MINOR NOTICES. 

ONE OF THE latest of the series of Temple Primers is a little volume on Plant 
life and structure translated from the German o r. E. Dennert by Clara L. 
Skeat.3 The author seems to recognize no organisms as plants but the seed 
plants, and continually makes assertions of the most comprehensive sort which 
are true only of green plants. Indeed he is rather given to generalizations, 
both regarding structure and function, and to teleology of the Paley type. 

n many minor matters and in some important ones the book is not up to the 
times ; Goethean metamorphosis, sexual terms for the sporophyll and micro- 
spores, and much faulty physiology might have been more easily forgiven in 
a book dated a decade or two ago. But the best to be said of this booklet 
is that it is better than many more pretentious works and good enough for one 
to regret that it is not better.— C. R. B 

AMONG the plastic products of plants one of the groups most — 
chemically and most valuable commercially is the volatile oils. In thei 
manufacture many firms have been engaged, but none so extensively, so 

3 DENNERT, E.: Plant life and structure. Translated from the German by Clara 
L. Skeat. 16 mo. pp. viii+-115, figs. “ie — : J. M. Dent & Co. (undated). 
New York: The Macmillan Company. 40c 


spree! 7 a 


278 BOTANICAL GAZETTE [ APRIL 


exclusively, or more scientifically than Schimmel & Co., of Leipzig. Not con- 
tent with much original work in the chemistry of these substances and costly 
experimentation with methods of manufacture, this firm commissioned Dr. E. 
Gildermeister, of Leipzig, and Dr. Fr. Hoffmann, of Berlin (the latter long 
and honored leader in pharmaceutical affairs in this country), to prepare a 
work on volatile oils,* treating them in an exhaustive and critical manner from 
the modern standpoint. This work, published about a year ago, has been 
oe yeaa into English by Dr. Edward Kremers, of the University of Wiscon- 
sii. The recent rapid development of the knowledge of the volatile oils 
sey the sti volume doubly useful. With the chemistry of these bodies 
it happily combines much interesting historical matter and a description of 
the modern processes of distillation. Abundant references to original chemi- 
cal sources make the work an important aid to the investigator. 

Though primarly chemical and technical, the list of plants, arranged 
according to families, from which volatile oils are obtained will be interesting 
for the botanist and the whole book is a mine of information. Four hundred 
and thirty oils are described, the botanical sources, percentage present in 
various parts, mode of preparation and composition being given.—C. R. B 

NOTES POR STUDENTS. 

HAROLD L. Lyon, of the University of Minnesota, has announced® that 
Nelumbo, “both in its anatomy and embryogeny conforms to the type of the 
monocotyledons.”’ His full paper will be awaited with interest.—J. M. C. 


LEWIN ° contradicts the view of Stahl that raphides are important as a 


means of protection against herbivorous animals. There is no evidence of » 


mechanical injury to animals, nor of poisonous effects soi produced 
through eating plants that contain these crystals.—H. C ES 
’" Mryosut? has performed a number of experiments in order to determine 
the influence of various substances in the soil or water upon flower colors. 
His results are not uniform, though he finds in general that aluminium com- 
pounds change lilac to blue (as Molisch previously observed), while potash 
changes lilac to green, and many acids change it to red. H.C: Cowes. 
W. C. WorSDELL® has concluded that the Bennettitales are more primi- 
tive than the modern cycads, as shown by their stem structure, radial 
*GILDERMEISTER, E. and HOFFMANN, FR.: The volatile oils. Written under the 
auspices of the firm of Schimmel & Co., Leipzig. Authorized translation by EDWARD 
KREMERS. Det pp. viii+ 733, fgs. 83, maps 4g. Milwaukee: Pharmaceutical Review 
Pub. Co. 
RE 13:470. Igor. : 
° Ber. deut. bot. Gesell. 18 : 53-72. 1900. 7 Bot. Centralbl. 83 : 345- 1900. 


* The affinities of the mesozoic fossil Bennettites Gibsonianus. Ann. Bot. 14: 717- 
721. 1900. 


Igor ] CURRENT LITERATURE 279 


sporophyll, and terminal ovule. In reference to the radial character of the 
primitive sporophyll the author accepts Celakovsky’s view, and also regards 
the peculiar ovuliferous stalks as foliar rather than axial structures.— 


D. H. Scorr® has described a new genus of lycopods, fideehie by 

name, which shows features indicated by the title of the is fr 

the Lower Coal Measures and also from a much lower oa pre alcifer- 
i h 


° 
B 


growth from the sporophyll, and completely encloses it except for a slit-like 
opening above. So much of the sporophyll persists as is necessary to com- 
plete this testa-like covering, and the whole seed-like structure falls off with 
its retained and germinated megaspore.—J. . 

Dr. E. O. JorDAN® has published recently a paper upon the bacterial 
ete of streams, the observations having been made during a 
study of the chemical and bacterial condition of the Illinois river and its 
tributaries, undertaken in behalf of the sanitary district of Chicago. The 
phase of self-purification considered is that which relates to the disappear- 
ance of the sewage bacteria. Although the conclusions are not to be regarded 
as final, a lessening of the bacterial content seems to admit of no question. 
In fact, the observations show that the Illinois river at Ottawa had become 
nearly free from the great mass of sewage bacteria with which it was originally 
laden. The probable factors in this result seem to have been dilution, sedi- 
mentation (leading to exhaustion of food supply), or action of sunlight. The 
influences of mechanical agitation and aeration, and of plankton, do not seem 
to be factors which enter into the problem in connection with the Illinois river. 
mares Lie. hy: 

JoHN WEINzIRL™ has recently published a paper upon “ The bacterial 
flora of the semi-desert region of New Mexico with especial reference to the 
bacteria of the air,” in which he records the result of a two-year study. 
Both quantitative and qualitative examinations were made. The quantitative 
data were obtained by the filter method, sand being used for the filtering 
substance. Five determinations made in the vicinity of the University of 
New Mexico gave an average of aboat forty-two bacteria per cubic meter of 
air. Eleven determinations in the residence district of Albuquerque yielded 

°On the occurrence me a seed-like fructification in certain paleozoic lycopods. 
Ann. Bot. 14 : 713-717. 

70 Some ee nage the Sages self-purification of streams. Jour. Exper. 
Medicine 5 : 271-314. p/. 20 

* Jour. Cin. Soc. Nat. Hist. 1g : 211-242. 1900. 


280 BOTANICAL GAZETTE [APRIL 


an average of one hundred and forty-three per cubic meter. Comparison is 
made with the Mont-Souris results, but no mention is made of Tucker’s 
observations in Boston. Ten organisms, presumably distinct, are described 
with some detail. It is not stated whether the “recommended procedures’’ 
have been employed. The organisms described are characterized as a whole 
by their inability to ferment sugar and to peptonize gelatin.—E. O. JORDAN. 

Miss ETHEL SARGANT™ has published an exceedingly interesting paper 
upon double fertilization, bringing together the contributions to date and 
discussing the questions that have been raised. The bibliography of the 
subject is represented now by at least eighteen titles, and the phenomenon 
has been recorded for about twenty-five species. The author inclines to the 
view that the triple fusion is a true act of fertilization between a male cell 
and an egg, interfered with and rendered abortive by the introduction of the 
non-sexual antipodal nucleus. The result is not a normal embryo, but a 
small and short-lived mass of tissue. Boveri’s experiments with the eggs of 
sea urchins, in which more than one sperm was forced to unite with egg 
nucleus, are cited to prove this probable result of more than double fusion. 
The theory is advanced that the third nucleus is introduced to secure the 
degeneracy of the resulting tissue. If this is true, the definitive nucleus has 
descended from one which was the result of true fertilization, and the partici- 
pation of the third nucleus is an added feature.—J. M. C 

ITEMS OF TAXONOMIC INTEREST are as follows: EDWARD L. GREENE 
(Pittonia 4: 242-284. 1901), in continuing his “ Studies in the Compositae,”’ has 
made a special attack upon Bidens. After an interesting account of the history 
of the genus, of &. frondosa L., and of B. cernua L., he describes twenty new 
species, thirteen of them being American analogues of B. cernua, and seven 
of them segregates of B. chrysanthemoides Mx. B. Beckii is given generic 
rank, and is associated with two new species under the name Megalodonta. 
In the same fascicle new species are described under Conoclinium (3), Eupa- 
torium (8), and Antennaria.—C. S. SARGENT (Rhodora 3: 19-31. Igo!) has 
described thirteen new species of Crataegus from the Champlain valley, 
remarking that this is one of the richest regions in the world for forms of 
Crataegus.—P. A. RyDBERG (Bull. Torr. Bot. Club 28: 20-38. Igo1), in the 
fourth paper of his series entitled “Studies on the Rocky mountain flora,” 
describes new species under Arnica (2), Artemisia, Picradenia, Antennaria, 
Aster, Townsendia, Eri igeron (3), Valeriana, Cam mpanula, Castilleia (3), 
Mimulus, Pedicularis, Pentstemon (3), Polemonium, Gilia, Phacelia, Lap- 
pula, Cryptanthe, Mertensia (4), Mentzelia, Impatiens, Geranium, Lupinus (2), 
Astragalus, Aragallus, Trifolium, and Lath hyrus.—C. V. Preer (dbzd. 39-45» 
in the fifth paper of his series entitled “New and noteworthy northwestern 


es ** Recent work on the results of fertilization in angiosperms. Ann. Bot. 14: 689- 
- 1900. 


1901] CURRENT LITERATURE 281 


plants,’ describes new species under Arabis, Trifolium, Astragalus, Solidago, 
Erigeron, Antennaria, Artemisia, Crepis, Dodecatheon, Gilia, Phacelia, Lap- 
pula, and Mimulus.—L. M. UNDERWOOD (2ézd. 46-47) has described a new 
Adiantum from New Mexico.—J. M. C 

BROWN and EscomMBE have under way one of the most important physical 
researches of recent years. They have set forth some of their results and 
have applied them to plant functions in a paper on the “Static diffusion of 
gases and liquids in relation to the assimilation of carbon and translocation 
in plants.” "3 Blackman’s interesting results (1895) concerning the diffusion of 
CO, through stomata are fully substantiated. The authors further find that 
with small openings a) the rate of diffusion varies as the diameter of the 
orifice; 4) multiperforate septa do not hinder diffusion when the openings 
are about ten diameters apart; ¢) decrease in per cent. of area of holes is not 
followed by proportional decrease in amount of diffusion; @) the laws hold for 
both gases and solutes. Assuming an analogy with lines of force about an 
electrified disk, they find these results accord with mathematical calculations. 
Applying these principles to plants the authors conclude that 1) in an ordinary 
leaf (Helianthus annuus) the stomata are sufficient to permit the diffusion of 
five or six times as much as CO, as is actually used by the plant; 2) resistance 
to the absorption of a greater quantity of CO, lies in the relatively slow diffu- 
sion of the CO, after solution; 3) the stomata are more than sufficient to 
permit the escape of the observed amounts of water vapor by diffusion with- 
out any mass movements; 4) continuity of protoplasm may have more to do 
with translocation of foods than has been supposed hitherto, since over 60 
per cent. of diffusion possible through an orifice the full size of the pit would 
take place through the many minute perforations, although their area. be 
only 0.84 per cent. of the total membrane of the pit.—T. C. Fry 

IN A RECENT PAPER by A. H. R. Buller™ on chemotaxis in fern sperms it 
is shown that these cells are attracted not only by malic and maleic acids 
and several of their salts (as was known before), but also by numerous salts 
of the inorganic acids. Potassium and sodium tartrate and potassium 
oxalate also attract. Sperms of Gymnogramme Martensit were used an 
were tested by Pfeffer’s method of capillary tubes. The concentration nectes- 


sary for attraction is usually = to + normal. With malic acid it is =, to 
<=» and with sodium malate *. to %. Nitrates and chlorids of sodium, 
ammonium, and calcium do not attract, nor does lithium nitrate. The choice 
of compounds to be tested was made, apparently, with the aim of making a 


catalogue of attracting substances rather than of determining the real nature 


3 Phil. Trans. Roy. Soc., London, B. 193 : 223-292. 1900. 
Contributions to our knowledge of the physiology of the spermatozoa of ferns. 
Ann. Bot. 14: 543. 1900 


282 BOTANICAL GAZETTE [APRIL 


of the attraction. The results are tabulated according to De Vries’ obsolete 
isotonic coefficients, and in order to be studied need to be rearranged by 
base and acid. The small number of salts of any given acid makes it utterly 
impossible to draw any conclusion as to the nature of the attraction. The 
author’s propositions in this regard may or may not be substantiated later; 
several hypotheses might be formulated which would explain equally well 
the cases cited. It is unfortunate that this paper (following that of Garrey* 
by nearly a year) should carry our knowledge of this important subject of 
chemotaxis such a little way further than it was carried by that writer. We 
have tried, with some success, to coordinate the results given in these’two 
papers, but nothing definite can be attained, so far as fern sperms are con- 
cerned, until we have more data. Chemical principles must be brought into 
requisition in the selection of salts to be tested as well as in the interpreta- 
tion of results. The author makes an interesting observation with regard to 
starch grains in the vesicle of these sperms. During the swarm period the 


oxidized for the production of kinetic energy. Unhappily, no observations 
on the hydrolysis of these grains were made. We may suppose them to be 
converted into glucose by an enzyme formed in the vesicle. Thus we might 
have the non-nucleated cytoplasm of the sperm mother cell doing nutritive 
work for the moving sperm. This, if shown to be the case, might throw 
some light on the phenomena occurring in the male cells of the lower gym- 
nosperms.— BURTON EDWARD LIVINGSTON. 

AMONG RECENT CONTRIBUTIONS to paleobotany the following papers may 
be mentioned: Lester F. Warp (2oth Ann. Rep., U.S. Geol. Survey, 1900), 
with the collaboration of Fontaine, Wanner, and Knowlton, has published an 
extended account of the Triassic and Jurassic floras of the United States. 
The various areas are systematically treated, and much new material is 
incorporated. A large number of plates accompany the work, many of which 
illustrate Ward’s recently described genus Cycadella. A similar report on the 
Cretaceous is to be expected soon from the same source, and will be heartily 
welcomed, inasmuch as the literature is considerably scattered. Ward has 
also published a very interesting popular report on the petrified forests of 
Arizona, which has been issued by the United States Geological Survey.— 
F. H. KNowLTOon has published on the flora of the Montana formation (Bull. 
163, U.S. Geol. Survey, 1900); this together with his previous papers on fossil 
plants from Idaho and Yellowstone Park adds greatly to our knowledge of 
the late Cretaceous and early Tertiary floras of that part of the United States. 
— Davip Waite (2oth Ann. Rep., U. S. Geol. Survey, 1900) has worked out 

*SThe effects of ions upon the aggregation of the flagellated infusoria. Am. 
Jour. Phys. 3: 291. 1900. 


tgor] CURRENT LITERATURE 283 


in great detail the stratigraphic succession of the fossil floras of the Pottsville 
formation in the Carboniferous of Pennsylvania.—A. C. SEWARD, whose 
study of the fossil history of Gingko has been briefly noted in this journal 
(30: 139), has published on the Wealden flora of Bernissart (Mem. Mus. 
Roy. Hist. Nat. Belg., 1900); the ferns dominate in these beds rather than 
cycads, but there are no angiosperms, although the Wealden is commonly 
regarded as equivalent to the Potomac beds of our country.—GRAND ’EuRY 
has given some very interesting accounts (Compt. Rend. 130: 871, 988, 1167, 
1366. 1900) of the Carboniferous forests of France. By his study of the 
plants preserved zm sztu, he concludes that most of the Calamites and some of 
the tree ferns grew in the water; the herbaceous ferns probably grew largely 
on hummocks. Even Cordaites seems to have grown in swamps, and 
Grand ’Eury suggests analogies with Taxodium and the Dismal swamp of 
today. The prevailing horizontality of the roots strikingly suggests swamp 
habitats. The author doubts if we have evidence of upland vegetation.—At 
the last meeting of the British Association (Geol. Mag., Jan. 1901) Seward, 
Kidston, and others discussed Carboniferous conditions, as indicated by plant 
fossils. Little new material was added, though Seward suggests that the 
xerophytic structures of Carboniferous plants may perhaps be accounted for 
by swamp habitats.—PENHALLOw (Brit. Assoc. Adv. Sci., 1900) has made 
an interesting report on the flora of the Canadian Pleistocene. Interest was 
aroused a few years since by Coleman's discovery of the papaw and osage 
orange in interglacial beds near Toronto, indicating a genial climate between 
ice-sheets. Over eighty Pleistocene species are now known from Canada. 
As Knowlton suggests in discussing this report in Pant World for January, 
there is a wide field for work in the study of Pleistocene floras in America. 
The poverty of our information in this respect is in striking contrast with the 
wealth of knowledge as to the Pleistocene of Europe—H. C. COWLE 

E FORMATION OF TETRADS has lately been investigated by H. O. 
Juel.“° The contribution consists of three distinct papers which may be con- 
sidered separately. ; 

I. Tetrad formation in the ovule of Larix.—The general relationships 
between the reproductive organs of pteridophytes and spermatophytes have 
long been known, but while it is accepted that the pollen grain arises by a 
tetrad division, just as the spores of pteridophytes, it is generally believed 
that the megaspore of spermatophytes is formed without such a division. 


development of the mother cell of the megaspore to the beginning of endo- 
Sperm formation. The paper is of special interest as the first to treat this 
portion of the life history of a gymnosperm from the standpoint of modern 

**Beitrage zur Kenntniss der Tetradenbildung. Jahrb. f. wiss. Bot. 35 : 626-659, 
bls. 15,16. 1900. 


284 BOTANICAL GAZETTE [APRIL 


cytology. In material collected about the middle of April, before the snow 
had disappeared, the mother cell of the megaspore is easily distinguished by 
its large size, and by the abundance of starch. The first division is hetero- 
typic, and shows the reduced number (12) of chromosomes. At the poles of 
the spindle are granular masses which may possibly represent centrosomes, 
although the author is not willing to commit himself to this interpretation. 
During the anaphase the starch disappears, a cell wall is formed, and each 
of the two daughter nuclei divides, this time by a homotypic division, giving 
rise to a row of four megaspores, the lowest of which germinates and pro- 
duces the gametophyte. By comparing this series with the development of 
the microspores, which has been thoroughly studied in Larix, the author 
reaches the conclusion that the two series are homologous, and that the 
megaspore, like the microspore, arises by a tetrad division. While the con- 
clusion is not new, the evidence supporting it is valuable. 

Il. Zhe tetrad formation in a hybrid plant — It has long been known that 
hybrids are inclined to be sterile, and that the pollen of hybrid plants is 
commonly imperfect. The present paper deals with the formation of the 
tetrad in Syringa Rothamagensis, a hybrid between S. vulgaris and S. Per- 
sica. The form did not prove to be favorable for such a problem, because 
the pollen of both parents is poor, in S. vudgaris about 50 per cent. of the 
pollen grains appearing to be incapable of functioning, and in S. Perstca 
normal pollen being quite rare. The latter form is almost as sterile as the 
hybrid. In all three forms the development is normal up to the formation of 
the pollen mother cells. In the hybrid it was found that while most of the 
divisions in these cells were mitotic, there were also numerous instances of 
amitotic division, and abnormalities in the chromatin and in the achromatic 
figure were frequent. 

Ill. The development of the pollen grain of Carex.—As a rule, the pollen 
mother cell of a spermatophyte gives rise to four pollen grains, but it has 
been reported that in the Asclepiadaceae and Cyperaceae the mother cell 
gives rise to one pollen grain only. A careful examination of Carex acuta 
showed that the wall of the pollen mother cell becomes the wall of the pollen 
grain. It also showed that the tetrad divisions take place, but the walls are 
imperfect and only one cell of the tetrad develops into a pollen grain, the 
other three being crowded out, just as in the megaspore series three potential 
megaspores are crowded out by the one that functions CHARLES J. CHAM- 
BERLAIN, 


DPER Le Lior... 


THE ROCHESTER CODE. 


In the course of his article on nomenclatorial principles, published in the 
March issue of the BOTANICAL GAZETTE, Mr. M. L. Fernald discusses at 
some length Professor Underwood's recent treatment of the fern genera in 
this country, making the following statement in that connection: 

In Britton and Brown’s ///ustrated Flora, published in 1896, fifty-nine species of 
true ferns are “eek eit nd the names, we are told, are those authorized by the 
Rochester code. But in Professor Underwood’s latest treatment more than 25 fer 
cent. of those very species appear under different names —still the names authorized 
by the Rochester code. 

Mr. Fernald then pieeenke in support of this statement, a comparative 
table of fifteen northeastern ferns, with the names used in Britton and Brown's 
Flora and those in the last edition of Underwood’s Our Native Ferns given 
in parallel columns. This table, as an illustration of the remark above 
quoted, cannot fail to be misleading, since Mr. Fernald evidently wishes the 
reader to draw the conclusion that the Rochester code is an uncertain guide, 
and that the same author may interpret it in different ways. In order to 
_ ascertain how far such a deduction is to be considered reliable, let us analyze 
briefly the supposed divergences between Professor Underwood's interpreta- 
tion of the code in 1896 and in 1900, the dates of the two works mentioned. 
I follow substantially the same order as that of the table 

1. Onoclea Struthiopteris, being regarded by Professor Underwood now as 
generically distinct from O. sensibilis, is separated under the first available 
generic name, Matteuccia since Struthiopteris had been earlier applied to 
Lomaria, 

2. Filix replaces C seneten and Phyliitis supplants Scolopendrium 
because the first mentioned names have been proven to be the older. This is 
in strict conformity to the Ganencu of the code, notwithstanding the fact 
that conservative botanists would not admit the change. 

3- The type of Dicksonia being a tropical tree fern, the American herba- 
ceous plant is very properly separated as a genus under the first available 
name. The various species of Polystichum are also segregated from Dryof- 
éeris, and the latter name is not abandoned, as might be inferred from Mr. 
Fernald’s table. It certainly involves no novel or unusual interpretation of 
the Rochester code to divide an original aggregate into what later study may 
Prove to be distinct elements; and in the light of numerous new species of 
Igor] 285 


286 BOTANICAL GAZETTE [APRIL 


Antennaria of the p/antaginifolia group that have been proposed from time 
to time by Mr. Fernald, it is pertinent to inquire whether he does not himself 
recognize the necessity for occasional segregations. Botanists of the last 
century, who were familiar with A. A/antaginifolia, as they supposed, would 
probably be fully as dismayed over the expansion of this species as modern 
fern students are likely to be over the division of Dryopteris. 

4. Phegopteris Robertiana and Notholaena dealbata are examples merely 
of the elevation of varieties to specific rank, an everyday practice among 
botanists of all shades of opinion. 

_ The case of Pel/aea is analogous to that of Dryopteris. 

6. Cheilanthes gracilis being a homonym, the first available name, C. 
Feet, is taken up for the species. 

Applying the process of exclusion to Mr. Fernald’s table in the light of 
the above remarks, it will be observed that of the fifteen species which he 
cites, the names of only two are the result of an altered interpretation of the 
code. These are Preridium for Pteris aguilina and its allies, and the restora- 
tion of Asplenium thelypteroides for A. acrostichoides. \f we assume the 
same ratio to hold good throughout Professor Underwood’s work, we shall 
find that all but about 4 per cent. of the changes are the legitimate outgrowth 
of added research, consisting of corrections of homonyms, division of aggre- 
gate genera and species, etc.; and the same changes would probably have 
been made by the same author under almost any system of nomenclature. 
On the other hand it may be considered excellent testimony to the soundness 
of the Rochester principles that after the application to them of Professor 
Underwood’s method of determining generic types so small a percentage of 
actual changes should occur. The citation of such cases as those discussed 
above affords no basis whatever for adverse criticism of the code, since, as 
already explained, the same course might be pursued even by opponents of 
the Rochester nomenclature.— CHARLES Louis POLLARD, U. S. National 
Museum. 


NEWS. 


THE Asa Gray Bulletin has been merged into the Plant World. Mr. 
Cornelius A. Shear becomes one of the editorial staff. 

PROFESSOR J, REYNOLDS GREEN’S work on Enzymes has been translated 
into German by Dr. Wilhelm Windisch, and published by Paul Parey, Berlin. 

Mr. SAMUEL M. COULTER, fellow in botany at The University of Chicago, 
has been appointed to an instructorship in the Shaw School of Botany of 
Washington University. ‘ 

PROFESSOR Dr. EDUARD STRASBURGER has been elected a member of 
the French Academy of Sciences, and the order of “ Rote Adler, III Classe, 
mit der Schleife ” has recently been conferred upon him. 

Mr. FRANK S. COLLINS, the phycologist, of Malden, Mass., has recently 
been elected a resident fellow of the American Academy of Arts and Sciences 
(Boston), and Mr. Cyrus G. Pringle, the veteran collector, of Charlotte, Ver- 
mont, has been elected an associate fellow. 

THE ALSTEAD School of Natural History is held at Alstead Center, N. H., 
for five weeks of the summer. In 1go! it offers instruction in general 
botany by Mr. M. L. Fernald of the Gray Herbarium; and in mycology 
(fleshy fungi) by Mr. Hollis Webster, of Boston. 

Mr. L. C. CorBerrt, horticulturist of the Agricultural College at Morgan- 
town, West Virginia, has accepted a position in the Bureau of Plant Industry 
in the United States Department of Agriculture. His work will include 
investigations in regard to floriculture and gardening. 

THE assistant professorship of botany in the University of Tennessee has 
been made a full professorship, and the present incumbent, Mr. S. M. Bain, 

s been promoted thereto. The department of botany is said to be 
thoroughly equipped and to be making satisfactory progress in every way. 

A HANDSOME silver loving-cup was presented by a number of teachers 
to Mr. Thomas Meehan, the veteran horticulturist and botanist of Philadel- 
phia, on the occasion of his seventy-fifth birthday. Mr. Meehan has been a 
member of one of the sectional school boards in Philadelphia for twenty-three 
years. 

R. JARED G. Situ, formerly in charge of the Section of Seed and 
Plant Introduction in the United States Department of Agriculture, has been 
selected to organize the territorial Experiment Station in the Hawaiian 
islands. Mr. Emst A. Bessey succeeds him as “ Assistant in Charge,” the 
Section having been placed under the Bureau of Plant Industry. 
T9o1]} 287 


288 BOTANICAL GAZETTE [ APRIL 


Tue Botanical Department of Wellesley College has received a gift of 
$25,000 from Mr. H. H. Hunnewell of Wellesley, who has already manifested 
his interest in the college by opening his pinetum and orchid conservatories 
to botanical classes. The money will be used as an endowment for the 
— and the income. will be used to meet the yearly needs. 

. Hircxcock, professor of botany at the Kansas Agricultural 
College, cain Kansas, has been appointed assistant agrostologist in the 
United States Department of Agriculture. r. H. F. Roberts, instructor 
in the Shaw School of Botany, and formerly a graduate student at The Uni- 
versity of Chicago, has been appointed Professor Hitchcock’s successor. 

THE DEPARTMENT of botany of Marine Biological Laboratory has issued 
its announcement for the season of Igo1 (to be obtained from Dr. Bradley M. 
Davis, University of Chicago). A course in cryptogamic botany is offered 
by Drs. Davis and Moore; phanerogamic botany by Dr. Shaw; plant physi- 
ology by Dr. True; and plant cytology by Dr. Davis and Mr. Lawson. An 
attractive course of open lectures will also be given. 

IT 1s proposed to establish a quarterly journal of biological statistics 
under the name Biometrika, which may serve as a means not only of collecting 
under one title biological data of a kind not systematically collected or pub- 
lished in any other periodical, but also of spreading a knowledge of such 
Statistical theory as’ may be requisite for their scientific treatment. The 
movement is in charge of Professor Karl Pearson, of University College, Lon- 
don, to whom tenders of support should be sent. 

THe Act of Congress, making appropriations for the United States 
Department of Agriculture for the year beginning July 1, 1991, contains 
several important items pertaining to investigations in botanical lines con- 
ducted by several divisions in that department. The Divisions of Vegetable 
Physiology and Pathology, Botany, Pomology, Agrostology, and Gardens and 
Grounds are grouped into a bureau to be known as the Bureau of Plant Indus- 
try. The divisions will retain their individual organizations as heretofore and 
will continue to work along lines similar to those which they have already 
developed. The total appropriation for the Bureau of Plant Industry is 
$204,680. The Division of Forestry is also made a bureau for which there is 
appropriated $185,440. 


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GOUT, RHEUMATIC GOUT, 
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‘|| BUFFALO 
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VOLUME XXxI NUMBER 5 


DOTANICAL _CGAgerre 
MAY, rooz 


THE GENETIC DEVELOPMENT OF THE FORESTS OF 
NORTHERN MICHIGAN; A STUDY IN PHYSIO- 
GRAPHIC ECOLOGY. 

CONTRIBUTIONS FROM THE HULL BOTANICAL LABORATORY. 

XVII. 
HARRY NICHOLS WHITFORD. 


(WITH EIGHTEEN FIGURES) 


Portions of the summers of 1898, 1899, and 1900 were spent 
by the writer in studying physiographic ecology at several 
points in northern Michigan. The work of the first two sum- 
mers was under the personal direction of Dr. H. C. Cowles, to 
whom the writer is indebted for many valuable suggestions. 
Indeed, the work set forth in the present paper is little more 
than the testing and working out in detail the ideas on physio- 
graphic ecology developed by Dr. Cowles in his lectures at The 
University of Chicago and in his published papers. Credit is 
due Mr. W. B. McCallum for the photographs used to illustrate 
this article, with the exception of figs. 7 and 4, which were 
obtained by Mr. E. N. Transeau. 

It should be clearly understood that the conclusions reached 
are based mainly on observations and are presented tentatively. 
They are published in the hope that they may be thoroughly 
tested by other observers, and if necessary modified or dis- 
carded altogether. It is the writer’s belief that only along lines 
Similar to those advocated here can the problems of forest dis- 
tribution be successfully solved. 

f 289 


290 BOTANICAL GAZETTE [MAY 


I. FACTORS. 

The factors to be taken into account in attempting to explain 
the relations that exist in different plant associations are so com- 
plex that it will be well to consider them in detail. Since the 
object of this paper is to answer some of the questions involved 
in the development of forests, special emphasis will be laid 
upon those factors which appear to be related to tree growth. 
In order to understand the distribution of trees the subject must 
be approached from both the negative and affirmative stand- 
points. In other words, the question is, why are there forests on 
certain physiographic formations and none on those which lie 
close by? Also within the forest itself there predominates now 
one and now another tree type. Insome places the coniferous 
forest is prominent, in others the maple-beech-hemlock type is — 
the chief feature. Indeed, if enough regions are studied an 
indefinite number of combinations may be observed. Thus not 
only must the presence or absence of trees be explained, but also 
where trees are present a reason must be given for the dominance 
of any particular kind of forest. If these questions can be 
answered satisfactorily, some light will be thrown on the origin 
and development of forests. In the answer three sets of factors 
are involved, climatic, ecological, and historical. 

A. Climatic factors. The greater part of the eastern half of 
the United States is a potential forest. Here the two great 
climatic factors, temperature and moisture, are favorable to the 
development of forest trees. When a climate makes possible 
the development of any predominant type of vegetation that 
type is called a climatic formation* To be specific, the eastern 
half of the United States has a forest formation. But if a bird’s- 
eye view of any portion of this formation be obtained, there 
will be found within it groups of other plant types. These are 
designated by Warming? plant societies. Within this vast forest 

a SCHIMPER, A. F. W.: Pflanzengeographie auf physiologischer Grundlage- 
I . 


* WARMING, E.: Plantesamfund. 1895. German edition, translated by Knob- 
lauch. 1896. 


1901 | GENETIC DEVELOPMENT OF FORESTS 291 


formation there are prairie, beach, dune, heath, swamp, and other 
plant societies ; also the forest itself may be divided into a num- 
ber of different forest societies. This leads naturally to the 
consideration of those factors that make up the plant society 
conditions. 

B. Ecological factors. For convenience this set of factors may 
be divided into edaphic, atmospheric, hydrodynamic, and biotic. 

Epapuic Factors. Schimper3 has regarded the soil and its 
properties so important that he has given the name edaphic for- 
mations to those societies which owe their existence to the 
peculiarities of the soil. More than any other form of vegeta- 
tion trees need a firm anchorage medium, and hence a deep 
soil. Since the roots are the absorbing as well as the holdfast 
organs, the water content of the soil is important. Warming’s 
classification of plant societies‘ into xerophytic, hydrophytic, and 
mesophytic is based upon the amount of water in the soil. 
More plant societies doubtless owe their characteristic physiog- 
nomies to the amount and condition of the water in the soil than 
to any other one factor. Since trees present a greater transpira- 
tion surface than other forms of plants they must occupy those 
positions where there is sufficient water to maintain the trans- 
piration current. This excludes them from those regions where 
the water content of the soil approaches the minimum; a stagnant 
condition of soil water is likewise injurious to trees. Probably 
here the exclusion of air and the presence of humic acids pre- 
vent the healthy growth and activity of the root system. In 
order that the forest condition may be obtained, therefore, the 
soil must be well drained, as well as watered. 

The physical properties of the soil play an important réle, 
for upon them depends the capacity of the soil to hold water. 
The water-holding properties of the different kinds of soil are 
too well known to need treatment here. The heat-absorbing 
and heat-retaining qualities of soils must be taken into account, 
for they often determine the presence or absence of certain 
plants. 

3 OD. cit. 4 OD. cit. 


292 BOTANICAL GAZETTE [MAY 


Again, the soil furnishes the plant certain organic and inor- 
ganic compounds, and hence its chemical properties should be 
considered. Asa rule, inorganic salts are present in sufficient 
quantities in the soil; indeed in salt marshes the presence of too 
great a quantity of salt in the water excludes trees, except ina 
few instances. Soils are more often deficient in organic com- 
pounds. Organic decay, since it furnishes most of the available 
nitrates, is of great importance. The amount of /umus is so 
essential that it often determines the character of a forest, not 
only in furnishing the nitrogen to the plants, but also in 
ameliorating the physical properties of the soil. Nitrifying 
bacteria are necessary for the conversion of organic matter into 
nitrates, and if these are excluded the processes of organic 
decay discontinue, and consequently a condition like that present 
in sphagnum swamps is developed. Here, generally speaking, 
trees are excluded, except a few characteristic species. 

We must look to the soil factors for an explanation of our 
most characteristic plant societies. There must be a soil. The 
water in the soil must be sufficient. The soil must be aerated. 
The amount of organic and inorganic compounds must not be 
too great or too little, and usually the soil must contain bacteria 
or other fungi for organic decay. If all these conditions are 
present in the right proportions, the soil is capable of supporting 
a luxuriant tree growth. 

ATMOSPHERIC FACTORS. By atmospheric factors are meant 
those which influence the aerial parts of plants. They include 
radiant energy in the form of heat and light, and also the influ- 
ence of wind. Of these light is the most important. The tree 
by virtue of its many planes of plagiotropic branches gives a 
greater surface on which the light may fall than is found in any 
other plant form. Principally because it has worked out this 
successful light relation, it is ecologically the culminating type 
of plant body. Once a dense forest is established, all forms of 
low vegetation, except those species that have special shade 
adaptations, are driven out. Attention has already been called 
to the necessity of heat as a climatic factor. Heat may also be 


1901 | GENETIC DEVELOPMENT OF FORESTS 293 


an ecological factor. In exposed situations the water and humus 
contents of the soil may be dried out. For example, a forest 
may be cleared and thus exposed to the sun’s rays to such an 
extent that the soil factors will be changed considerably. It is 
obvious that it is difficult to separate the light and heat factors. 
While it is a benefit for a plant to get as much light as possible 
up to a certain point, along with the light the plant may absorb 
too much heat. 

The carbon dioxid content of the air is so constant that it 
plays little or no part as an ecological factor. Those plants 
nearest the ground are slightly favored by the greater amount of 
carbon dioxid in the lowest layers of the atmosphere. The 
effect of winds on trees, however, is often pronounced in exposed 
situations. Besides destroying trees the wind may injure them 
to such an extent that in the struggle with disease and with 
other trees they will be the first to succumb. The wind is of 
great importance also in that it causes excessive transpiration. 
The effect of wind on seed distribution is so pronounced that, 
other things being equal, those plants whose seeds are most 
easily distributed will stand the best chance in a given area. In 
conclusion, therefore, the atmospheric factors of light, heat, and 
wind must be taken into account in trying to explain the presence 
or absence of certain plant societies. 

HypRopyNAMIc FACTORS. The term hydrodynamic is used 
here to designate the action of tides and waves upon strand 
vegetation, and the action of stream and ocean currents in dis- 
tributing seeds. These factors may play an important paft in 
determining the peculiarities of plant societies, hence the neces- 
sity of keeping them in mind. 

Bioric Factors. Two plants cannot occupy the same soil 
at the same time. The struggle for a foothold in the soil may 
take place between species of the same kind or of different 
kinds. Again, the struggle may be between plant societies, as 
the forest and heath, or forest and prairie. The line along which 
two societies meet has been called the tension line. Here it is 
that the struggle is most pronounced. If the other ecological 


294 BOTANICAL GAZETTE [MAY 


factors remain constant, the tension line does not change. In 
that case, for instance, the forest does not advance on the heath 
nor the heath on the forest. But, as will be shown in the dis- 
cussion of the historical factors, the conditions as a rule are 
changing constantly. Not only may the struggle be between 
the forest on the one hand and some other type of plant society 
on the other, but it may be between different kinds of forests. 

The struggle between plants and animals may have an impor- 
tant bearing on the explanation of any given floral region. In 
civilized communities man has changed the whole nature of the 
vegetation. His influence is seen where forests are cleared for 
timber and fuel, or for purposes of cultivation; also indirectly 
where forest and prairie fires are active. Again in regions where 
stock is raised trees are damaged or kept from spreading by the 
ravages of domestic animals. Insect life may also be an impor- 
tant factor in explaining the floral character of a given region. 

In concluding this discussion of the ecological factors, it is 
well to note again that the three great physical media—soil, air, 
and water—are all influential in bringing about certain plant 
physiognomies. These, together with the biotic factors, make 
that variety in the landscape of any region which is shown in 
the plant societies that are present. 

C. Historical factors. The third great set of factors that play 
a part in the understanding of plant associations has been desig- 
nated historical, for it involves the element of time. It means 
simply that the geological and physiographic forces have 
changed and are changing the factors so far considered as to 
make absolutely necessary a readjustment of plant formations 
and plant societies to meet the new conditions. Diastrophic 
movements may submerge large areas of land and thus destroy 
all terrestrial vegetation or force it to migrate to unsubmerged 
parts. On the other hand, when there is an addition to the land 
area a condition is obtained where there can be noted the suc- 
cessive stages in the reclamation of such an area by vegetation. 
Again, in the last glacial epoch the ice sheet in its advance des- 
troyed vegetation and modified the climate beyond the limits of 


1901] GENETIC DEVELOPMENT OF FORESTS 295 


its advance, so as to bring about an arctic vegetation where there 
formerly existed a temperate floral formation. 

Erosive forces also come into play. The advance and retreat 
of the ice sheet changed the physiographic features. Drainage 
lines were dammed up and lakes were formed. It is the belief 
of glaciologists that the great lakes, at least in part, were 
formed in this way. Oscillations in the change of the front of 
the ice sheet changed the levels of lakes and left old beaches, 
other sand formations, and cliffs, thus bringing into existence 
new topographic forms and new soil conditions. With the final 
retreat of the ice sheet the normal processes of water erosion 
began again. Drainage lines became reestablished, and with the 
aid of decaying vegetation the lakes began to be filled up. The 
longer an area of land has been free from the ice sheet, the more 
nearly have its drainage lines become firmly established. It is 
evident then that the edaphic conditions in a region recently 
vacated by the ice will be decidedly different from those that 
have been exposed to the erosive force of water. The more 
nearly a region reaches base level the more stable will be 
the edaphic conditions. The recent work of Cowles’ on the 
Chicago region shows clearly the successive stages in the 
advance toward the climax condition. He was also the first ® 
to bring out clearly the dynamic conditions due to physio- 
graphic changes. It is evident that the existing plant societies 
must not be looked upon as fixed, but rather as changing from 
year to year; sometimes slowly, as in the case of swamps; or 
sometimes with rapidity, as in the case of dune societies. 


II. THE SAND SOCIETIES. 


The islands of North Manitou and Beaver at the northern 
end of Lake Michigan, the adjoining mainland on the Michigan 


LEs, H. C.: The physiographic ecology of Chicago and vicinity; a study of 

the i, development, and classification of plant societies. Bor. GAZ. 31: 73-108, 
145-1 Igol. 

“cows s, H. C.: The ecological relations of the vegetation of the sand dunes of 

e Michigan Bot. Gaz. 27:95-117, 167-202, 281-308, 361-391. 1899. /dem: 

The physiographic ecology of Northern Michigan. Science 12: 708, 709. 1900. 


296 BOTANICAL GAZETTE [| MAY 


side of the lake, and the southern shore of Lake Superior in the 
neighborhood of Marquette, Michigan, offer exceptionally good 
fields for a study of the stages in the life history of the forest 
societies. North Manitou island, in general, consists of a clay 
core covered over and added to by sand formations, principally 
beach and dune. So complex are these in the interior that it is 
difficult to trace any definite time relations, and no attempt was 
made to do this. On the south and southeast ends of the island 
however, where the beaches and dunes are in the process of for- 
mation, it is not so difficult to discover the time relations. The 
northern portion of Beaver island and the mainland in the 
vicinity of Charlevoix, Michigan, have similar formations. On 
the southern shore of Lake Superior, while the beaches are pres- 
ent, the dunes are for the most part absent. The beach and 
dune societies have been so thoroughly studied by Cowles that 
attention need only be called to the points in his work essential 
to the understanding of the various stages in the life history of 
the forest of these regions. 

THE LOWER BEACH. The lower beach is defined as that por- 
tion washed by the summer waves. There are a number of fac- 
tors here, any one of which may exclude vegetation. Since the 
beach is the shore drift in transit, the very movement of the 
sand or pebbles may prevent plant life. The mechanical wash 
of the waves would uproot any plants that had obtained a foot- 
hold. Again, the border between the land and water presents 
at one time a xerophytic and perhaps the next moment a 
hydrophytic habitat. However, in protected places, or where 
the water is shallow for a distance from the shore, the breaker 
line thus being distant from the shore, a condition is obtained 
which may bring into existence a hydrophytic society. Wher- 
ever sand is accumulating along the shores in the region under 
discussion, the lower beach is prominent. With a further reces- 
sion of the lake it may pass into the middle beach. It is con- 
sidered, therefore, the first stage in the life history of the forest, 
for by the action of the waves a soil is made, the first requisite 
for any forest society. 


1901] GENETIC DEVELOPMENT OF FORESTS 297 


THE MIDDLE BEACH. The middle beach is that portion of 
the shore drift in transit only during the winter months. It is 
consequently free from the waves during the growing season. 
Here, therefore, is a new land formation in which all the factors 
which prevent growth in the lower beach are eliminated for the 
brief space of one growing season. While the hydrodynamic 
factors have been eliminated, certain of the atmospheric, edaphic, 
and historical factors are detrimental. Since the former are 
considered in the initial stages of the upper beach society, they 
will be discussed under that head. In the short period that this 
beach is left untouched by the wave action, no plants of any 
consequence can obtain a foothold. This, together with the 
sterile soil of sand or pebbles and the extreme exposure to 
insolation, often prevents the growth of any species. In a few 
instances the annuals, Cakile Americana and Corispermum hyssopt- 
folium, and the first year’s growth of the biennial, Oenothera 
biennis, are present. If the new land thus made contained an 
alluvial soil stocked with seeds, one might expect a more or less 
abundant growth of annuals, though the insolation would still 
be very great, and would undoubtedly prevent a rich growth. 
Since the middle beach, by a further recession of the waters of 
the lake, may become a fossil beach, and thus reach, as will be 
shown later, a condition more near the forest, it is regarded as 
the second stage in the life history of the forest. 

FossiL BEACH. The fossil beach is that portion of the shore 
beyond the reach of the hydrodynamic factors. In the initial 
stages the fossil beach is as barren of vegetation as the middle 
beach, and if in an exposed situation the wind (fig. 1) shifts 
the sand repeatedly, this condition is maintained for a greater 
or less length of time. If fairly well protected from winds, 
successive plant societies appear rather rapidly. 

The well-known physical and chemical qualities of sandy soils 
need not be discussed at length here. While the upper layers 
of sand dry out rapidly, observations seem to show that the 
lower layers are moist up to high levels. However, the first few 
layers are extremely dry after a short period of drouth, due to 


298 BOTANICAL GAZETTE [MAY 


the rapid evaporation from the loose soil. The soil heats up 
and cools off rapidly, so that the variation in temperature in a 
short space of time is great, a condition unfavorable to plant 
growth. Nevertheless, in spite of the strong insolation and dry 
soil, if sufficient time elapses a definite fossil beach society is 
obtained. The enumeration of species is not of any consequence. 
They are at first mostly herbs, both annual and perennial. By 
their death, and in some instances by the decay of old logs left 
by storms when the beach was nearer the lake, humus conditions 
of the soil are gradually obtained. This is at first slow because 
of insolation. Nevertheless, each year adds its growth of herbs, 
and even though a greater part is dried out or blown away, in 
time the better humus condition will bring about the possi- 
bility of a higher type of plant society, so the fossil beach 
society gradually develops into a heath. 

Heatu. The heath society is considered, in the normal 
development, the fourth stage in the life history of the forest. 
It will be well to keep in mind the changes that make possible 
the heath. These are the increase in the amount of humus and 
the element of time. Juniperus communis, J. Sabina procumbens, 
Arctostaphylos Uva-urst, Pleris aquilina, Zygadenus elegans, Solidago 
nemoralis, Campanula rotundifolia, and Comandra umbellata are 
some of the characteristic plants of the heath. /ig. z shows a 
heath at the foot of a coniferous forest on the lake side of a 
sand dune. 

Now for the first time biotic factors assume some importance, 
though slight. In the first three stages the struggle of plants 
against adverse physical conditions gave plenty of room for all 
able to survive. Although the physical conditions are still 
unfavorable, in places plants like the junipers and bearberry 
may occupy considerable areas to the exclusion of others. In 
the open spaces between the individual shrubby plants, however, 
there is sufficient room for a large number of herbs. These by 
decay contribute humus to the soil, and since the insolation is 
not so great as in previous societies, a larger portion of the 
decaying vegetation is prevented from drying out. Thus as 


| 


igor] GENETIC DEVELOPMENT OF FORESTS 299 


the vegetation increases there is a gradually increasing incre- 
ment of humus, and this constitutes a condition favorable to a 
higher type of plant society. 

THE CONIFEROUS FOREST. It is not always easy to determine 
the conifer that first comes to occupy a place in the heath. If 
conclusions can be based upon a limited number of observations, 
the jack pine (Pinus Banksiana) is the most xerophytic (fig. 2). 
This is followed closely by the red pine (P. vesinosa) and the white 
pine (P. Strobus) in the order named. This does not mean that 
anyone of these always appears to the exclusion of all others, 
though that is sometimes the case (fig. 2). Other things being 
equal, the more xerophytic the conditions the more likely is it 
that the jack pine will be the predominant tree, and the less 
xerophytic the conditions the more abundant the white pine. 
fog. 3 is a photograph of a pine forest in which the red pines 
form a large proportion of the trees. In this forest are a num- 
ber of white and jack pines. If the succession as given above 
be correct, it means that this forest is probably a transition 
between the jack pine society and a white pine society. In the 
ideal case, then, the succession of predominant species is in the 
order named. The drier and perhaps the colder the climate up 
to a certain point, the more likely is this succession to be real- 
ized. Thus, in the Marquette region almost pure jack pine for- 
ests are found, while at North Manitou island, where the climate 
is more uniform, the jack pine stage is almost eliminated. Very 
often the balsam (Abies balsamea) (fig. 1) and hemlock ( Zsuga 
Canadensis ) occupy a prominent place in the coniferous forest. 

The first stages of the pine forest society are seen in the appear- 
ance of a number of trees in the later stages of the heath. For 
the first time in the normal development of the plant societies 
the tree has to be taken into consideration. In the tension zone 
between the heath and forest the trees are more scattered near the 
heath, and become gradually thicker as the forest is approached. 
The territory between the trees, where not shaded, is usually occu- 
pied by a growth of the heath plants already named. Thus, 
gradually the pines advance on the heath, and in time come to 


300 BOTANICAL GAZETTE [MAY 


occupy its territory. In the young pine forest near the tension 
zone there are often found scattered trees of the white birch 
(Betula papyrifera), oak, and the poplars (P. tremuloides and P. 
grandidentata). As the forest reaches maturity, these are usually 
crowded out, the birch being the last to disappear. 

Thus, with time and with the gradual increase of humus, the 
pine forest society is established. As a rule, the white pine 
seems to occupy the tension zone between the coniferous and 
the deciduous forests. If the climate be unfavorable for a higher 
type of forest tree, however, a stage beyond the white pine for- 
est is not reached. If the forest be not too open, and if surface 
fires are absent, the deep shade established by the pines means 
that the edaphic and atmospheric factors have become favorable 
for establishing the climax forest of the region. 

MAPLE, BEECH, AND HEMLOCK FOREST. Beginning with the 
middle beach we have seen that in each successive society a 
gradually increasing quantity of humus has been added to the 
soil, until a white-red pine forest is established. The pines add 
their quota of vegetable decay to the humus. The xerophytic 
soil thus becomes more and more mesophytic, and pari passu it 
has become more and more shaded. The humus means that the 
almost sterile soil of the beach has become richer in organic 
compounds, and by the aid of fungi and bacteria more nitrates 
are added. The capacity of the soil to hold water is likewise 
increased. The better edaphic and atmospheric conditions 
make a natural nursery for the growth of beech (Fagus fer- 
ruginea) and maple (Acer saccharinum Wang.) seedlings. The 
white pine does best in the richer soil conditions, but its seed- . 
lings require more light than they can obtain in the shade 
of the forest. As the maple and beech attain maturity, they 
come to occupy a prominent place in the forest, and a mixed 
pine-deciduous forest is the result. The broad-leaved decidu- 
ous trees produce a still deeper shade, so that if any pine seed- 
lings are able to survive in the pine forest, the increasing shade 
will finally make it impossible for a new generation of pine trees. 
The development of beech and maple seedlings, however, is not 


a ae 


Pe ee 


1901 | GENETIC DEVELOPMENT OF FORESTS 301 


affected by the shade of the parent trees. In the midst of the 
deciduous forest isolated white pines are often seen rising above 
their neighbors, and thus stand as relics of a previous pine for- 
est. A pine starting from a seedling condition with young 
deciduous trees as a rule never reaches a height greater than 
that of the trees among which it is growing. At the same time 
the girth of such a tree is much greater, owing to the better soil 
conditions, than it would be were it growing among its kind, 
where it must necessarily grow taller or be crowded out in the 
struggle for light. 

The place of the hemlock in the pine and deciduous forests 
is a peculiar one. It often composes 10 per cent. of the trees 
in a white pine forest, and may reach as high as 50 per cent.” It 
seems to be of about the same age as the trees with which it grows, 
and hence probably started with them. This probably means that 
its seedlings are able to develop like those of the pines, without 
being shaded. Yet it often occupies a prominent place in the 
deciduous forest (fig. g), even when the shade has become so 
dense as to exclude any chance for the development of white 
and red pine seedlings. It apparently occupies this position 
among the maple and beech because its seedlings can endure 
shaded conditions, although they do not thrive in such situations. 
Dwarfed young trees fifteen or more years old are often found 
in the densest shade possible. These develop slowly, until an 
open place is made by the death of the older trees around them, 
when they spring rapidly into prominence and occupy a position 
equal to that of the maple and beech. Like the white pine, the 
hemlock will flourish in the more xerophytic places, but prefers 
the deciduous woods and does best in them. The balsam also 
occupies a position similar to that of the hemlock, although it is 
seldom present in a mature climax forest. The yellow birch 
(Betula lutea) has a constant place with the beech, maple, and 
hemlock. It is more plentiful in the Marquette region than far- 
ther south, but seldom constitutes more than a small percentage 

7SPALDING, V. M., and FERNOW, B. E.: The White Pine Bull. 22. U. S. 
Department of Agric., Div. of Forestry. 1899. 


302 BOTANICAL GAZETTE [MAY 


of the total number of trees. The beech is absent in the 
Marquette region, and the significance of this is not clear. It 
may be due to the fact that the climate is not favorable for its 
development. It is present, however, in the northern peninsula 
east of Marquette. 

The whole interior of North Manitou island (fg. z), except 
in clearings and undrained areas, is covered with a mature maple- 
beech-hemlock forest. The presence of seedlings and young trees 
of these three in abundance and the absence of all other young 
trees in their shade indicate that the future forest growth will 
be the same as the present. The climax forest in places reaches 
nearly to the shore of Lake Michigan, restricting the coniferous 
and heath societies to very narrow belts. If the present shore line 
should remain constant, and if the natural succession of plant 
societies were not interfered with by man, undoubtedly the whole 
island would in time become completely covered with a 
deciduous forest save a narrow strip, the last remnants of a 
coniferous forest, next the water’s edge. 

The undergrowth in the dense shade of the maples, beeches, 
and hemlock is scanty. Taxus Canadensis and Mitchella repens 
are usually the most abundant. The spring plants are character- 
istic, but their vegetative period is confined to the leafing time 
of the trees under which they grow, so that by midsummer 
only traces of the many forms survive. The loose sandy soil is 
favorable for creeping underground stems. Lianas are entirely 
wanting. This is probably due to the lack of sufficient light and 
heat, for in the open woods farther south the liana habit is com- 
mon, and as one approaches the tropics the increasing warmth, 
even in dense shade, favors a luxuriant growth of lianas. 
Epiphytes, except mosses, liverworts, and lichens growing on the 
bark of trees, are also absent. 

Aspidium spinulosum intermedium, A. marginale, Lycopodium 
lucidulum, L. inundatum, Actaea alba, Goodyera pubescens, Osmor- 
rhiza brevistylis, Maianthemum Canadense, Monotropa uniflora, 
Corallorhiza odontorhiza, and Epiphegus Virginiana are among 
the most common plants that occupy a place in these forests. 


SE cnet 


1901 | GENETIC DEVELOPMENT OF FORESTS 303 


The humus in the soil is usually variable in amount, but is 
most plentiful where fallen logs have decayed. Hummocks of 
soil rich in humus mark places where dead trees when uprooted 
brought with them quantities of earth. The decaying logs are 
covered with fungi and mosses. Indeed the soil itself is full of 
a living mass of hyphe. The recent researches of Stahl® 
show that mycorhiza is the exception rather than the rule. The 
role that mycorhiza plays is problematic. Some plants, the 
European beech among them, are entirely dependent upon these 


.root-fungi, for when grown in sterile soil they perish. This 


seems to indicate that the beech cannot thrive in soil without 
humus, for its root-fungus is dependent upon organic decay for 
its existence. In other words, the absence of humus means the 
absence of mycorhiza, and its absence excludes the beech. 
Indeed, future investigation may show that a large majority of 
forest trees and other forest plants have established an obligate 
mutualistic relation with root-fungi. 

THE DUNE sociETIES. In discussing the sand societies the 
conditions favorable to the formation of dunes have been dis- 
regarded. Where the wind sweeps across the fossil beach the 
succession of plant societies is retarded. The extreme exposure 
to the xerophytic influence of the wind often prevents the pres- 
ence of plants. Nevertheless, when not too severe, a definite 
beach society is finally attained. The origin and development 
of dunes and dune societies have been clearly shown by Cowles.? 
It is only necessary to emphasize the fact here that the plant 
succession on dunes is similar to that on the beaches, and that 
in time the climax maple-beech society is reached. Fig. 1 is 
from a photograph, showing a fossil beach society in the fore- 
ground. On the right is a dune clothed with a coniferous forest. 
At the base of the dune is a heath encroaching upon the beach. 
On the landward side of this dune there are indications of the 
beginnings of a maple-beech forest, and farther inland are dunes 

*STAuL, E.: Der Sinn der Mycorhizenbildung. Jahrb. f. wiss, Bot. 34 : 539-668. 
1goo. 


°Sand dunes of Lake Michigan, Bor. Gaz. 27:95 ff. 


304 BOTANICAL GAZETTE [ MAY 


which have passed through the first stages into mature climax 
societies. Dunes clothed with maple-beech forests are also 
found along the Michigan coast in a number of localities. It is 
difficult to understand how such a mesophytic plant society is 
attained, unless it be assumed, as observation seems to indicate, 
that by capillarity, water may rise from the ground water level 
to considerable heights in sandy soils. 


lll, THE CLAY SOCIETIES. 

There are greater difficulties in tracing the life-history of 
clay societies than of sand societies, principally because there is 
not such a perfect sequence of clay formations at the present 
time as of sand formations. With the exception of sea-cliffs 
and clearings, clay soil formations free from vegetation date 
back to the time of the retreat of the last ice sheet. Sea-cliffs, 
however, formed by the action of the waves on glacial clay, are 
a common feature along the shores of Lake Michigan. Here the 
waves are constantly undermining the cliff, and thus prevent 
anything more than a transient vegetation. But should the 
débris at the foot of the cliff become too great for the shore 
current to transport, or should a recession of the waters of the 
lake leave a cliff beyond the reach of the waves, the undermin- 
ing would cease. Here then at the outset is a clay bluff almost 
free from vegetation. 

The reclamation by plant growth begins at once. The first 
stage in the life-history is represented by herbaceous plants like 
Solidago humilis, Aster laevis, Elymus Canadensis, Prenanthes alba, 
etc. Later on xerophytic shrubs like Shepherdia Canadensis and 
Juniperus communis give a heath-like physiognomy to the bluff. 
Again, a third stage is reached in the appearance of conifers, 
poplars, and white birch. These stages are more rapid, for clay 
soil retains moisture more readily than the sandy soils. The 
action of humus on clay is exactly the reverse of its effect on 
sand, for it loosens up the soil and renders it much less soggy 
than it would be wére it free from organic decay. Where the 
cliff has been long enough freed from the action of the waves, 


1901 | GENETIC DEVELOPMENT OF FORESTS 395 


in due time a maple-beech-hemlock condition is the result. 
Thus, starting with an herbaceous vegetation similar to that on 
the fossil beach, there is a transition through the heath and 
coniferous forest to the climax society. Making due allowance 
for difference in climate, it is probable that all the glacial clay 


- I.—The lakeward slope of a sand dune on North Manitou island covered by 
a forest. Balsam is the most common tree. The undergrowth is very dense, and the 
deep shade furnishes favorable conditions for the development of maple and beech. 
The evergreen forest is encroaching on the evergreen heath seen at the foot of the 
slope, and the latter, in turn, is encroaching on the fossil beach society situated in the 
foreground. : 


lands in the region under discussion have passed through some- 
what similar stages, since they were formed by the action of the 
last ice sheet. As a rule, where the vegetation has not been dis- 
turbed by man, they are clothed with maple-beech-hemlock 
forests. Where clearings are made, as will be shown subse- 
quently, they rapidly attain the same forest condition if let 
alone 


306 BOTANICAL GAZETTE [MAY 


IV; (fHE- ROCK. SOCIETIES: 


The pre-Cambrian rocks of the Marquette region furnish an 
excellent field for the study of the succession of plant societies. 


2.—Jack pine forest on a fossil beach 
near Manes e. The — of undergrowth 
is due to repeated surface fire 


The rocks are mostly gran- 
ites and quartzites. The 
chemical nature of the rocks 
seems to have little influ- 
ence on the ultimate plant 
society that is obtained, but 
the ease with which they 
disintegrate is different, 
therefore the rapidity of the 
succession of plant growth 
is influenced. Other things 
being equal, granites, be- 
cause of their heterogene- 
ous structure and conse- 
quently differential weath- 
ering, will furnish a soil 
more quickly than the 
homogeneous quartzite. In 
the region under discussion 
the rocks have not only 
been worn and_ polished 
smooth by the action of 
the last ice sheet, but since 
then have lain beneath the 
waters of the former exten- 
sion of Lake Superior. In- 
deed, some of them have 
only recently emerged from 
the lake. Thus just as 
there are beach lines of dif- 
ferent ages, so there are 
rock areas of different ages. 
Other things being equal, 


1901 | GENETIC DEVELOPMENT OF FORESTS 3°97 


the longer the rocks have been subject to sub-aerial influences 
the more nearly they approach the condition of the climax 
plant society. 

On the sand formations at the contact of the lake with the 
shore the hydrodynamic factors absolutely prevent plant growth. 


This forest alse 


—Norway pine forest on an old beach near Marquette. 
pontithe. a top white and jack pin 


Here likewise, where the rocks dip beneath the lake, there is a 
zone of no vegetation. However, often within the reach of not 
too active waves lichens are found. Many lichens need no soil ; 
such are essentially lithophytes and are soil makers. They are 
fastened firmly to the rocks by. holdfasts which. secrete acids 
that aid in disintegrating the rocks on.which the lichens grow. 
Lichens receive moisture from the rain that falls on them, or, 
when near the water’s edge, from the wash of the waves. 
Between rains they dry and curl up, but are uninjured, for after 


308 BOTANICAL GAZETTE | MAY 


the next rain they are as fresh as ever. Nitrogen compounds 
are obtained from the rain water or from particles of dust which 
lodge around them and other requisite inorganic compounds come 
from the rocks. Thus these pioneers of vegetation, as has been 


Fic. 4.—A maple-beech-hemlock forest in sandy soil 6n North Manitou island. 
he diene consists principally of maple, beech, and helmlock seedlings and 
young trees; also Taxus Canadensis and Mitchella repens. The young trees indicate 
that the next generation of trees will be the same as the presen 


shown by various writers, are able to flourish in the most 
xerophytic situations. They not only aid in disintegrating the 
rocks, but by decay furnish humus constituents to the soil, and 
in these ways, pari passu, both organic and inorganic soil is 
made. : 


1901 | GENETIC DEVELOPMENT OF FORESTS 309 

If the lichens be on not too steep a slope they will in them- 
selves furnish soil enough to maintain a higher form of plant 
life. Xerophytic mosses gain a foothold as soon as a slight soil 


is made. These, too, by decay aid the lichens in forming a soil 


1 Cladonia rangiferina, Moss, and 


Fic. 5.—A granite rock covered by the licher 
herbaceous plants. To the right are conifers growing in the crevices of rocks 
that will support still higher forms. /ig. 5 shows such a society. 

It is even possible, in time, without the aid of inorganic proc- 

esses of weathering to establish a soil capable-of supporting a 
tree vegetation. However, inorganic processes of weathering 

4 . 7. > 1 } Z 

are going on continually. Changes of temperature, etc., crumble 
the rocks and form crevices in which soil lodges. Since most of 


310 BOTANICAL GAZETTE [MAY 


the rock surfaces are not level, the soil made by lichens and 
mosses is washed into the crevices or into hollows made by the 
action of ice or water erosion. At once a herbaceous and 
shrubby vegetation springsup. Plants like huckleberries, golden- 
rods, Potentilla tridentata, Campanula rotundifolia, and the bear- 
berry gain a foothold in the crevices. These are followed by 
arbor vitae (Zhuwja occidentalis), the junipers, and the pines. ig. 
6 shows a rocky island in a bay near Marquette. At a distance 


Fic. 6.—A granite island in Lake Superior near Marquette. At a distance it 
appears destitute of ~~ growth. 


it looks barren of vegetation, but a closer view (jig. 7) shows 
that it is not altogether destitute of plant life. /%g. 8 is from a 
photograph of a larger island near by which shows a much higher 
type of vegetation, for here are found red and white pines. The 
weathering process is greatly aided by the prying action of roots. 
fig. 9 shows a red pine whose roots have split and raised a 
granite rock. 

One of the noticeable features of rock vegetation is its 
variety. Within the limits of a few square rods miniature repre- 
sentations of all the great plant societies are found. Near a 
roche moutonnée, as smooth and as barren of vegetation as the 
day the ice sheet left it, is a small depression containing a 
swamp, in which sphagnum moss and the cranberry are found ~ 
growing. Near by is a lichen society; this may grade into a 


1901 | GENETIC DEVELOPMENT OF FORESTS 311 


heath such as is shown in fig. 5, and this in turn may border on 
a small group of pines. 

As in the sand and clay series, the mesophytic forest is not 
reached until first preceded by a coniferous forest. The conif- 
erous forests on the rocks are at first very open, but ultimately 


On the faces of 


Fic. 7.—A near view of a portion of the island shown in fg. 6 
In the crevices are a number of herbaceous 


the rock are shown lichens and mosses 
arbor vitae, huckleberry, 


and woody plants; among them are goldenicld. poplar 
ninebark, Juneberry, wild red cherry, Campanula rotundifolia, and Potentilla trt- 
di i 

denaata 


the gaps are filled up, sometimes with birch and poplars, and 
thus a condition is obtained for the maple-hemlock torests. 
Some rock hills near Marquette illustrate these points. /ig. ro 
is a view from Sugar Loaf. The top of this hill is almost barren 
of vegetation. Obviously this would be the case, for the soil 
that is formed descends ‘to lower levels; in the case of steep 


312 BOTANICAL GAZETTE | MAY 


slopes gravity alone is sufficient, while on all slopes the trans- 
porting power of rain water comes into play. Usually at the 
foot of the hill and sometimes well up the slope the conditions 
are favorable for the climax society. Other hills (fg. 77) 


as 


catia oa” 


i 


Fic. 8.—Portion of an island near the one shown in figs. 6 and 7. Here the 
vegetation is further advanced. Besides lichens, etc., white and Norway pines are 
found. 


farther in the interior are completely covered with pine and 
deciduous forests. 


Vv. THE SWAMP SOCIETIES. 

One of the most characteristic features of a young glacial 
topography is the large number of lakes. In general these may 
be divided into two classes according to their mode of origin. 
One class is due to the action of the ice sheet. Depressions 


IgoT | ° GENETIC DEVELOPMENT OF FORESTS 313 


may be made directly by the moving ice, or glacial deposits 
may dam up drainage lines. A second and subsequent class of. 
lakes is made by bars cutting off lagoons from the larger lakes, 
or by spits formed across their embayments. Both classes 


have a_ similar _life- 
history. Some find out- 
lets and ultimately pass 
out of existence through 
the normal stages in the 
life-history of a river. 
A larger number, how- 
ever, never find outlets, 
but are silted up by the 
wash of the surround- 
ing soil, and by the ac- 
cumulation of vegeta- 
tion. Thus swamps are 
formed, and since they 
may represent a stage 
in the life-history of 
the forest they deserve 
treatment here. 

The zonal distribu- 
tion of plants in swamps 
is one of the stock illus- 
trations of ecologists, so 
that development of the 
various zones need only 
be mentioned briefly. 
Most swamps start as 
ponds or lakes, in which 
water lilies and other 
pond plants dominate. 
On the borders of the 
pond sedges appear ; 
these, by their decay, 


Fic. 9.—A Norway pine growing from a crevice 
in granite ; the rock has been cracked and uplifted 
by the growth of a root. 


314 BOTANICAL GAZETTE | MAY 


build up a soil, and thus prepare the way for the next zone, the 
Cassandra-sphagnum vegetation. The sedges encroach farther 
on the original lake. The Cassandra-sphagnum zone makes 
conditions possible for a tamarack-spruce zone (fig. 77). Thus 
each successive zone is pushed farther and farther toward the 
center. Finally the lily center disappears, and then successively 


Fk —A general view of granite rock vegetation as seen from a granite hill 
(Sugar Loaf) near Marquette. On the margin of the lake the rocks show the first 
stages in the life-history of a forest. As the foot of the hill is approached the vegeta- 
tion becomes more and more mesophytic until a mixed conifer and deciduous forest is 
attained. Probably in places this condition is reached more rapidly because of the 
presence of glacial drift. At the top of the slopes the xerophytic condition of the lake 
border is again attained, 


the sedge and Cassandra zones, until a tamarack forest may 
come to occupy the whole territory. 

Attention has already been called to the probable factors 
unfavorable to a high development of plant life in these swamps. 
These are due in the main to undrained conditions. The accumu- 
lation of humic acids may Cause, osmotically, a drying-up effect. 
Insufficient aeration of the soil prevents a healthy growth of the 
root system of highly organized land plants, and also bars the 


T1901] GENETIC DEVELOPMENT OF FORESTS 315 


presence of nitrifying bacteria. These probably bring about the 
xerophytic structures of plants so commonly seen in hydrophytic 
habitats. 

The areas around the swamp, whether clay, sand, or rock, 
have been undergoing their normal changes, so that the swamp 
is eventually surrounded by a forest. The swamp is finally built 


IG. 11.—A lagoon cut off from Lake Superior by a beach line, resulting in an 
undrained swamp. In the foreground is seen a portion of the original lagoon; back 
of this is the sedge vegetation, and near the conifers is a Cassandra zone. The coni- 
fers are mostly tamaracks (Zarix Americana). 


up high enough above the water level to permit a higher type 
of tree to occupy the area. Thus the tamarack is gradually 
crowded out, and a climax forest is the final stage. 

The life-history is not always as indicated above. , The 
Swamp may become partially drained, but still be too wet to 
Support the highest tree society. In that case the abor vitae 
gradually replaces the tamarack. Associated with the arbor 
vitae are the ash ( Fraxinus sp.), the balsam, the white pine, and 
the yellow birch. Gradually, however, the arbor vitae swamps 


316 BOTANICAL GAZETTE [MAY 


give away before the encroaching mesophytic forests. ig. 12 
shows a hemlock-maple forest bordering an arbor vitae swamp. 
In wet springy soils the arbor vitae forest is also found. fig. 73 
shows a young forest in such a situation. 


VI. CLEARING SOCIETIES. 

Thus far what has been called the normal life-history has 
been traced. The influence of human agencies, with few 
exceptions, has been disregarded. By far the greater number 
of areas studied show unmistakable signs of the devastating 
influence of man, though less so on the islands visited than on 
the mainland. These clearing areas cannot be disregarded if a 
complete history of forest development is to be written. At first 
the difficulties of getting any order out of the tangle seemed 
insurmountable, and the notes were usually headed ‘‘clearings, 
conditions artificial.” The literature on the subject of plant 
succession in clearings is confusing. It consists, as a rule, of 
records of isolated observations without any attempt at correla- 
tion, That there is some solution to the problem was the writer's 
firm belief, and after collecting a large number of notes the study 
began to yield results. 

If the factors controlling the normal development as given 
above are correct, they ought to dominate in the clearing socie- 
ties. With two exceptions, the stages observed in the develop- 
ment of these clearing societies are represented in the normal 
history. These exceptions are the “fireweed” and the poplar- 
birch societies. To explain these it was found necessary to 
give prominence to the fact that some plants migrate more easily 
than others. 

If a deciduous forest be destroyed and burned over so that 
the surface débris is partially reduced to ashes, it will usually be 
found that the humus conditions of the soil are then reduced 
toward, but not necessarily to the conditions obtained in the 
heath. The equilibrium that had been established has been dis- 
turbed, and other things being equal all plants will have an equal 
chance provided their seeds are present. Shade plants, however, 


1901 | GENETIC DEVELOPMENT OF FORESTS ii te 


including young plants of beech and maple, are excluded 
because the insolation is great. Some few plants that seem to pre- 


fer more xerophytic conditions likewise will not thrive. Then the 


struggle is nar- 
rowed to those 
plants that can 
endure strong in- 
solation, that pre- 
fer tolerably 
good edaphic 
conditions, and 
whose seeds are 
present. It is ob- 
vious that those 
plants in the 
neighborhood 
that have the 
lightest seeds will 
have the largest 
representation of 
seeds on the 
ground first, and 
of these the most 
rapid growerswill 
prevail the first 
year ortwo. The 
so-called ‘fire- 
weeds’’ meet 
the requirements 
best. Epilobium 
angustifolium,Eri- 
geron Canadensis, 
and certain gold- 


Fic. 12.—A hemlock-maple forest bordering onan 
arbor vitae swamp. The tangled growth of the arbor vitae 
seen in the nagar The hemlocks ,are 


Swamp is 
encroaching on the arbor vita 


enrods spring at once into an occupancy of ‘the field. and there 
is given a “ fire-weed ” physiognomy to the clearing ( fig. 14). 
The poplars (P. tremuloides and P. grandidentata) and the 


318 BOTANICAL GAZETTE [MAY 


white birch are the trees which have seeds adapted for rapid 
migrations, and are also rapid growers. Because they are per- 
ennial, and can thus get an early start each year after the first, 
they soon overtop the fireweeds and gradually shade them out. 
Ostrya Virginica, Prunus Pennsylvanica, P. serotina, and Quercus 
yubra are in some places associated with the poplars and birches. 


—A spring hillside with young growth of arbor vitae. ‘Ibe hilf in the 
ig oe been cleared of a maple-hemlock forest and is used for a pasture. 


In the absence of pine trees in the vicinity to furnish pine seeds, 
this poplar-birch society is maintained until deep shade condi- 
tions are again obtained for the growth of the maple-beech seed- 
lings. Then, just as in the normal development the pines are 
replaced by the maples and beeches, so in the clearing develop- 
ment the poplar and birches are likewise replaced. The pop- 
lars go first, then usually the birch and hornbeam, and finally 
the cherry trees drop out. 

If, however, pine trees in the vicinity of a clearing can fur- 
nish seeds, the pines occupy a place in the life history. The 


Igor] GENETIC DEVELOPMENT OF FORESTS 319 


more unfavorable the humus conditions up to a certain limit, the 
more likely is the clearing to have a pine aspect. The humus 
in the soil may be so reduced as to be incapable of supporting a 
poplar-birch growth. fig. 15 shows a young jack pine growth. 


Fic. 14.—A hemlock -maple forest destroyed by fire. The rich growth of fireweec 
Aonapeel oe indicates the first stage in the life-history of a second- 
growth for, 


In this repeated fires have probably so reduced the humus con- 
ditions of the soil that the most xerophytic of the pines alone is 
able to gain a foothold. If favorable for the presence of both 
pine and poplar-birch growth (fig. 76), the pines in time over- 
top the poplars and birch, until a condition corresponding to the 
normal life-history is attained, viz., a pine society preceding the 
maple-beech society. 


320 BOTANICAL GAZETTE [MAY 


It can be seen readily that it is possible to obtain all stages 
of reduction between mesophytic and xerophytic conditions. 
This depends on what stage in the normal life-history is attacked 
by the fire, and on the intensity or entire absence of subsequent 
fires. Thus all possible combinations are represented. ig. 77 


Fic. 15.—A young jack pine forest on a fossil beach near Marquette. The 
humus content of the soil has been much reduced by insolation and repea ated fires. 
The undestroyed Norway pines in the background probably indicate that the beach 
had formerly attained the Norway pine stage in the life-history series. 


shows a maple-hemlock forest that has been nearly destroyed 
by fire. This is situated on the slope of a quartzite hill, and had 
probably reached the climax condition. In places the fire has 
left remnants of the forest, and even where the trees are 
destroyed certain geophilous forest herbs like Clintonia borealis 
have maintained themselves in the shade of half burned logs, 
while near by fireweeds, poplars, and pines are found. Fug. 18 
shows an almost pure birch forest. Not far away, however, are 


1901 | GENETIC DEVELOPMENT OF FORESTS 321 


found poplars mixed with the birch. In this neighborhood 
stumps of pines indicate the former forest. In places repeated 
fires have reduced the area to heath conditions. This is shown 
by a number of heath plants like Preris aguilina, Gaultheria procum- 
bens, Cladonia rangiferina, etc. The presence of few seed-produc- 


1G. 16.—A second-growth forest of pines and poplars on a fossil beach near 
Marquette. The pines are slowly gaining ascendancy over the short-lived poplars 
and these will in time disappear. 


ing pines in the neighborhood will probably account for the’ 
absence of many pine seedlings. 

Thus, no matter how far toward the first stage in the life-his- 
tory a forest is reduced, if not interfered with by human agencies, 
ultimately the climax forest society is reached, but not without 
first having passed through a pine or a poplar-birch stage. A 
maple forest was never observed to follow a maple forest except 


322 BOTANICAL GAZETTE [MAY 


when the stools of a cleared forest are capable of producing 
sprouts. These may in time reach tree dimensions, though 
usually not before a considerable number of pines, birch, or pop- 
lars spring up between them. 

Thus it will be seen that if human agencies should cease to 


Fic. 17.—A partially destroyed hemlock-maple forest on the slope of a quartzite 
hill (Mt. Mesnard) near Marquette. The forest had attained the climax stage in the 


life-history series. 


operate, all classes of topographic forms will ultimately come to 
be clothed with the climax forest growth. There is no reason 
why even the jack pine barrens may not in time reach this 
state. It must be remembered that these are late topographic 
features, or are reduced by repeated burnings to a soil poor in 
humus. 


ies) 


Tgo1 | GENETIC DEVELOPMENT OF FORESTS 32 


VII. CONCLUSIONS 


The life-history of the vegetation of four sets of physio- 
graphic formations has thus been traced. It has been shown 
that in each series the climax plant growth is a deciduous-hem- 
lock combination. These physiographic formations have been 
treated separately because usually they are distinct from one 


a 
M 
H 
f 
f 


ev 


IG. 18.—A white birch clearing growth on a fossil beach near Marquette. The 
undergrowth consists of Preris aguilina, Diervilla trifida, etc. 


another. In some instances, however, clay is mixed with rock 
débris or underlies the sand. The only effect such a mixture 
has is to hasten the succession of stages, for the advance toward 
the climax is more rapid on clay than on other soils. Princi- 
pally for this reason the clay soils on the whole are already 
covered with the climax forest. In many instances the sandy 
soils have likewise attained the last stage in the life-history. 
Especially is this so where free from the influence of man as on 


324 BOTANICAL GAZETTE [MAY 


North Manitou island. But the succession of plant societies is 
much slower on sand than on clay soil, hence the prevalence of 
pine forests on the former. The reasons for believing that ulti- 
mately these pine forests will give way to the deciduous have 
already been given. 

It is the belief of the writer that with some modifications the 
principles brought out in the foregoing pages will hold for all 
regions climatically capable of supporting a tree vegetation. 
The region under discussion is physiographically young. If the 
terms of physiography be adopted for ecology, a region having 
scanty vegetation may be considered young, while from this 
there will be all stages through maturity to old age, viz.,a meso- 
phytic climax forest. It would appear to the casual observer 
that northern Michigan is a region physiographically young and 
ecologically old. But no inconsiderable portion of the area of 
both Michigan and Wisconsin is occupied by swamps and lakes 
(including the great lakes). These will remain hydrophytic and 
hence ecologically young until filled up or drained. This con- 
dition is not attained until the region reaches maturity. The 
whole region, from an ecological standpoint, therefore, cannot 
reach maturity until it does so physiographically. Thus the 
development of the forest is in a measure held back until the 
normal physiographic processes extinguish the swamps and lakes. 
When these are eliminated, the region will reach old age from the 
vegetation standpoint much sooner than it does physiograph- 
ically, because the climate is favorable for the extension of meso- 
phytic forests into altitudes which would otherwise support only 
a xerophytic society. 

In an area where the climate is more severe for tree growth 
the life-history stages are less rapid, for even though the region 
may be nearer base level the climax stages are more restricted. 
For example, Cowles* has shown that in the Chicago region, as 
a rule, the river bottoms contain the mesophytic plant societies, 
while the clay hills have only attained a semi-mesophytic forest 

” Bot. GAZ. 31 : 88 ff. 1901. ; 


Igor | GENETIC DEVELOPMENT OF FORESTS 325 


of oaks and hickories. Such clay hills in northern Michigan are 
usually covered with the climax forest. 

Again, as one approaches the semi-arid regions of the West, 
he will observe that the forest growth on the hills becomes less 
and less mesophytic until finally it gives way altogether to the 
prairie society. At the same time, the river bottom forests also 
become less mesophytic and more restricted to the banks of the 
streams. Ultimately, they, too, in the region of the great plains 
pass from existence and the prairie reaches to the very margin of 
the drainage lines. The writer has made some studies at vari- 
ous places in Kansas in reference to these points, and it is his 
intention to discuss them in full in a future paper on the eco- 
logical relations of prairie and forest. 


THE UNIVERSITY OF CHICAGO. 


MEXICAN FUNGI. III. 
E. W. D. HOLWay. 

THE descriptions of the following new species were sent to 
me in German by Dr. Dietel. I am under many obligations to 
Professor Robinson of the Gray Herbarium, and to C. G. Pringle 
tor the determination of the host plants. A large collection of 
Uredineae yet remains to be studied. 


Uromyces Celosiae Diet. & Holw., n. sp.—Sori hypophyllous, 
sometimes on indistinct yellow spots, small or medium-sized, 
scattered or often thickly covering the leaf; uredosori cinna- 
mon-brown; uredospores elliptical or globose, sometimes obo- 
vate, 27-34 X 24-26; epispore thick, thinly covered with strong 
Spines, with two germ-pores, brown; teleutosori dark brown, 
teleutospores elliptical or globose, coarsely verrucose, chestnut- 
brown, apex with a light colored rounded cucullate or papilla- 
like thickening, 28-38 X 22-30u; pedicel hyaline, as long as the 
spore, or a little longer, easily separating at the base from the 
we ‘plant, swelling in water. 

“On Celosia latifolia; Oaxaca, October 17, 1899, no. 3641. 


Uromyces venustus Diet. & Holw., n. sp.— Aecidia hypophyl- 
lous, single or in small groups, hemispherical, opening at apex 
by a small pore; aecidiospores globose or elliptical, 20-25 X 20H, 
finely verrucose, teleutosori epiphyllous, opposite the aecidia, 
often in a circle, the center of which is the aecidium on the 
opposite side of the leaf, variable in size, sometimes confluent, 
naked, pulverulent, dark brown; teleutospores ovate or ellip- 
tical, sometimes almost globose, 32-40 X 20-28, chestnut-brown, 
apex light brown, beak-like; epispore thick, with longitudinal 
lines; pedicel about the length of the spore, thin, hyaline, easily 
breaking at the base from the host-plant. _ 

AAA { In Cestrum nitidum, Amecameca, October 31, 1899, no. 3759- Easily 
¢ 326 [Ae 


1901] MEXICAN FUNGI 327 


distinguished from U. Cestri Mont. by the papilla at apex, and by the lines 
on the epispore. 


Uromyces Oaxacanus Diet. & Holw., n. sp.—Sori epiphyllous, 
irregularly scattered (occasionally a sorus on the under side of 
the leaf), small, pulverulent, black; teleutospores ovate, ellip- 
tical or almost globose, frequently angular and irregular, 25-35 
x 18-24; epispore chestnut-brown, not thickened at apex, ver- 
rucose; pedicel hyaline, hollow, somewhat longer than the 
spore. 

On Jatropha urens, Oaxaca, October 21, 1899, no. 3690. 


Uromyces dolichosporus Diet. & Holw., n. sp.— Uredosori 
mostly hypophyllous, scattered, dark brown, partly with spermo- 
gonia on the upper surface of the leaves; uredospores obovate, 
dark brown, 33-47 23-27; epispore thickened at apex and 
often at the base, echinulate, with three germ-pores; teleuto 
sori of medium size, hypophyllous, rarely epiphyllous, scattered 
or in small circular groups, orange-yellow when fresh, becoming 
white and felt-like; teleutospores long-fusiform or long-clavate, 
45-65 X 12-18; with a thin hyaline smooth epispore, and ger- 
minating as soon as mature; pedicel firm, about the length of 
the spore. 


On Tournefortia velutina, Oaxaca, October 18, 1899, no. 3655. 


Uromyces Rubi Diet. & Holw., n. sp.—Spots circular, about 
in diameter, occasionally confluent, yellow ; sori epiphyllous, 
small, white when dry; uredospores elliptical, obovate, or glo- 
bose, 23-35 x 18-25; epispore hyaline, with stout spines ; teleu- 
tospores elliptical or clavate, 26-35 X17-24#, with a very thin 
hyaline epispore; pedicel short; spores germinating at once. 

On Rubus, Cuernavaca, May 17, 1898, C. G. Pringle. 


Tar 


Uromyces PoLyMNIAE (P. Henn.) Diet. & Holw.—Uredo is U. 
Polymniae P. Henn. Teleutosori scattered, hypophyllous, single 
Sori here and there on the upper surface, brownish black, naked ; 
teleutospores elliptical or almost globose to pyriform, sometimes 


328 BOTANICAL GAZETTE | MAY 


truncate at apex, 30-45X20-304; epispore smooth, strongly 
thickened at apex, brown; pedicel firm, tinted, up to 75 long. 

On Polymnia maculata ?, Rio Hondo cafion, near city of Mexico, October 
4, 1899, no. 3562; October 30, 1899, no. 3562B. On Polymnia maculata, 
Patzcuaro, October 10, 1899. The latter host shrubby, 10-15 feet high; the 
former herbaceous only, 4-5 feet high. 


Uromyces Indigoferae Diet. & Holw., n. sp.— Sori epiphyllous, 
sparingly hypophyllous, scattered or in little groups on dead 
brown spots, and also on the petioles and young fruit, small, 
naked, surrounded by the ruptured epidermis, dark brown; ure- 
dospores globose or elliptical with short spines and three germ- 
pores, brown, 20-25 X 18-23; teleutospores elliptical or globose, 
smooth, chestnut-brown, strongly thickened at apex, 22-30 
X 18-25; pedicel long and firm, hyaline. 


On Indigofera Mexicana, Oaxaca, October 23, 1899, no. 3722. 


Uromyces GALPHIMIAE Diet. & Holw.—Uredo on G. Humboltiana, 
Guadalajara, September 14, 1899, no. 3409. 

UromyceEs VIGNAE Barclay ?—On Vigna stroboliphora, Guadalajara, 
September 15, 1899, no. 3424. 

Uromyces TRIFOLIAE (Hedw.) Lev.—On Trifolium, Pachuca, October 
6, 1899, no. 3588. 

UROMYCES GLoBosus Diet. & Holw.—On Sapium biglandolosum, Cuer- 
navaca, September 28, 1899, no. 3517. This is the host plant of the original 
collection. 

Uromyces Caapir (Schw.) Farl.—On Arisaema macrospathum, Cuer- 
navaca, September 29, 1899, no. 3522. The 1896 specimens are on this host, 
and not on A. Dracontium. 

URoMYCES SOLANI Diet. & Holw.—On SS. appendiculatum, Amecameca, 
October 31, 1899, no. 3761. 

Uromyces AEGOPOGINIS Diet. & Holw.—On Aegopogon cenchroides, 
Tizapan, Va!ley of Mexico, September 27, 1899, no. 3506. 


Puccinia Berberidis-trifoliae Diet. & Holw., n. sp.—Spots 
brown, or blackish-purple; sori hypophyllous, firm, strongly 
pulvinate, black, about 1-3" in diameter, linear on the petioles; 
teleutospores brown, quite variable in form and size, ellipti- 
cal, oblong or fusiform, rounded at apex, or conical, or often 


1901 | MEXICAN FUNGI 349 


prolonged into a point, mostly rounded at base, little constricted, 
24-45 X 14-24, smooth, variously thickened at apex according 
to the form of the spore; pedicel rather longer than the spore, 
tinted, firm; one-celled teleutospores rather common. 

On Serberis trifolia, Rio Hondo, near City of Mexico, October 4, 1899, 
no. 3570. The Aecidium of Puccinia graminis occurs on some of the leaves. 


Puccinia Aniscanthii Diet. & Holw., n. sp.—Sori on both 
sides of the leaf, scattered, small; uredosori brown; teleutosori 
black, naked ; uredospores elliptical or obovate, 22-28 X 18-224, 
light brown, with distant short spines, and two germ-pores ; 
teleutospores elliptical, rounded at both ends, slightly con- 
stricted, 38-48 X 25-31, chestnut-brown, at the apex and often © 
on the lower cell a cucullate or papilla-like light-colored thicken- 
ing, verrucose; pedicel longer than the spore, hyaline, apd 
separating at the base, sometimes laterally inserted. 

On Antsacanthus, probably 4. Wrightii, near Acamboro, October 8, 1899, 
no. 3595. This species is much like Puccinia Ruelliae-Bourgaei Diet. & 
Holw., but that has almost all the pedicels laterally inserted, and the mem- 
brane is not thickened over the germ-pores. Puccinia Blechi Lagerheim 
appears to be very similar, but according to the description the teleutospores 
are differently colored, and also without the thickening over the germ-pores. 


Puccinia Ruelliae-Bourgaei Diet. & Holw., n. sp—Aecidia in 
irregular groups along the veins; cells of the pseudosporidia 
oblong ; aecidiospores elliptical or globose, often angular, 
25-38 X 20-26, yellowish, strongly verrucose, much thickened 
at apex; uredospores in these specimens few, with the teleuto- 
spores, brown, echinulate, 20-25; teleutosori hypophyllous on 
yellow spots, sparingly epiphyllous, scattered, 0.5—2™™ in diam- 
eter, black, pulverulent, naked; teleutospores elliptical, rounded 
at both ends, not at all or only slightly constricted, 35-41 x 27— 
324, dark chestnut brown, with large tubercles ; pedicel as long 
or longer than the spore, hyaline, rough, inserted at one side. 

On Ruellia Bourgaei, Chapala, September 20, 1899, no. 3471. Closely 
resembles Puccinia laterifes B. & R., but has much larger and darker 
teleutospores ; the aecidiospores are also larger and remarkable for their 
Strong apical thickeni ing. 


330 BOTANICAL GAZETTE | MAY 


PUCCINIA HETEROSPORA B. & C.—On Adbutilon crispum, Oaxaca, October 
20, 1899, no. 3684; on Anoda hastata, near Tula, September 21, 1898, no. 
3186; Cuernavaca, September 28, 1898, no. 3093; on Axoda, Patzcuaro, 
October 17, 1898, no. 3004; Orizaba, October 6, 1898, no. 3194; Chapala, 
September 22, 1899, no. 3477; on Anoda acerifolia, Oaxaca, October 18, 1899, 
no. 3656; Cuautla, October 12, 1898, no. 3044; on Szda Hol/wayt, Cuautla, 
October 12, 1898, no. 3043. 

PucciniA GALI (Pers.).— On Galium uncinulatum var. obstipum, Oaxaca, 
October 18, 1899, no. 3654. 

Puccin1IA SEYMERIAE Burrill.—On Seymeria virgata, Oaxaca, October 
22, 1890, nO, 3721. 

PuccinIA PITCAIRNIAE Lagerh.—On /Pitcairnia Palmert, Chapala, 
September 1899, no. A; Uruapam, October 11, 1899, no. 3619. These two 
' specimens are not exactly alike, the latter having somewhat smaller more 
regular spores less thickened at apex. 

PUCCINIA MENTHAE Pers.—On Monarda?, Jalapa, October 3, 1898, 
no. 3211. 

PUCCINIA PHILIBERTIAE E. & E.—Tizapan, bs of Meixco, October 
28, 1899, no. 3749, on Metalstelma angustifolium. 


Puccinia Marsdeniae Diet. & Holw., n. sp.—Sori on yellow 
or brownish spots, medium-sized or small; uredosori scattered, 
cinnamon-brown, on both sides of the leaves; uredospores 
broadly elliptical, almost globose, 28-33 x 25—29p, light brown; 
epispore thick, with widely separated short spines and numerous 
germ-pores; teleutosori epiphyllous, scattered, naked, black, 
pulverulent ; teleutospores elliptical, rounded at both ends, very 
little constricted, 33-45 28-32", dark chestnut-brown, with 
large tubercles; pedicel up to 75m long, easily breaking off at 
the base, often inserted at the side of the spore. 


On Marsdenia Mexicana, Cuernavaca, September 29, 1899, no. 3529. 


PuCCINIA XANTHII Schw.— On Xanthium, Chapala, September 17, 1899, 
no. 3440; Cuautla, October 12, 1898, no. 3042; Oaxaca, October 17, 1899, 
no. 3644; Cuernavaca, September 28, 1899, no. 3516; on Zinnia tenutfiorg, 
Chapala, September 17, 1899, no. 3441; Oaxaca, October 17, 1899, N0- 
3646. 


PucciniA KUHNIAE Schw.—On Barroetea sabuligera, Aguas Calientes, 
September 12, 1899, no. 3400. 


So eee ee ee le 


Igor] MEXICAN FUNGI 331 


Puccinia Oaxacana Diet. & Holw. n. sp.—Sori hypophyllous, 
small, scattered, brown; uredospores elliptical or ovate, 23~29 x 
23m, with echinulate epispore ; teleutospores oblong or elliptical, 
rounded at both ends or narrowed to the base, little constricted, 
31-50 X 18-25, smooth, pale brown, slightly thickened at apex, 
germinating at once; pedicel the length of the spore or a little 
longer. 

On Baccharis hirtella DC., Oaxaca, October 18, 1899, no. 3673. Aectdtum 
Jragile, n. sp., occurs with this, but it is doubtful whether it is a stage of the 
Puccinia. 


Puccinia Baccharidis-multiflorae Diet. & Holw., n. sp.—Sori 
hypophyllous, small, scattered; uredosori light chestnut-brown ; 
uredospores obovate, 32-42 X 22-28, yellowish-brown, echinu- 
late, apex with a cucullate thickening (up to 7#) and with 3 
germ-pores, which are covered with a convex hyaline thickening ; 
teleutosori dark brown; teleutospores oblong, apex rounded or 
conical, mostly narrowed to the base, somewhat constricted, 
38-60 X 21-30p; epispore smooth, yellowish-brown, apex strongly 
thickened, and mostly somewhat paler; pedicel short, or up to 
60 long, hyaline, rather fragile. The spores germinate soon 
after maturity. 


On Baccaris multiflora, Amecameca, October 31, 1899, no. 3757: 


CAEOMA PUNCTATO-STRIATUM Diet. & Neg.— On Baccharis, Guadalajara, 

September 16, 1899, no. 3435, with Puccinia Baccharidis, Diet olw. 
Puccinta BaccHaripis Diet & Holw.—On Baccharis, Patzcuaro, 

October 17, 1898, no. 3003; Guadalajara, September 16, 1899, no. 3435- 


Puccinia Baccharidis-hirtellae Diet. & Holw., n. sp.—Sori 
hypophyllous, scattered, punctiform, pulvinate, chestnut-brown ; 
uredospores (mixed with the teleutospores) elliptical to globose, 
or obovate, 22-27 x 19-25, with a thin light brown echinulate 
€pispore ; teleutospores elliptical, rounded at both ends, slightly 
constricted, 32-46 X 20-30n; epispore yellowish-brown, slightly 
Or not at all thickened at apex, finely punctate; pedicel long, 
thin, hyaline. 

On Baccharis hirtella, Amecameca, October 31, 1899, n0. 3756- 


332 BOTANICAL GAZETTE [MAY 


Puccinia subglobosa Diet. & Holw., n. sp.—Sori on both 
sides of the leaves, particularly on the under side, scattered, 
punctiform, naked; uredosori chestnut-brown ; teleutosori black, 
pulverulent ; uredospores elliptical or obovate, sometimes almost 
- globose, 18-25 X 17-20 brown, with short spines; teleutospores 
broadly ellipsoidal, often almost globose, rounded at both ends, 
not at all or only slightly constricted, apical thickening slight, 
smooth, dark chestnut-brown, 29-38 X 24-29; pedicel hyaline, 
easily breaking from the host plant. 

On Viguiera Palmeri, Chapala, September 23, 1899, no. 3488. 


Puccinia praemorsa Diet. & Holw., n. sp.—Sori on the under 
side of brown roundish spots, which are somewhat depressed, 
solitary, medium-sized, dark brown, pulvinate, mostly made up 
of several small sori which have crowded together; teleutospores 
oblong-clavate, apex truncate, conical, or very irregular, narrowed 
at the base, somewhat constricted, 40-73 long, upper cell 15— 
30m wide, smooth, brown, thickened at apex ; pedicel short, 
firm, brownish. The spores germinate, at least in part, as soon 
as mature. 

On Brickellia veronicaefolia, Oaxaca, October 20, 1899, no. 3686. This 
is like Puccinia Asteris, but differs in several particulars. 


Puccinia inanipes Diet. & Holw., n. sp.—Sori on both sides 
of the leaf, particularly on the upper, scattered, punctiform ; 
uredosori brown; uredospores elliptical, rounded at both ends 
and when dry both ends depressed, scarcely constricted, apex 
with a very slight cucullate thickening, smooth, dark chestnut- 
brown, 34-42 X 28-31 #, with long hyaline hollow pedicels which 
easily break from the host plant. 

On Exupatorium brevipes, Oaxaca, October 18, 1899, no. 3677- 


Puccinia espinosarum Diet. & Holw., n. sp.—Sori on both 
sides of the leaf, scattered, small or medium sized, naked; 
uredosori brown; uredospores ovate, epispore light brown, with 
short spines, 30-36 X 20-234; teleutosori black; teleutospores 
broadly elliptical, rounded at both ends, apex with a slight cucullate 


1901] MEXICAN FUNGI 333 


thickening, smooth, dark chestnut-brown, 40-50X 32-364; 
pedicel long (up to 125), tou thick, hollow at the base SHY 
easily breaking from the leaf. 


On Eupatorium espinosarum, Oaxaca, October 17, 1899, no. 3651. 


PUCCINIA VIGUIERAE Peck.—On IV. ficta, near Tula, September 20, 
1898, no. 3136; on V. excelsa, Rio Hondo cafion, near City of Mexico, Septem- 
ber 22, 1898, no. 3160; City of Mexico, October 9, 1898 ; no. 3039; on Gymno- 
lomia Ghiesbreghti?, Oaxaca, October 21, 1899, no. 3700; on Viguiera, Toluca, 
September 20, 1898, no. 3178; on Gymnolomia subflexuosa, Oaxaca, October 
17, 1899, no. 3645; on Verbesina trilobata, Oaxaca, October 24, 1899, no. 
3731; on Verbesina virgata, Rio Hondo cafion, near City of Mexico, October 
30, 1899, no. 3751; on Verbesina montanifolia, Patzcuaro, October 10, 1899, 

0. 3604; on Viguiera, Chapala, September 20, 1899, no. 3469; on Calea 
Zacetechichi var. rugosa, Cuernavaca, September 30, 1899, no. 3534; Sep- 
tember 28, 1899, no. 3512; on Cadea hypoleuca, Oaxaca, October 17, 1899, 
no. 3648. 


Puccinia ferox Diet. & Holw., n. sp.—Sori pulverulent, large, 
irregular, brown, attacking the upper leaves and occasionally 
the flowers, often destroying the plants, the leaves being much 
thickened and deformed and often entirely covered with the 
spores on both sides; teleutospores rounded at both ends, 
Strongly constricted, with epispore uniform in thickness, pale 
brown, finely verrucose, 33-43 X 20-26; pedicel short, fragile. 
The spores germinate at once. 


On Verbesina aera Oaxaca, October 21, 1899, no. 3704. A very 
destructive specie 


Puccinia Electrae Diet. & Holw., n. sp.—Sori on various 
colored spots on the upper side of the leaf, less numerous on 
the lower side, small, scattered; uredosori dark brown; uredo- 
Spores obovate or elliptical, 28-35 x 21-25 4, echinulate, brown; 
teleutosori naked, black; teleutospores elliptical, rounded at 
both ends, scarcely constricted, verrucose, apex not thickened, 
36-48 X 24-31; pedicel long, hyaline, or next the spore 
somewhat tinted, easily separating at the base from the host 
plant. 

On Electra Galeottii?, Oaxaca, October 18, 1899, no. 3664. 


¢ 


334 BOTANICAL GAZETTE | MAY 


Puccinia Desmanthodii Diet. & Holw., n. sp.—Sori on the 
under side of large irregularly limited violet or yellow spots, 
small, but closely crowded into large groups in a kind of stroma 
formed of dark brown firm united paraphyses; teleutospores 
oblong or mostly fusiform, narrowed to both ends or truncate at 
apex, not at all or only slightly constricted, 40-60 X 10-174, 
smooth, thickening at apex conical or cucullate, light brown; 
pedicel short or up to 40» long, firm, tinted. 

On Desmanthodium ovatum, Oaxaca, October 18, 1899, no. 3665. Aecidia 
occur on some of the leaves but are too old to describe. 


Puccinia Iostephanes Diet. & Holw., n. sp.—Sori mostly 
epiphyllous on small purple spots, less numerous on the under 
side of the leaf, scattered, black; teleutospores elliptical, 
rounded at both ends, scarcely constricted, chestnut-brown, ver- 
rucose, a light-colored cucullate thickening at the apex and on 
the side of the lower cell, 37-50 x 25-35 »; pedicel long, hyaline, 
easily breaking at the base from the host plant, often laterally 
inserted. 

On lostephane heterophylla? Cuernavaca, September 30, 1899, no. 35435 
on Viguiera dentata, Oaxaca, October 21, 1899, no. 3543, and October 25, 
1899, no. 3744. Type is on no. 3543, the host being an Iostephane which is 


br eoees Ll. heterophylla, although it differs somewhat from other Mexican 
specimens 


Puccinia Guardiolae Diet. & Holw., n. sp.—Sori on purple 
spots, hypophyllous, rarely single sori epiphyllous, punctiform, 
scattered; uredosori dark brown; uredospores nearly globose, 
elliptical or obovate, 24-32 22-25, dark brown, echinulate, 
with two germ-pores; teleutosori blackish, strongly convex, 
firm, small; teleutospores clavate, rounded at apex, or more 
rarely truncate, with a hyaline cucullate thickening, narrowed to 
the base or rounded, constricted, 45-60 X 18-25 , smooth, pale 
brown, germinating at once, whitening the sori; pedicel hyaline, 
firm, mostly shorter than the spore. 


On Guardiola Mexicana, Cuernavaca, September 28, 1899, no. 3513- 


a 


1901] MEXICAN FUNGI 335 


Puccinia conjuncta Diet. & Holw., n. sp.—Sori epiphyllous, 
a few scattered ones epiphyllous, of medium size, except on the 
petioles and stems, where they break through the epidermis in 
large patches, pulverulent, dark brown; teleutospores strongly 
constricted, of two almost globose cells, 34—45 X 22—28y, brown, 
echinulate, apex with a slight cucullate thickening; pedicel 
short, deciduous. 

On Lippia Pringlei, Oaxaca, October 23, 1899, no. 3719B. Collected at 
about 10,000 feet, where there were no specimens of Uredo Lippiae D. & H. 
to be found. The latter occurred at about 6000 feet. 


Puccinia Coulterophyti Diet. & Holw., n. sp.—Spots yellow 
or wanting; sori hypophyllous, small, scattered; uredosori light 
ochre color; uredospores obovate, 29-36 X 22-29P; epispore 
hyaline, apex very strongly thickened, covered with distant, 
globose warts; teleutosori blackish-brown, naked, pulverulent ; 
teleutospores long elliptical or irregular, rounded at both ends, 
scarcely constricted, 33-50 X 23-30; epispore chestnut-brown, 
verrucose, sometimes irregularly rugose; pedicel deciduous. 
One-celled teleutospores are not uncommon. 

On Coulterophytum laxum Rob., Chapala, September 19, no. 3463; 
Uruapam, October 11, no. 3621. These hosts were determined for me at the 
Gray Herbarium, The leaflets differ, however, the former being smooth, 
while the latter are densely white-tomentose. 


Uredo Lippiae Diet. & Holw., n. sp.—Sori hypophyllous, 
scattered, dark brown, pulverulent; uredospores globose or 
broadly elliptical, 25-32 x 25-28; epispore dark brown, closely 
echinulate, to 3u thick, with two germ pores. Hyaline ampulla- 
ceous paraphyses are found in the sori. 

On Lifpia Pringlei, Oaxaca, October 23, 1899, no. 3719. Collected at 
about 6000 feet. No trace of Puccinia conjuncta D. &. H. could be found at 
this elevation. 


UREDO Arbut! Diet. & Holw.— On Arbutus diversifiora, Oaxaca, Octo- 
ber 18, 1899, no. 3662. 


RAVENELIA EPIPHYLLA Schw.—On Brongniartia, Guadalajara, September 


336 BOTANICAL GAZETTE [MAY 


14, 1899 no. 3415; on Tephrosia Talpa, Oaxaca, October Ig, 1899, no. 
3679. 

RAVENELIA INDICA Berk.—On Cassia Aédsus, Tequila, September 29, 
1893, C. G. Pringle. 

RAVENELIA BRONGNIARTIAE Diet. & Holw.—On B. sericea, Oaxaca, 
October 18, 1899, no. 3663; on Brongniartia, Cuernavaca, September 28, 
1898, nos. 3170 and 3022; September 29, 1899, no. 3519; on B. intermedia, 
Tizapan, Valley of Mexico, September 27, 1899, no. 35044. 

RAVENELIA INDIGOFERAE Tranzschel.— On /ndigofera Palmer, Oaxaca, 
October 19, 1899, no. 3682, forming swellings on the stems, and apparently 
quite destructive; on /udigofera Cuernavacana, Cuernavaca, September 22, 
1898, no. 3120. 


Ravenelia spinulosa Diet. & Holw., n. sp.—Sori on both sides 
of the leat, breaking forth from beneath the epidermis, of 
irregular shape and size; uredosori ochre-colored; uredospores 
elliptical or almost globose, 18-23 X 16-Igm, light brown, with 
short spines, germ pores numerous, paraphyses sparingly inter- 
mixed ; teleutosori black, heads hemispherical, 75-110 in diam- 
eter, with 7-9 spores in cross-section, chestnut-brown, with 
numerous globose hyaline cysts on the under side; spores II— 
17m broad,-one-celled, each with a long (up to 84) pale brown 
point at the apex. 

On Cassia multiflora, Oaxaca, October 18, 1899, no. 3675. This species 
is much like the African 2. Stuhmanni P. Henn., but has smaller uredospores. 


PUCCINIOSIRA PALLIDULA (Speg.) Lagerh. (P. triumfetta Lagerh.).— On 
malvaceous plant, Jalapa, April 22, 1899, C. G. Pringle. Is probably on 
Triumfetta sp. 

PUCCINIOSIRA BRICKELLIAE Diet. & Holw.—On Brickellia tomentella, 
Amecameca, October 31, 1899, no. 3767. These specimens show that the 
fungus also occurs on the under side of the leaves in various sized groups, 
sometimes annular, or elongated along the veins, and that the spores are not 
smooth, as originally described, but finely verrucose. 


Endophyllum singulare Diet. & Holw., n. sp.—Sori covering 
large areas of the stems and leaves, with pseudosporidia strongly 
developed, conical, up to 2™™ long, irregularly splitting; spores 
ochre-colored in mass, elliptical or oblong, not rarely pointed at 


1901 | MEXICAN FUNGI 337 


apex, 40-55 X 24-30"; epispore pale brown, strongly thickened 
at apex, closely verrucose. 


On ericaceous plant, Jalisco, Marcus E. Jones. 


Stichospora Mentzeliae Diet. & Holw.,n. sp.—Sori hypophyl- 
lous, scattered, small; uredosori orange-yellow when fresh, white 
when dry ; uredospores formed in short chains, 20-28 X 20-24p ; 
epispore verrucose, hyaline; teleutosori waxy, blood-red, 100— 
120M high; teleutospores formed in a few series, one over the 
other, cylindrical clavate or elliptical, 24-45 X 14-20p, at first 
one-celled, shortly before germination divided by vertical septa 
into four cells; epispore of the upper spore of each series with 
a hyaline thickening (up to 8); sporidia elliptical, often nar- 
rowed to one end, 15 x 10-13. 

On Mentzelia hispida, Chapala, September 18, 1899, no. 3452. 


Coleosporium Verbesinae Diet. & Holw., n. sp.— Sori scattered 
or in irregular groups, hypophyllous; uredosori golden-yellow ; 
teleutosori bright red; uredospores elliptical to globose, 26-33 X 
23-26m, with colorless verrucose epispore; teleutospores cylin- 
drical, up to 130M long, 12X18 thick, strongly thickened at 
apex. 

On Verbesina virgata, with Puccinia Viguierae, Rio Hondo cafion, near 
City of Mexico, October 30, 1899, no. 3751 ; on Verbesina, Cuernavaca, Sep- 
tember 30, 1899, no. 3542. 


Coleosporium paraphysatum Diet. & Holw., n. sp.—Sori hypo- 
phyllous, on yellow or chestnut-brown spots, small, scattered ; 
uredospores long-elliptical to clavate, with sharply projecting 
warts, 26-43 X 17-24; teleutospores at first ellipsoidal and one- 
celled, at length cylindrical or clavate, and divided by horizon- 
tal septa into four cells, 45-65 X 17-22; in many teleutosori 
there were numerous filiform paraphyses. 

On Liabum discolor, Chapala, September 23, 1899, no. 3483. 


Coleosporium anceps Diet. & Howl., n. sp.—Uredosori punc- 
tate, scattered or in circular groups, hypophyllous, white when 


338 BOTANICAL GAZETTE [MAY 


dry ; uredospores elliptical, oblong, or almost globose, 17-25 Xx 
15-204, with long cylindrical tubercles; teleutosori hypophyl- 
lous, on yellow or brown dead spots, single, or often in annular 
or irregular groups, honey-colored, waxy; teleutospores cylin- 
drical or clavate, 90H long, 18-25" wide, 4-celled; pedicel 
hollow, 60-70 long; sporidia mostly cylindrical, 24-30 x 12- 
15. 

n Verbesina sphaerocephala, Chapala, September 24, 1899, no. 3492, 
mostly Uredo, and September 25, 1899, no. 3501, only teleutospores. 


DeEcorAH, Iowa. 


a“ 


OVULE ‘ANDEMBRYO OF POTAMOGETON NATANS. 


CONTRIBUTIONS FROM THE HULL BOTANICAL LABORATORY. 


G. M. HOLFERTY. 
(WITH PLATES II AND III AND ONE FIGURE) 


THIs investigation was begun several years ago, and was 
continued as time permitted. Material from a number of stations 
in the vicinity of Chicago was collected, and other collections 
have been added. From this material about seven hundred 
slides of microtome sections have been prepared and studied. 
The work was done under the direction of Professor John M. 
Coulter and Dr. Charles J. Chamberlain. 

— 


MATERIAL AND METHODS. 


The collections were made in June, July, and August. Those 
made in June show some young spikes still within the cone- 
shaped spathe, while others are just issuing from it. Early in July 
the flowers open and pollination is effected, while in August the 
seed comes to maturity. 

The spikes were cut off with a sharp scalpel and dropped at 
once into the 1 per cent. solution of chrom-acetic acid. After 
remaining in the solution from 36 to 48 hours, the material was 
washed in warm water, and then allowed to remain in running 
tap water 20 to 24 hours. After imbedding in paraffin the 
material was cut on a Minot microtome, the sections being from 
6~—13m in thickness. 

Combination stains were used for all work, cyanin and 
my throsin proving most satisfactory, particularly for sporogenous 
.Ussues. The safranin gentian-violet orange G combination gave 
very satisfactory results for chromosomes and all phases of 
mitosis. The length of time sections should be left in each 
‘er 339 


34° BOTANICAL GAZETTE [MAY 


stain became a matter of some experimentation. In general, 
30 to 40 minutes in cyanin and 30 to 40 seconds in erythrosin 
gave the bests results. The use of turpentine between absolute 
alcohol and xylol was found to be of decided advantage in many 
cases. 

FLOWERS AND SPIKES. 

The flower consists of four similar cup-shaped sepals, four 
opposite stamens, and four alternate carpels. In early stages 
the sepals completely overlap the other organs, but later they 
‘spread sufficiently to 
allow the protruding 
styles to expose the 


a 
OS 


branches, bearing an 


> 
sO 
> Se 


ee anther on each branch. 
(\ : 
fect The anthers are biloc- 
Phd Teh 

reg ulate, and hence the 


) 


& 
= ie 
SLED 


—s 


eo 
es 
relat: 


i} 


group of four carpels 
is surrounded by six- 
teen pollen-sacs. The 
flowers are nearly ses- 


g 


Fic. 1.—Illustrations of spikes: a, X 3 size; 3, 


X #36 Xd x2 sile on the vertical ' 


; spike, alternating in 
such a way that the spiral] arrangement is clearly seen (fig. I). 
From these spikes longitudinal and transverse sections were 
cut. It can be seen that the first and last longitudinal sections 
from such a cylindrical spike would give cross-sections of stigmas, 
styles, and ovaries; while those cut from the center would give 
two rows of longitudinal sections. Cross-sections of the spike 
would of course give longitudinal sections of ovaries, styles, and 
stigmas, but in a plane at right angles to the longitudinal sections 
mentioned above. 
THE OVULE. 
The embryo sac is much more deep-seated than in many other 
plants, occurring in the center of the nucellus, and surrounded 


1got } OVULE AND EMBRYO OF POTAMOGETON 341 | 


on all sides by a mass of tissue from five to eight cells in depth. 
This deep-seated position may bring added protection of some 
sort, or it may have no such significance. In the earliest stages 
of the nucellus no differentiation of cells can be distinguished. 
Its growth appears to be due almost entirely to divisions of 
nucellar tissue other than epidermal, since after the formation 
of the epidermis no layers were observed cut from it by peri- 
clinal walls. In fact, the occurrence of periclinal walls in the 
epidermis is exceedingly rare. 

The archesporial cell and its two daughter cells — Longitudinal 
sections of the nucellus, about the time the primordium of the 
inner integument makes its appearance, show a single hypodermal 
cell having archesporial characters. This cell is larger than the 
others and has a larger nucleus, and its protoplasm is usually 
less dense (fig. 7). When this hypodermal cell is barely distin- 
guishable from the surrounding tissue, it divides by a periclinal 
wall. The outer cell is the so-called ‘tapetal cell” (figs. 2, 3), 
from the progeny of which an extensive region of sterile tissue 
is subsequently formed. The inner cell is the primary sporog- 
enous cell or megaspore mother cell, its sporogenous character 
being plainly shown by the rapid changes which soon take place 
in both its nucleus and cytoplasm. 

The sterile tissue—-The subsequent history of the tapetal 
cell is as follows. The increase in size is followed at once by a 
periclinal division (fig. g),and this by an anticlinal wall (fg. 5). 
Doubtless an anticlinal wall follows in each of these cells, mak- 
ing a plate of four cells lying next to the epidermis. Wiegand* 
Says that the anticlinal walls may precede or follow the periclinals, 
usually preceding, but this was not verified in P. zatans. Anticlinal 
walls now follow in the second layer, thus completing a two- 
layered tapetum. In fig. 7 a three-layered tapetum is shown; 
in fig. 13 one that is five-layered. The process continues until 
in some cases eight layers lie between the embryo sac and the 
epidermal layer. As mentioned before, the embryo sac now 
lies in the very heart of the nucellus. 

* WIEGAND Kari M.: The tonnes of the embryo sac in some monocotyle- 
donous plants. Bor. Gaz. 30:25-47. pls. 6-7. 1900. 


342 BOTANICAL GAZETTE [MAY 


The primary sporogenous cell—At the same time the primary 
sporogenous cell is undergoing important changes. It increases 
very much in size, its contents becoming less and less dense, 
until in some cases vacuoles make their appearance (jig. 6). 
The nucleus also increases in size, and is often surrounded 
by an area of hyaline cytoplasm (kinoplasm), while radiating 
strands of granular protoplasm extend from the nucleolus to the 
periphery of nucleus (figs. 2, 3). The chromatin of the nucleus 
now collects in a closely compacted mass, with a dense non- 
granular nucleolus by its side (figs. 4, 5, 6), the cell having 
passed into the synapsis stage so characteristic of mother 
cells. 

That the development of the primary sporogenous cell has 
been simultaneous with the formation of tapetal tissue is shown 
by the fact that in many cases the former cell has been found in 
early synapsis with only a two-layered tapetum (fig. 4). As the 
nucleus of the mother cell passes to the spirem stage, the chro- 
matin threads are usually more on one side than on the other. 
The nucleoli never appeared granular, though often one to several 
very large granules or black grains could be seen. The cyto- 
plasm was never uniform in appearance, at times having a retic- 
ulated appearance (fig. 5), and again a stringy appearance 

Fig. 6). 

It may be stated that if the manipulation of the stains has 
been fortunate, one finds in all phases of synapsis an excellent 
illustration of the value of cyanin and erythrosin as differential 
stains, the chromatin being always a deep red and the nucleolus 
a deep blue. There was in the case of Potamogeton no differ- 
ence in the way in which the stains were taken up by nuclei of 
the megaspore and microspore mother cells, thousands of the 
latter showing the highly erythrophilous chromatin and the 
highly cyanophilous nucleoli seen in the former. 

Development of potential megaspores.—\t was some time before 
any positive evidence of an axial row could be found; but the 
presence of a cap of densely staining protoplasm at the micro- 
pylar end of the sac was a strong indication of its presence. 


| 


1901} OVULE AND EMBRYO OF POTAMOGETON 343 


Wiegand? demonstrated a two-celled row for P. foliosus, but 
found the fate of the two cells somewhat unusual. He says, 
“the first division of the archesporial cell is immediately fol- 
lowed by a second nuclear division in each of the daughter cells, 
but without the formation of a wall between the two nuclei. 
The upper of these two cells soon perishes, while the lower goes 
on through various phases forming at last the complete embryo 
sac.” 

In P. natans some irregularity appears in the number of cells 
of the axial row, but it is always more than two, brought about 
by the division of one or both daughter cells of the primary 
sporogenous cell. The early stages of the division of the sporog- 
enous cell are shown in figs. g-7._ In the stage shown in fig. 8, 
the chromosomes have passed to the poles, the spindle fibers are 
still intact, and the cell plate is clearly marked. A spindle is 
also formed in each daughter cell (fig. 9). It is at this point 
that the irregularity mentioned above appears. That a spindle 
is not always formed in each cell may be inferred from fig. 77, 
where the lower cell shows by its size, age, and nucleus that it is 
the functioning megaspore. The middle cell, its sister cell, is 
being resorbed along with the upper cell. The spindle shown 
in fig. ro indicates that the two resulting potential megaspores 
will lie side by side. The preparations referred to thus far do 
not show any completed four-celled axial row, but they do show 
several three-celled rows, with spindles to indicate four poten- 
tial megaspores. The spindle in the upper cell of the axial row, 
in both cases shown, is transverse rather than longitudinal. 

The germination of the megaspore—The divisions leading to 
the ante-fertilization stage of the gametophyte appeared perfectly 
regular and normal, although Wiegand} states that in P. pauciflorus 
both egg- apparatus and antipodals were somewhat abnormal. 
He says, “although the normal number of cells in each was 
present, they were formed irregularly. The polar nucleus and 

? WIEGAND, Karu M.: doc. cit. 32. 


3 WIEGAND, Karu M.: Notes on the embryology of Potamogeton. Bot. GAz. 
25: 116-117, 1888 


344° BOTANICAL GAZETTE [MAY 


first and second synergids seem to have been cut off successively 
from the mother nucleus of the egg. The synergids disappear 
almost immediately. A similar irregularity was found in the 
antipodals.”” This could not be verified in P. natans, in which 
the process seems quite normal (figs. 73,74, 77). At this time 
there is considerable plasmolysis even when great care is taken 
in the fixing and subsequent processes (jigs. 75, 16, 77). In fig. 
4 traces of the first spindle of the megaspore can still be seen 
connecting the recently formed daughter nuclei, but there is no 
trace of a wall. By the growth of the sac the antipodals are 
left ina small pocket, and are of short duration (jigs. 16, 77); 
but no traces of a wall shutting them off from the rest of the 
sac. at any time could be found. Nor could a wall be found 
cutting off the egg-apparatus from the opposite end of the sac. 
The polar nuclei never meet in the center of the sac, but always 
nearer the antipodal end (figs. 76, 17). The endosperm 
develops as a parietal layer of cytoplasm in which free nuclei 
are imbedded, and no walls were observed in the most advanced 
stages studied (figs. 19, 21d). 


THE EMBRYO. 


The first division of the oospore is transverse, resulting ina 
_ large vesicular suspensor cell and the first cell of the embryo 
proper (fig. 78). The three-celled embryo shown (fig. 79) has 
probably resulted from the division of the embryo cell. The 
single suspensor cell becomes remarkably large and vesicular, and 
its nucleus and nucleolus correspond in size. The first division of 
the terminal cell in the row of three is longitudinal (jig. 20), 
and this is followed by a second longitudinal division at right 
angles to the first (fig. 27). This quadrant stage is also shown 
in transverse section in fig. 27d. Further views of the embryo 
at this period are shown in figs. 2ra—21d, which are consecutive 
transverse sections from such an embryo as that shown in jig. 
21, in which figure the points at which the sections are made 
are lettered a, 6,c,d. It is seen from the sections that the 
young embryo is cylindrical rather than flattened. 


_— 


eka eames > Sg eae 


1901 | OVULE AND EMBRYO OF POTAMOGETON 345 


In two or three cases a very large nucleus was found near 
the antipodal end of the embryo sac, the embryo being in the 
stage just described. Its origin could not be determined, but it 
seems reasonable to consider it a derivative of the primary 
endosperm nucleus, and possibly the lower nucleus of the first 
division, as in Sagittaria+ and Potamogeton paucifiorus.$ 


THE. UNIVERSITY oF CHICAGO. 


EXPLANATION OF PLATES II AND III. 


Fig. 1. Micropylar end of nucellus, showing the archesporial cell. 
Fic. 2. Primary sporogenous and tapetal cells. 
Fig. 3. The same at a later stage; nucleus with radiating strands of pro- 


_ toplasm, and surrounded by a hyaline a 


Fig. 4. Tip of a young nucellus; inner i: duteeeinnont and primordium of 
outer integument; sporogenous cell with nucleus in early synapsis stage; 
chromatin massed, dense (deeply erythophilous); nucleolus non-granular 
(deeply cyanophilous) ; two-layered tapetum. 

Fig. 5. Primary sporogenous cell; nucleus in synapsis; a two-layered 
tapetum with an anticlinal wall recently formed. 

Fic. 6. Same with two-layered tapetum; nucleus in synapsis. 

Fic. 7. Same with three-layered tapetum. 

Fig. 8, Same with spindle and cell-plate; epidermis with double layer 
but taking no part in tapetal structure. ; 

Fic. 9. Same with two spindles; upper one nearly transverse; beginning 
of axial row. 

Fig. . Axial row showing two potential megaspores, and the upper 
daughter ne with a spindle. 

Fig. 11, Functioning megaspore more advanced, but no sign of further 
division in either middle or upper cells. 

FIG. 12. Functioning megaspore resorbing the potential megaspores. 

Fig. 13. Embryo sac after the first division; daughter nuclei widely 
separated, with large vacuole between them; six-layered tapetum 

1G. 14. Embryo sac, after second division; remnants of the spindle 
between the nuclei at antipodal end of sac; eight-layered tapetum not all 
shown. 

Fig. 15. Micropylar end of embryo sac, showing cells of the egg-appa- 
ratus; from section adjacent to fg. 16. 

*SCHAFFNER, JOHN H.: Contribution to the life history of Segi#taria variabilis. 
Bor. Gaz. 23 : 252-273. pls. 20-26. 1897. 

5 WIEGAND, Karu M.: Bor. Gaz. 25: 117. 1898. 


346 BOTANICAL GAZETTE [MAY 


Fic. 16. Antipodal end of embryo sac; antipodals in pocket, and polar 
nuclei meeting in lower end of sac; from section adjacent to fig. 75. 

Fig. 17. Embryo sac after third division, showing the eight-celled stage; 
synergids with vacuoles, egg cell beneath; antipodals in pocket, two with 
nuclei, remnant only of the third one; polar nuclei meeting below the center. 

Fic. 18. A two-celled embryo, showing the large vesicular cell and first 
cell of ico prope 

1G. Ig. A three- nee embryo imbedded in endosperm and suspended 
from the pe? vesicular cell; endosperm of free nuclei embedded in cyto- 
plasm ; each nucleus with two nucleoli. é 

Fic. 20. A four-celled embryo; the end cell divided longitudinally; very 

large vesicular cell with correspondingly large nucleus an nucleolus 

21. An eight-celled embryo; end-cell divided into four cells; very 
large vesicular cell; letters denote points of cross-sections shown in follow- 
ing figures. 

Fic. 21d. Cross-section of end cell through d@ in fig. 27. 

Fic. 21¢. Cross-section through ¢ in fig. 27. 

Fic. 214. Cross-section through 4 in fig. 27. 

FiG. 21a. Cross-section through a in fig. 27. 


PLATE df 


SIMIC err, del. . \ 


HOLFERTY on POTAMOGETON 


PLATE FT 


BOTANICAL GAZETTE, XXXI 


% 
cs 


? 


Sat 4 erty 


N 


HOLFERTY on POTAMOGETO 


ne ee ee 


i nh a SE a 


BRIEFER ARTICLES. 


FERN VARIATION IN GREAT BRITAIN. 


Up to a comparatively recent date, the departures from the normal 
type of ferns which were found growing wild under perfectly natural 
conditions received little or no attention from professional botanists, 
but were simply dubbed “ monstrosities” and left to amateurs to collect, 
cultivate, and observe. Eventually, however, the discovery that these 
“sports” did not always merely involve a superficial change of form, 
but also that this change was in some cases correlated with hitherto 
unknown modifications and abridgments of the normal life cycle, led 
to a greater interest being taken, with the result that theories as to the 
origin of alternation of generations were seriously affected. Before 
these investigations and discoveries, it was assumed that the life cycle 
was necessarily spore, prothallus, fertilized egg, and finally the sporo- 
phyte or fern proper. Apogamy, however, discovered by Professor 
Farlow, eliminated the sexual act, a vascular structure originating in 
the prothallus which resulted in an asexual bud, whence at once arose 
the sporophyte, the life cycle then being spore, prothallus, sporophyte. 
This, though first remarked in a normal Preris cretica, was subsequently 
found by DeBary to occur with seeming constancy in an abnormal 
tasselled form of Lastrea (Z. pseudo-mas cristata) and several other 
species normal and abnormal. The next discovery was that of soral 
apospory by the writer, on a form of Athyrium filix-foemina, which 
shortened the life cycle in another way altogether, by eliminating the 
Spore, inasses of prothalli being produced, as Professor F. O. Bower 
subsequently ascertained, from the stalks of aborted sporangia, on the 
ordinary soral sites... Here the life cycle runs thus, sporophyte, sorus, 
prothallus, fertilized egg,sporophyte. No sooner was this phenomenon 
announced than Mr. G. B. Wollaston reported the still more remarkable 
case of apical apospory ina form of Polystichum angulare (P. ang. 
pulcherrimum), in which the abnormally long sickle-shaped pinnules 

*DRUERY, C. T.: Jour. Linn. Soc. 21: 354. 1884; 22:427-440. 1885. Bower, 
F. O.: Trans. Linn. Soc. 2: 301-326. 1887. 
eet 347 


348 BOTANICAL GAZETTE [ May 


had their terminal and segmental apices dilated into prothalli, which 
when layered ran the normal course, with the exception that resulting 
plants were invariably defective and depauperate. Here the sorus is 
eliminated, and the prothalli are produced altogether independently 
of the usual reproductive sites, by a modification of the tissue of the 
sporophyte. Several quite independent finds of the pudcherrimum type 
existed, and in every case apical apospory was found to be correlated 
with it, and furthermore soral apospory existed on the fertile fronds. 
The modifications of the life cycle, however, were not even yet 
exhausted, for curiously enough a sporeling of the apogamous variety 
of Lastrea above mentioned was found by the writer to bear a well 
developed prothallus at the tip of its first frond, and the second and 
third bore prothalli, even profusely, on their surfaces,a sort of pro- 
thallic rash as it were.* These produced a brood of plantlets, but one 
and all lost this aposporous character and assumed the merely crested 
type of the parent as their later fronds arose. Here as we have apogamy 
and apospory associated, the life cycle dwindles down to sporophyte, 
prothallus, sporophyte, the shortest possible cut except the bulbils on 
the fronds of viviparous ferns, which have no intermediate stage at all. 
Subsequently, the writer found both soral and apical apospory on 
another variety of Athyrium, and apical on a fimbriate form of Scolo- 
pendrium.? Curiously enough, subsequent to the discovery of the 
aposporous Lastrea, a sporeling of the same parentage originated in 
Mr. Cropper’s collection, which was and is profusely prothalliferous 
from all apices, the smallest piece of frond forming a mass of prothalli 
when layered, which creeps about Marchantia-like and yields a perennial 
crop of typical plants. Finally, in this connection, Professor Farlow 
found an aposporous (soral) type of Preris aguilina. Subsequently, 
at Professor F. O. Bower’s suggestion, Mr. W. H. Lang commenced a 
series of investigations in connection with the prothalli produced from 
spores of abnormal varieties, and thereby practically completed the 
series of abnormal modifications of the life cycle, by finding in more 
than one species prothalli bearing developed sporangia and spores,’ 
thus cutting out the sporophyte, and reducing the life cycle to spore, 
prothallus, spore, an absolute minimum. I must refer to Mr. Lang’s 
* DRuERY, C. T.: Jour. Linn. Soc. 29: 480-482. 1892. 
3 DrueEry, C. T.: Jour. Linn. Soc. 30:281-284. 1892. 

Lana, W. H.: Proc. Roy. Soc. 63:56-61. Phil. Trans. Roy. Soc. Ser. B. 190: 

ae 1898 


1901 | BRIEFER ARTICLES 349 


papers for the most interesting details of these and other vagaries, and 
will merely add that on one and the same prothaJlus several different 
modes of reproduction were found, namely, asexual bulbils, some pro- 
ducing fronds and others merely roots; other prothalli budding out 
from the surface; and sporophytes sexually developed; all these on 
prothalli which themselves were thick cylindrical fleshy masses instead 
of the normal flat cordate thallus, while archegonia, antheridia, and 
sporangia were indiscriminately dotted about among the other varied 
growths. 

The above facts are cited merely as a prelude and justification for 
the suggestion that in the study of American species the varieties should 
have due attention and not be labeled “ monstrosities,” and ignored 
as they were in Great Britain for a very long period. The writer has 
repeatedly urged and urges once more that it is quite as much, if not 
more, in the direction of nature’s exceptions that we should seek the 
key to many of her secrets. Certainly in the study of the abnormal 
fern types alluded to, a number of new and unexpected facts have 
cropped up, which largely modify previous ideas regarding fern repro- 
duction, not one of the links in the normal life cycle being absolutely 
essential. Spore, prothallus, egg, sporophyte, have all in turn been 
shown to be supertluous, the fern in one shape or another reproducing 
itself without them. 

As regards the great number of varietal forms which have originated 
in Great Britain, there is little doubt that it is due far more to the con- 
tinued existence of a coterie of persistent variety hunters than to any 
predisposing elements in the environment. The traveling botanist is 
rarely a variety hunter, but a species hunter, hence he does not in a 
habitat full of known species examine them plant by plant, or scan them 
with a peculiarly trained eye, as would the hunter for sports. If he 
have such an eye and a taste for varieties, we have the evidence that 
“ finds” will crop up abroad as well as at home. Witness Mr. George 
Brown, who found Woodwardia radicans cristata, two finely crested 
forms of Zastrea dilatata, and two tasseled forms of Aspidium hemionttis 
in the Azores; while the writer, hunting in all for about two hours 
recently in Fayal (Azores), found a clump of Aspidium hemtonitis beauti- 
fully tasseled throughout, and a fine pendulous polydactylous variety 
of Pteris aguilina which lined the high road for fifty yards. This being 
so I would advocate variety hunting in the United States as a pursuit 
likely to reward its votaries not merely with very beautiful types for 


350 BOTANICAL GAZETTE [MAY 


their collections, but also with material well worthy scientific attention. 
Thanks to some short notes sent to the Fern Bulletin, | have received 
already fronds of a crested Athyrium found by a lady, and plants of 
Denstaedtia punctilobula cristataand Phegopteris hexagonoptera truncata, 
the latter found by Mr. W. R. Maxon on the Potomac, which have 
originated in the United States as wild sports, a proof that such sports 
exist. At the same time, however, I note that much stress is laid upon 
doubtful subvarieties, which a wider knowledge of marked ones would 
probably minimize considerably. In my own fern hunting expeditions 
I invariably come across subvarieties in which the cutting is more or 
less modified ; but with the abundance of really marked types in mind, 
these are simply noted and left. As exemplifying this abundance the 
figures from Mr. G. I. Lowe’s British Ferns, a descriptive list published 
in 1891, may be appropriately quoted. 


No. of vars 
Pteris aquilina - - - a ‘ ei 17 
Adiantum capillus-Veneris - - - - - 34 
Asplenium adiantum nigrum : - - - 16 
trichomanes - - é z é 27 
maximum~— - : : - = E 28 
Athyrium filix-foemina ss : = Z ‘ : 313 
Scolopendrium vulgare - - : . : 450 
Polystichum aculeatum ~— - - : 4 : 34 
angulare” - - $ “ . F 304 
Lastrea filix-mas - - : * - 5 54 
pseudo-mas - - H : - 42 
propinqua s us as 4 - 28 
montana (reeptene) - ‘ - - 77 
dilatata and others - - - = 57 
Polypodium vulgare - _ : x 74 
Other varieties of sundry species - - - . 214 
Total - x Z is bi 1859 


Upon analysis it is found that 1119 of these were wild finds, without 
reckoning additional independent finds of forms too similar to dif- 
ferentiate. This list cannot be regarded as exhaustive, and we may 
safely reckon the distinct varieties at over 2000, and the wild finds at 
r500. As a concrete example of what one man can doin a single 
district, Mr. J. Moly, of Langmoor in Dorset, is credited with no less 
than 600 distinct finds in that and adjoining counties. His near 


1901 | BRIEFER ARTICLES 35% 


neighbor, Dr. Wills at Chard, found several hundred more. ‘The num- 
bers, however, represent the outcome of many years of persistent 
search, which, however, could not have been exhaustive, as it is recorded 
that Mr. Patey, visiting Mr. Moly, found one of the finest of all types 
(P. ang. plumosum Patey) in Mr. Moly’s own hedge, while Dr. Wills 
was indebted toa farm laborer for that unique fern P. ac. pulcherrimum, 
also found in a hedge close to his house, and thought to have a funny 
look about it by the finder. Finally as an inducement to fern hunt- 
ing, we have the incontrovertible fact that many of these wild sports 
are far more beautiful than the normal forms, and as such constitute 
decorative foliage plants of highest merit. 

A word may be added in reference to the soral bulbils, as these 
occurrences should afford good material for the morphological study 
of the sporangium. Professor Bowers’ monograph on apospory and 
allied phenomena, already cited, gives some illustration of this, and the 
writer’s previous paper’ also alluded thereto. Since then, however, 
such soral bulbils have been recorded as occurring on Adiantum capillus- 
Veneris vars. daphnites and imbricatum, and on Polypodium vulgare 
elegantissimum, while most of the superdum section of plumose Athyria 
have inherited the capacity from the original wild Axminster find. In 
all these cases the bulbils are seated on the soral sites, and are usually 
accompanied by sporangia grading from imperfect and aborted ones to 
perfect ones with full complement of perfect spores which germinate 
freely and yield fairly typical plants. In the case of the Polypodium, 
such bulbils occur only on the most highly developed fronds, and on 
pinnules of extremely fine cutting, the terminals of which run out into 
nearly inch long lingual extensions, pointing I think to aposporal 
tendencies. The sori are massive and consist of filamentous processes 
some of which lengthen out into fronds, while others form perfect 
Sporangia of normal golden yellowcolor. Here then do not appear those 
massive cellular growths which are found on the Athyria, but in time 
one bulbil gets the predominance and a little plant of several small 
fronds is developed. Unfortunately, neither my leisure nor my train- 
ing permit me to investigate properly the transitional stages which 
must exist in cases like these, but I should be happy to provide material 
to any one who desires to follow up this line of research.— CHARLES 
T. Druery, rr Shaa-road, Acton, London, W. 


‘Jour. Linn. Soc. 21: 254. 1884. 


352 BOTANICAL GAZETTE | may 
NOTES OF TRAVEL. IV. 


COFFEE GROWING IN BRAZIL AND THE GIANT JEQUITIBA TREES. 


Ir was the writer’s pleasure to accompany Colonel Charles Page 
Bryan, American minister to Brazil, on a visit to Sao Paulo and one of 
its large coffee estates. Santos, the port of Sado Paulo, and therefore 
the greatest coffee port of the world, is connected by an excellent line 
of railway with the latter. Sao Paulo isa rapidly growing town of 
some 65,000 inhabitants and is surrounded by many square miles of 
plantations of Arabian coffee. 

The geological and geographical commission of the state of Sao 
Paulo is situated here at the capitol, and Dr. Orville Derby, a gradu- 
ate of Cornell, is its originator and present chief. Under Dr. Derby’s 
direction a botanical department has been established and an eco- 
nomic botanic garden started. with Mr. Alberto Léfgren at its head, a 
competent and thoroughly enthusiastic Swedish botanist. Mr. Lofgren 
is assisted by a systematist, Mr. Gustavo Edwall, who has charge of 
the rapidly increasing herbarium, and also by a young Belgian gar- 
dener. ; 

At Tremembé, an hour’s ride by steam train from the city, is the 
young botanic garden, now with about four acres in cultivation and 
many more available, a small laboratory, cold frames, and a convenient 
house for orchids and other shade loving plants. In this garden Mr. 
Léfgren is planting native forage plants, fruit trees, and ornamentals, 
with the view of introducing them into Brazilian culture. As in all 
new countries, the field here for such work is quite open, and this 
garden, if properly supported, will be of great importance to the 
country. 

The best varieties of East Indian mangoes, the southern varieties 
of alfalfa, and the best sorts of oranges are all quite unknown in this 
immense region that is so admirably suited to their cultivation, and to 
that of many other economic plants. The pioneer work of plant 
introduction has scarcely been begun in Sao Paulo, and will not make 
much progress until coffee ceases to be what wheat is in our prairie 
states, the one lucrative crop. 

At the invitation of Dr. Eduardo Prado, owner of one of the 
largest coffee estates, as well as owner and editor of one of the promi- 
nent newspapers in Brazil, Minister Bryan and his party spent a day 
on the Berg6n coffee estate, some eight hours’ ride by train from Sao 


Igor | BRIEFER ARTICLES 353 


Paulo. ‘To any one who has seen Ceylonese, Javanese, or even 
Hawaiian coffee plantations, the Brazilian method of culture will be in 
striking contrast, as no shade trees are employed. The sight of 
thousands of acres of any perennial plant is impressive, but a planta- 
tion of 770,000 trees of coffee, loaded with dark red berries, is really 
beautiful as well as impressive. 

The Brazilian coffee soil is more like New Jersey red clay than any 
other American soils I know, but it breaks up into an impalpable 
powder and rises in brick-red clouds about the horses and wagons 
driving through the plantation, and stains every thing a bright red 
that is very difficult to remove. To a man of fastidious tastes this 
choking, sticky dust would be a decided drawback to life on a 
Brazilian coffee estate. 

From the railway station, the party were driven several miles 
through a broad stretch of coffee trees, and in the very heart of the 
plantation found a bit of virgin forest several acres in extent, that had 
been spared the axe, to show how nature clothed that fair land in the 
days when coffee had no market value. Under the monster trees we 
passed, marveling at their size and beauty, until without a word of 
warning we found ourselves in the presence of two giant trees towering 
above the lesser giants as those would tower above our grandest elms 
and oaks. It has never been my fortune to stand beneath such 
majestic forms of plant life as these Jequitiba trees of the Brazilian 
jungle. The largest specimen measured nearly sixty- two feet in cir- 
cumference six feet above the ground, or over twenty feet in diameter. 
Its height was unknown, but certainly exceeded two hundred and fifty 
feet. The trunk was covered with a regular clear gray bark, and was as 
columnar as if taken from a Grecian temple, tapering very gradually 
to the immense crown of spreading branches. Nothing could have 
given a better idea of the magnitude of this crown than one of the 
fallen branches which lay like a large tree trunk on the ground, meas- 
uring at least four feet in diameter and fifty feet in length. From 
beneath, we could see that in this tree top a veritable forest of epi- 
phytes and parasites was growing, which added very much to its 
graceful outlines. 

These Jequitiba trees, I believe, challenge the world for majesty of 
size and form. They are certainly more beautiful than the slender, 
spire-like Eucalyptus of Gypsland, Victoria; their crowns are far more 
picturesque than those of the Kauri gums (Jammary) of New Zealand, 


354 BOTANICAL GAZETTE [MAY 


although their trunks are similar in shape; and the columnar form of 
their trunks is more graceful than the rapidly tapering ones of the 
Sequoia, whose crowns in comparisonsuare as the Norway spruce to the 
cedar of Lebanon in picturesqueness. On the following day a second 
visit to this wonderful grove confirmed both Mr. Lathrop and me in 
the opinion that there are nowhere in the world more beautiful trees 
than these specimens of Couratari legalis (Myrtaceae). I believe there 
are very few such giants in Sao Paulo, and any botanist traveling in 
this state would be repaid by a visit to these trees, which have only 
been easily accessible to visitors for two years past. 

The coffee estate of Dr. Prado is equipped with modern machinery; 
and many signs of ingenuity in the arrangement of the drying floors, 
pulping machines, and utilization of dried parchment for fuel, indicate 
decided progress since the days when the whole berry was dried and 
the seed removed by a species of husker. 

No seed selection is practiced in the planting of the trees, and as 
yet all these coffee estates, like those of the East Indies, are composed 
of unselected seedlings, although it is an admitted fact that there is a 
great variation in the productive powers and other advantageous 
qualities of the different individuals. As yet no fertilizer save the 
refuse pulp is applied to the soil, although the latter is gradually 
becoming exhausted. 

With Brazilian coffee at the price it now holds in the New York 
market there is little money in coffee raising, and the time is near 
when more labor-saving machinery and improved economical methods 
of culture will necessarily be employed in order to make the business 
pay.—Davip G. Faircuitp, Agricultural Explorer of the U. S. 
Department of Agriculture. 


CURRENT LITERATURE. 
BOOK REVIEWS 
The morphology of spermatophytes.* 


TuHIs volume forms the first part of a treatise on seed plants. The repu- 
tation of the senior author as a lucid and interesting writer is well borne out 
in the present instance. The illustrations, which in many cases are obviously 
due to the junior author, are admirably executed when original, and well 
chosen where they are drawn from earlier publications. The typography and 
general make-up of the book are creditable. 

s is stated in the preface, the present volume grew out of a course of 
lectures, accompanied by laboratory work, and it shows throughout a com- 
mand of the most recent literature and a knowledge at first hand of all the 
morphological facts which may be investigated in a well equipped laboratory 
in the temperate region of the United States. It presents original views in 
regard to morphological terminology and phylogeny, while concerning Ginkgo 
and the Coniferales a considerable amount of original information is supplied. 

The extant Gymnosperms are divided in accordance with the results of 
recent investigations into four classes, the Cycadales, Ginkgoales, Coniferales, 
and Gnetales. The information supplied concerning the first class incorpo- 
rates the important recent works of Ikeno, Webber, and Lang on the gamet- 
ophyte. The interesting genus Ginkgo is elevated on the basis of the 
researches of Hirasé, Webber, and Seward into the representative of a group. 
(Ginkgoales) distinct from the Coniferales. We find here a considerable 
amount of original information concerning the earlier and later stages of the 
seed, including several good figures and photomicrographs. More space is 
naturally given to the Coniferales than to any other class of Gymnosperms,. 
since they are the representative group of the present day. In this connec- 
tion we find an éxcellent photograph illustrating the Heteromorphy of certain 
coniferous seedlings. The account of the reproductive organs is mainly based 
on the original examination of Pinus Laricio. The photomicrographs and 
figures in this section are particularly fortunate, especially those saike 
illustrating the development of pollen and of the embryo. autho 
express the opinion that the genus nds is perhaps the most sdecilieed 
representative of the group. Towards the much disputed question of the 

OULTER, JOHN M., and CHAMBERLAIN, oa J.: Morphology of Sperma- 
Spices Part I. Sv nucanaras 8vo. pp.x + 188. figs. 706. New York: D. Apple~ 
ton & Co., 1900. $1.75. 
Foon 355 


356 BOTANICAL GAZETTE [MAY 


morphology of the ovuliferous scale of the Coniferales they occupy a judicial 
attitude, although obviously inclining to the view that in the Pinaceae at any 
rate it is to be regarded as a modified shoot in the axil of the sterile bract. 
Concerning the treatment of the Gnetales little need be said. Although of 
necessity presenting no original features it summarizes the most recent litera- 
ture. 

In its treatment of the fossil Gymosperms the present work is far beyond 
any previous American botanical work. The Cordaitales receive a sufficient 
consideration, illustrated chiefly by Renault’s well-known figures. The 
extinct cycadoid group, the Bennettitales, is fully discussed from the stand- 
point of the existing literature, and an interesting account, in part original, is 
given of the recently discovered microsporophylls of the group, which as yet 
have been found in a state of suitable preservation only in the United States. 
The microsporangia occur in synangial sori on the abaxial surfaces of the 
sporophylls, thus resembling at once Cycas and the marattiaceous ferns. A 
further interesting feature is that the strobili in this group were not unfre- 
quently bisporangiate, bearing both megasporophylls and microsporophylls. 

The work closes with chapters on the comparative morphology, phy- 
logeny, and geographical distribution of the Gymnosperms. As regards 
comparative morphology the authors have almost entirely excluded vegetative 
anatomy from their work, and among so many admirable illustrations of the 
external features and reproductive organs of living and fossil Gymnosperms 
there are few or none representing anatomical structures of the vegetative 
organs. The present volume is destined to have such an important influence 
on the study of Gymnosperms on our continent, that it is impossible not 
to regret that its authors have not done something to direct the attention of 
American botanists to the importance of anatomical studies, so much neg- 
lected as yet in North America. The work of European palaeobotanists has 
apparently finally set at rest the claim that the reproductive organs of the 
higher plants are the only trustworthy guide in matters of morphology and 
phylogeny. It appears to be established as a result of the work of the late 
Professor Williamson and his followers that the fibrovascular skeleton of 
plants is quite as important phylogenetically as the osseous skeleton has 
er to be in the case of animals. The Calamities and Sigillariae, for 
example, were put by Williamson, on account of the constant and character- 
istic features of structure of their vegetative organs, with the Equisetales 
and Lycopodiales respectively, in opposition to Brongniart who placed them 
among the Gymnosperms. The subsequent discovery of their reproductive 
organs by Williamson and Zeiller only confirmed Williamson's predictions. 
At the present time the Cycadofilices, a group on anatomical grounds alone 
considered transitional between the Filicales and Gymnosperms, are in the 
same position as that formerly occupied by the Calamites and Sigillariae. 


1901] CURRENT LITERATURE 357 


The actual condition of our anatomical knowledge seems to justify the gen- 
eral statement that the skeletal features of plants are even more conservative 
than their reproductive organs, and of quite as great importance in estab- 
lishing the phylogenetic grouping of the Vasculares. 

Turning to phylogenetic matters, the authors consider with Potonié and 
Scott that the Cycadofilices form the connecting link between Ferns and 
Gymnosperms. This group they believe to have given rise in all probability 
to two series, the Bennettitales and the Cordaitales. From the former the 
Cycadales were derived at a later date, while from the latter stock branched 
off subsequently the Ginkgoales and Coniferales. The significance of this 
phylogenetic tree would be more apparent had the earlier part of the book 
given a better illustrated account of the Cycadofilices and the Cycadofilicinean 
features of the living and extinct Gymnosperms. 

The present admirable volume is indispensable to every botanist, and 
the reviewer may perhaps express the hope that in the second edition, which 
will doubtless soon be called for, the authors will add to its usefulness by a 
well illustrated summary of the relevant points of vegetative anatomy.— 
E. C. JEFFREY, Zoronto. 

Bergen’s botany. 


WHEN there began a revulsion against the teaching of botany from the 
floral standpoint, Bergen’s Elements of Botany was one of the most helpful 
text books, because it looked toward the introduction of the student to the 
more vital phases of botany. The book met instant and increasing success, 
because it was just different enough from the former texts to attract the 
teacher who felt the need of a change, and not so different as to repel the 
teacher who was willing but not anxious to find a new guide. 

The Foundations of Botany recently issued? is in some sense a revision of 
the older book. It is said to be written upon the same lines, but it differs 
from the Elements in the extension of the portion treating of cryptogams, and 
in the introduction of a section on ecology. The adoption of these features, 
which characterize several of the more recent texts, is an acknowledgment 
that the lines on which they proceed are approved by teachers, and evidence 
that the publishers desire to meet this demand. The //ora, which in the 
earlier book was wholly inadequate, has been increased fivefold, and, judging 
from the title-page, is issued in editions adapted to various sections of the 
country. 

The illustrations are all new and almost without exception admirably 
selected and well made. Many improvements in the text are also evident, 
and the book is probably as accurate as any text-book now on the market 

BERGEN, JosePH Y.: Foundations of botany. 12mo. pp. xii+412. #, 
With which is combined : Bergen’s botany, key and flora; northern and we ao states 
edition, PP- 257. #gs. 25. Boston: Ginn & Co. 1901 


358 BOTANICAL GAZETTE [MAY 


We regret that the author did not use the metric system ; his objection that 
itis not familiar outside of chemical and physical laboratories is really an 
indictment of botanical laboratories. 

But the book still lacks the treatment of cryptogams which alone can 


those of seed plants, and the unfortunate name “spore plants”’ 
the two sharply apart. The chapters on thallophytes, bryophytes, and 
pteridophytes contain directions for enough laboratory work for a year, but 
they are not well organized pedagogically, nor in such fashion as to indicate 
the more important parts. A single chapter on the evolutionary history of 
plants attempts to furnish a thread of philosophy which has been wanting 
earlier, but the student is left almost unaided to string his pearls of fact, great 
and small, into a shapely whole. The great pregnant ideas of alternation of 
generations, heterospory, and the seed habit lose all their significance unless 
they are presented in connection with the facts they coordinate. 

One naturally expects the Foundations of Botany to exemplify, as most 
books do, the subjects and mode of presentation which the author thinks best 
for the schools. But we find it difficult to interpret Mr. Bergen. One looks 
for the present book tu show advance along the lines which characterized the 
Elements. On the contrary it seems to encourage the teacher of ‘“ analysis” 
by bringing to his hand a better flora; it meets the demand for ecology by 
organizing a section on this subject; it gives the histological teacher ample 
scope for the use of microscope and reagents; it does not develop further the 
physiological features, What is the teacher todo? Is the book a guide to 
wise botanical instruction or simply a book catering to all tastes? This 
uncertain pedagogical sound seems to us to make it doubtful whether /owzt- 
dations will prove as satisfactory as Elements, which had in its time a mission 
that it excellently fulfilled C. R. B. 


Two books on mushrooms. 

PoPpuLAR books on mushrooms multiply. The latest two issue from New 
York. There are many good things to be said for Miss Marshall’s account? 
of the more conspicuous Ascomycetes and Basidiomycetes. In the first place, 
the figures and diagrams are excellent and so numerous that eighty-three of the 
hundred and more species described are well illustrated. A number of the 
half-tone plates are printed in colors and the remainder in black and white. 
The descriptions are simple and clear, and there is also much interesting infor- 
mation appended concerning habits and the edible and poisonous properties 

3 MARSHALL, NINA L.: The*mushroom book. A popular guide to the identifica- 
a8 and study of our commoner fungi, with special emphasis on the edible varieties. 
mp. 8vo. pp. xxvi+167. With many illustrations in color and black and white. 
eatin: Page & Co. New York, 1901. $3.00 


1901] CURRENT LITERATURE 359 


of the important species. Miss Marshall has introduced an elaborate key 
at the beginning of the book, designed to lead the reader directly to the 
enus. This naturally presents the complexities inherent in a subject of such 
acknowledged difficulty. One notes some inconsistencies. ‘For con- 
venience,” Craterellus is described among the “fungi with gills,” but even 
then the writer failed to find its place in the key. Such looseness reacts in 
the end on the general worth of the book. Then it seems a pity that Miss 
Marshall should attempt to describe in general language, and sometimes even 
figuratively, structural characters and physiological activities that are funda- 
mental to all careful observations of fungi. Such expressions have a pseudo- 
simplicity, which deceives and misleads instead of enlightening the novice. 
The second work is issued as a Memoir of the New York State Museum, 
by the botanist, Mr. Charles H. Peck. The forty-eighth report for 1894, con- 
taining quarto colored plates and descriptions of edible fungi, was in great 
demand, so that several editions were exhausted almost as soon as issued. 
The forty-ninth, fifty-first, and fifty-second reports, contained illustrations and 
descriptions of thirty-three species. The work of the past year adds four- 
teen species to the list. On account of the great demand for these reports by 
mycologists and mycophagists, the parts on edible fungi are brought together 
to form the present memoir, illustrations and text having been revised when 
necessary. Thus the forty-eighth report with sixty-nine species and the 
present memoir with forty-eight species illustrate to date the edible and 
poisonous fungi of the state. The character of descriptions and plates is 
like that of former reports. The figures are stiff and mechanical in drawing 
and rather crude in coloring. A comparison of the two books in this respect 
shows the great superiority of photographs and half tones over anything but 
the most expert and artistic drawing, and the most expensive reproduction.— 
B. M. Davis. 


MINOR NOTICES. 

THE REPORT of the state botanist of New York for 1899 has just appeared 
in its usual dilatory fashion. It contains descriptions of numerous new fleshy 
fungi, and three colored plates.— J. M. C. 

(E THIRD FASCICLE of Schumann’s Blihende Kakteen (Iconographia 
Cactacearum) has appeared, containing beautiful illustrations of chee - 
cactus longihamatus Gal., E. Monvillei Lem., E. Fordii Orcutt, and £. Knip- 
pelianus Liebn.— J. M. C. 

THE SEVENTH FASCICLE of the first volume of “ Illustrations de la Flore 
du Congo,” by Wildeman and Durand, has just appeared, containing plates 

* PECK, CHARLES H.: Report of the State Botanist on Edible Fungi of New 
York, 1895-1899. 4ta. pp. 133-234. pls. gz-68. Albany: University of the State of 
New York. Igoo, 


360 BOTANICAL GAZETTE [MAY 


73-84. The publication continues its high character, and the plates represent 
the best of the lithographer’s art.—J. M. C 

ONE of the most recent additions to the local floras of Britain is the Flora 
of Cheshire, prepared from the manuscripts of the late Lord de Tabley. It 
gives in great fullness an account of the floral districts, the ecological condi- 
tions, and a list or species accompanied by a detailed account of stations.— 
i, Sas 3 

‘THE FIRST FASCICLE of the third volume of Pittier’s Flora of Costa Rica® 
red. It contains the “ Filices, Equisetaceae, Lycopodiaceae, 
esc aliavead and Rhizocarpaceae,” by H.Christ. Seventeen new species 
and ten new varieties of ferns are described, besides one new species and two 
new varieties of Lycopodium.—]J. 5 

BILTMORE BOTANICAL STUDIES is the title of a new botanical journal, 
to be “issued at irregular intervals,” and to include papers by the director 
of the Biltmore Herbarium and his associates. The first number was issued 
April 8, 1891, and includes the following papers: C. D, BEADLE and F. E. 
Boynton, Revision of the species of Marshallia (7 species, 3 of them new); 
C. L. Boynton and C. D. BEADLE, Notes on certain coneflowers (including 
5 new species of Rudbeckia); T. G. HARBISON, New or little known species 
of Trillium (3 new species); C. D. BEADLE, New species of thorns from the 
southeastern states (21 new species of Crataegus); C. D. BEADLE, a shrubby 
oak of the southern Alleghanies oe eats The journal is very hand- 
somely printed and illustrated.— J. M. C. 

PROFESSOR E. L. GREENE has begun the publication of a series of 
papers bearing the title Plantae Bakerianae. It is to contain lists of plants 
collected by Mr. Carl F. Baker and his colleagues and distributed to various 
herbaria on both sides of the Atlantic. The first volume is to include the 
collections of 1898 by Messrs. Baker, Earle, and Tracy in southern Colorado, 
and the fascicle now at hand extends from Fungi to Iridaceae. A prefatory 
narrative by Mr. Earle describes clearly the region traversed, and gives an 
adequate background for the list of plants which follows. The fungi are by 
Messrs. Earle and Tracy, and the list includes descriptions of thirty-one new 
species. The Polyporaceae have been determined by Professor Underwood, 
the lichens by Professor Fink, and the grasses by Professor Tracy; otherwise 
the determinations are by Professor Greene, who includes descriptions of new 
species of Zygadenus and Allium.—J. M 


TABLEY, Lorp : The flora of Cheshire. Edited by Spencer cogs and with 
a biographical notice of the author by Sir Mountstuart Grant Duff. 8vo. p. cxiv-t 
399. London: Longmans, Green, & Co. 1899. $3.50. 


6 PITTIER, H.— Primitiae Florae Bdahaciceec. Vol. III. pp. 1-69. San José de 
Costa Rica. Ig01. 75 cents. 

7Plantae Bakerianae 1: 1-52. 23JaIgol. descr to Iridaceae. Catholic Uni- 
versity of America, Washington, D.C. Price 40 ¢ 


Igor | CURRENT LITERATURE 361 


NOTES FOR STUDENTS, 

W. A. WHEELER ® has published an ecological account of the vegetation 
of southeastern Minnesota in connection with a list of the species collected 
there. The paper is accompanied by several excellent heliotypes.—H. C. 
COWLES. 

E, M. WILCox has investigated? a rhizomorphic root-rot of fruit trees 
that is doing great damage to the peach, apple, and cherry trees of the south- 
west. The fungus concerned is described as a new species, C/itocybe 
parasitica, and is also found infesting oak trees. A very extensive bibli- 
ography and several plates accompany the report.— H. C. CowLEs. 

CORRENS has shown experimentally” that in Mirabilis Jalapa only one 
pollen grain out of five, and two ovules out of three, are fit, while in Mirabilis 
longifiora one pollen grain out of four, and one ovule out of two are fit. 
‘Therefore the chances of fertilization increase with the number of pollen 
grains used in pollination, and the progeny are stronger.—C. R. B 

IN A RECENT number of Engler’s yearbook ™ there is an account of the 
recent attempts to secure a uniform nomenclature in plant geography. 
Warburg’s address before the Berlin geographical congress in September 
1899 is given, as is also the resolution passed by the Paris botanical congress 
in 1900. Engler makes an appeal for contributions that will tend to clarify 
the various questions, offering to publish them in the yearbook. Particularly 
with reference to the use of the word “formation” is there need of a general 
understanding.—H, C. Cow.es. 

Von SCHRENK has published a preliminary report on diseases of New 
England conifers’ which contains a good deal of interesting material. 
General remarks are made on the conditions in the New England forests, and 
on the relation of fungi to forest problems. Several fungi, mainly Polypori, 
are described with especial reference to the mode of occurrence and the effect 
on the tree. A number of excellent plates accompany the report. Dr. von 
Schrenk’s work has often been noticed in this journal, and must be highly 
commended both for its botanical importance and its practical aspect. —H. C. 
CowLes. 

IN A SHORT paper entitled “A contribution to the natural history of 
marl, C. A, Davis™ shows conclusively that water plants, especially Chara 


® Minn. Bot. Studies, ser. 2, pt. 4. 1900. 

* Bull. 49, Oklahoma Agric. Ex. Sta. 1got. 

* Berichte d. deutsch. bot. Gesells. 18 : 422-435. 1900. 

**Eng. Bot. Jahrb. 29 : Beiblatt 66, 23-30. 1900. 

"* Bull. 25, U. S. Dept. of Agric. Div. of Veg. Phys. and Path. 1900. 
3 Jour. Geol. 8: 485-497. 1900. 


362 BOTANICAL GAZETTE [MAY 


and blue-green algae, are of great importance in marl formation. The cal- 
careous incrustations, which give rise to marl upon the decay of the plants, are 
formed by the deposition of CaCO; when the CO,, which caused it to remain 
in solution, is used in photosynthesis. The oxygen set free in photosynthesis 
also causes the precipitation of CaCO;. One interesting fact brought out by 
the study is that plants vary widely in respect to the incrustations, aes 
selective processes not yet understood.— H. C. COWLES. 


Miss ELizABETH DALE, in a communication to the Royal Society, 
London, through Professor H. Marshall Ward, shows that the abnormal out- 
growths, or intumescences, in A/zbiscus vitifolius Linn. are due to patho- 
logical conditions, being formed in a moist atmosphere, provided there is 
also adequate light and heat. The immediate effect of the damp atmosphere 
is to check transpiration. This, in its turn, by blocking the tissues with 
water, disturbs the normal course of metabolism, and so leads (when the 
light and heat are sufficient) to abnormal development of certain regions. 
The formation of these outgrowths is accompanied by the production of oil, 
which is not found in normal leaves. Its presence suggests that events 
similar to those occurring in succulent plants are taking place, viz., reduced 
respiration and the development of osmotically active substances in excess. 
It is therefore probable that the intumescences are due to the local accumula- 
tion of osmotically active substances, produced under the abnormal con- 
ditions.—H. M. WarpD 


THE LITERATURE of the first mitosis of the spore mother cell of Lilium 
shows a wide divergence of opinion in regard to the phenomena involved, but 
there aré certain stages which have been constantly observed. How these 
stages are derived from one another is the most debated question. Ina 
recent paper Professor Dixon™ figures and describes six well ascertained 
stages, and then proceeds into the debated territory. Nearly all observers 
describe a longitudinal splitting of the entire thread just before the seg- 
mentation into chromosomes, but Professor Dixon believes that the stage S0 
constantly observed arises from the looping on each other and approxima- 
tions of two portions of the thread. Several very suggestive objections are 
urged against the commonly accepted interpretation. Although believing 
that each of the two twisted portions undergoes a longitudinal splitting while 
stillin the spirem stage or immediately after differentiation into chromo- 
somes, regarded as a second longitudinal splitting by Guignard and others, 
the author believes that this is the first and only longitudinal splitting. A 
series of very clear diagrams illustrates the author’s interpretation of the 
composition of the chromosomes and their behavior during the later phases 


4On the first mitosis of the spore mother cells of Lilium. Notes from the Botanical 
School of Trinity College Dublin 4: 129-139. pls. 7-8. 1901 


T901] CURRENT LITERATURE 363 


of mitosis. According to this interpretation there is no qualitative reduction 
during the first division of the spore mother cell.—CHARLES J. CHAMBERLAIN. 

THE LIFE HISTORY of Schizaea Pusilla has been investigated recently, 
and the results form the first fairly complete account of this interesting fern.’ 
The material was collected at Forked river, New Jersey, in July tg00. Sec- 
tions do not seem to have been made except in the study of the root, stem, 
and leaf. While the peculiar gametophyte and the general aspect of the 
young sporophyte are shown more clearly without sections, one cannot help 
feeling that the account of the development of the antheridia and arche- 
gonia, and also of the very young sporophyte, would have been more satis- 
factory if the study had been made from microtome sections. The gametophyte 
is composed of numerous erect branching filaments which have a somewhat 
uniform diameter and bear a striking resemblance to the protonema of a moss. 
The filaments persist until the young sporophyte has attained considerable 
size. The archegonia are not imbedded, but are entirely free, in general 
appearance suggesting the archegonia of certain liverworts. The arche- 
gonium originates as a single superficial cell which gives rise to a row of 
three cells. From the outermost of these is formed a neck consisting of four 
tiers of cells with four cells in each tier. From the middle cell comes the 
central cell which gives rise to the neck canal cell, the ventral canal cell, and 
the egg. The basal cell forms the venter. One figure illustrating the devel- 
opment of the antheridium shows a row of three cells. The outermost cell 
“becomes largé and globular and cuts off a cap cell at the summit, with the 
wall oblique. The large cell divides up into the mother cells of the anthero- 
zoids and one ring cell.” The anatomy of the root, stem, and leaf is described 
in detail. CHARLES J. CHAMBERLAIN. 

Dr. A, Kerr * has recently studied anthrax to ascertain if it is possible 
to produce the spores in a culture of the bacillus grown under artificial ana- 
erobic conditions, and to observe the effect of nitrogen and hydrogen upon the 
growth of the colonies. For the nitrogen experiments Buchner’s tubes con- 
taining pyrogallic acid and caustic potash were used. By this means the 
oxygen and carbon dioxid of the atmosphere in the sealed tubes were 
absorbed, leaving nothing but nitrogen. For the hydrogen experiments he 
employed Kipp’s apparatus for generating the hydrogen, and Botkin’s appa- 
ratus for growing the cultures in plates and in liquid media. Although every 
Precaution was taken against the possibility of error, Klett shows by careful 
experimentation that spore formation in anthrax is independent of the pres- 
ence of oxygen. He was enabled to obtain a rich growth of the bacillus with 

*S BRITTON, ELIZABETH G. and TAYLOR, ALEXANDRIA: The life history of 
Schizaea pusilla. Bull. Torr. Bot. Club 28: 1-19. pls. 7-6. 1901. 

** Die Sporenbildungen des Miltzbrandes bei Anaérobiose. Zeit. f. Hyg. u. Infec- 
tionskrank. 35 : 420. 1900. 


364 BOTANICAL GAZETTE [ MAY 


abundant spores in an atmosphere of nitrogen. In the hydrogen atmosphere, 
on the other hand, there appeared only a very meager growth of the colonies, 
without a trace of spores. From these observations Klett opposes the more 
commonly accepted view that the presence of free oxygen is a necessary 
condition for the formation of spores in the anthrax bacillus, and holds that 
spores cannot be formed in an atmosphere of hydrogen owing to the retard- 
ing effect of this gas upon the growth of the colonies. He further concludes 
that it is not the presence or absence of oxygen that determines the growth 
and the development of spores in the bacillus, but that every gas has its 
specific influence in this respect. € paper is a most interesting and valu- 
able contribution to the controversy of spore formation in anthrax, and it 
also has an important bearing on many experiments where hydrogen, con- 
sidered as an inert gas, has been employed to bring about an anaerobic con- 
dition.—A. A. LAwSson. 

ITEMS OF TAXONOMIC INTEREST are as follows: L. DIELS (Engler’s Bot. 
Jahrb. 29:577—659. 1901) has completed his account of the flora of central 
China, the closing part extending from Bignoniaceae to Compositae. Besides 
numerous new species, Kolkwitzia (Caprifoliaceae) and Hoeckia (Valeri- 
anaceae) are described by Graebner as new genera.—O. E. SCHULZ (idem 
660-735. /s. 6-8) has published a monograph of Je/ilotus. A discussion of 
the history, morphology, teratology, biology, and geographical distribution of 
the genus is followed by a detailed presentation of the twenty-two recognized 
species, three of which are new.—S. SoMMIER and E, LEvIER (Acta Horti 
Petropolitani 16 : 1-586. /s. 7-49. 1900) have published an elaborate account 
of the plants collected in the Caucasus in 1890, illustrated by forty-nine fine 
lithographic plates. The list includes cryptogams as well as seed plants. More 
than a hundred new species are described, and still more numerous new 
varieties.— W. Lipsky (¢dem 18:1-146. 1900), in a contribution to the flora 
of middle Asia, has described numerous new species, and with them two 
new genera of Umbelliferae, Korshinskia and Galagania.— N. L. BRITTON 
(Torreya 1:21. rgor) has described a new Senecio (S. Crawfordiz) from 
Pennsylvania.— M. L. FERNALD (Rhodora 3 : 43-56. 1901) has published a 
synopsis of the northeastern species of Carex of the subsection Vesicariae, 
recognizing eleven somewhat polymorphic species and describing seven new 
varieties.— F. LAMSON-SCRIBNER (Div. Agrost. Circ. 30. 8 Mr 1got) has 
igi Sus new species of lespane he Gc cate, Bouteloua (2), and Danthonia, 
—E. L. Morris (Bull. Torr. 
Bot. Club 28 : 112-122. Db £2. Igor), in his second paper on N, Am. Plantagi- 
naceae, has described six new species of Plantago.—G. N. Best (dem 
123-131. Pls. 17-74), in a revision of the N. Am. Species of Heterocladium, 
recognizes six species and varieties, three of which are described as new.— 
J. Me &. 


= ‘ a sive 
eae ine cinta whos 


OPEN LETTER. 


THE NAMES OF OUR FERNS. 

THE deadly parallel has always been a striking method of impressing a 
point. When its bald statements are unqualified it seems to mean more 
than the facts will warrant, and it is expected to kill at long range. Ina 
recent issue of the BOTANICAL GAZETTE, Mr. Fernald has made use of it to 
show the supposed instability of the “ reform movement,” but does not note 
the fact that ten of the fifteen changes of which he accuses me were made 
in accordance with changed conceptions regarding the limitations of genera, 
and consequently have no bearing whatever on the point he is attempting to 
illustrate. His pee iens in other respects is very happily chosen, and its full 
import will become apparent when we add the part he forgot to use. For the 
benefit of the younger seins of botanists who, like Mr. Fernald, might be 
misled to suppose that all the changes of plant names have been proposed by 
the “reform movement,” I will add from the same group the changes of 
trans-Carline origin that have been made in the past generation as follows: 


FERN NAMES USED IN GRAY’S MANUAL, FERN NAMES USED IN GRAY’S MANUAL, 
FIRS ON. 


T EDITION, SIXTH EDITI 

Polypodium Phegopteris L. (2, 3, 4).* Phegopteris polypodioides Fee. 
Polypodium hexagonopterum Michx. (2, Phegopteris hexagonoptera Fee. 

mi 

olypo odiu um Dryopteris L. (2, 3, 4). Phegopteris Dryopteris Fee. 
okie eris Germanica Wil “(2 3,4,5). Onoclea paneer 28 Hoffm. 
Allosorus chlnciatd henge (2, Pellaea gracilis Hook. 
Pteris at An rpur Pellaea _ietion nie Link. 
Scolopen m ofcing (2, 3,4). Scolopendrium vulgare Smith. 
Staite ie tilobula. Hook. Gs 4,5). | Dicksonia pilosiuscula Willd. 
Dryopteris s Thelypte eris [A. Gray]. Aspidium Thelypteris Swartz 
Dryopteria Noveboracensis [x Gray]. Aspidium Noveboracense Swart 
Dryopteris intermedia [A. Gray]. Aspidium spinulosum peeing Dp. C. 

aton. 

Dryopteris dilatata [A. Gray]. —— spinulosum dilatatum D. C. 
Dryopteris rigida [A. Gray]. Asp um Boottit Tuckerm. 
Dryopteris cristata [A. Gray]. spidium cristatum Swartz. 

ryopteris Goldiana [A. Gray | Asbidi Goldian 
Polystichum aculea m Roth. idium aculeatum Braunii nt 
Polyst —~ acrostichoides Schott. Aspidium acrostichoides Swart 
Polystichum Lonchitis Roth. Aspidium Lonchitis Swartz 

pende: spectabils Willd. (2, 3, 4) Osmunda regalis 


Botrychium lu arioides s Swartz (2.3.4, 5). Sot ae oiee terna atum Sw 
ht ium Vir inginium = bg cig a 5). Botrychium Virginianum Genk: 
ditions in which this same form appeared. 
S passed eben the intermediate stage of Allosorus atropurpureus [A, 
Gray] before reaching its final form. 
tia 365 


366 BOTANICAL GAZETTE — | MAY 


Or in other words, out of the fifty species of ferns from the northeastern 
states described in the first edition of Gray’s Manual, 21, or 42 per cent., bore 
different names in the last issue of* the same work, and more than that, eight of 
these suffered a change in the specific name also, not including those whose 
relative rank was changed from variety to species or vice versa. Surely the 
trans-Carlines will have to admit that this exceeds the 25 per cent. which 
Mr. Fernald cites as such a horrible example. But after all, what matters it 
if 99 per cent. are changed so long as the change is an evolution towards a 
more stable system based on principles less unseaworthy than the personal 
preference hit-or-miss system of Kew and Berlin? 

LuciEN M. UNDERWOOD. 


NEWS. 


Mr. B. E. FERNow, Director of the New York State School of Forestry, 
Cornell University, will give a course of lectures on forestry at The Univer- 
sity of Chicago during the summer quarter. 

Dr. Roscor Pounp, whose studies on the phytogeography of Nebraska 
have made him well known as a botanist, has been appointed a member of 
the new Supreme Court Commission, which is to assist the Supreme Court of 
Nebraska to clear up its docket. The work of the court is far in arrears, and 
the commissioners are de facto justices. Dr, Pound is winning laurels in his 
vocation as well as in his avocation. 

Mr. Epwarp F., BIGELow, the editor of the Nature and Science depart- 
ment of St. Wicholas, has had prepared compressed tablets after Sachs’ for- 
mula for making nutrient solution. Two of the tablets make a pint of the 
solution of usual strength. Two forms are made, those with and without 
ferric chlorid. The tablets are exceedingly convenient. Doubtless they can 
be purchased through Mr. Bigelow. 

THE DEATH of J. G. Agardh on January 17, in his 88th year, removed a 
notable student of marine algae. His first paper on marine algae was pub- 
lished in 1836, and his last paper has just come to the writer’s desk, not all 
the proof of which did the author live toread. His greatest work is the Species 
Genera et Ordines A lgarum, and to this his last paper was a supplement. A 
brief résumé of his work appears in the Journal of Botany for April. His 
great collection, full of types, passes under the immediate contro] of the 
University of Lund 

THE following circular letter has been sent to many botanists in this 
country. As the committee find it difficult to ascertain the addresses of 
American botanists they desire that all botanists who may see this notice send 
their application for membership in the proposed association to DR. ; 
Lotsy, Wageningen, Holland, by postal card, even though they may not 
receive a copy of the circular: 


The undersigned, padiain that a better organization of the botanists of the dif- 
ferent countries ptalie ribute in a most desirable manner to our mutual aim, viz., 
the progress of bo tany, te ave the honor to invite you to become a member of a new 


Society to be called the /néernational Botanical Associati 

eneral meeting will take place at Geneva, doelinciiesid, on the 7th of August 
next in the botanical laboratory of the university at 10 A. M 
Igor ] 367 


368 BOTANICAL GAZETTE [MAY 


seein this meeting several questions will be submitted to the judgment of its 
members, and you are invited to propose rege or in writing such measures as you 
think it desitable that the new society should ad 
The chief object of the association will be ai establishment of a bibliographic 
ied abstracting in a perfectly impartial manner all botanical publications in 
such a pe that the important will be separated from the less 
will not — as some periodicals do—devote page after page to publications of 


questionable value, st most important works are put off with two or three lines or. 
all. 


not mentioned a 
The abstracts ea the desire of the contributors—be published in English, 
_ French, or German. All will be submitted to the judgment of an editor nominated 
by the association and responsible to 

is most desirable that the ene be as wide as possible, since this is the 
only way of making membership inexpensive. Under no circumstances will the 
membership cost more than $6, including the free delivery of the periodical. 

We wish to call your attention to another great advantage of the new society; by 
its means members who live in different parts of the globe will be brought into more 
intimate contact one with another and this will greatly facilitate the procuring of 
material for investigation and demonstration. 

is der certain conditions to be discussed, accepted, or rejected at the general 

ing in Geneva, Switzerland, the proprietors of the Botanisches Centralblatt have 
eae that their journal become the property of the association. 

The call is signed by Bornet, Borzi, Bower, Celakovsky, Chodat, Fair: 
child, Farlow, Goebel, Lotsy, Nawaschin, Raciborski, Rauwenhoff, Schimper, 
Stahl, Warming, and v. Wettstein 


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_~ GOUT, RHEUMATIC GOUT, 

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ie. Laird, Resident Ph sician, Hot Springs, N.C.: : 

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» gpthenions Chica 
183 


1 Tremont Sheest, Boston. 


Vol. XXXI JUNE, 1901 


sah 
z 


JOHN M. COULTER axp EARLS Ri 


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Vol. XXXI, No. 6 Issued June 20, 1903 


CONTENTS 


A CONTRIBUTION TO THE LIFE HISTORY AND CYTOLOGY OF ERYTHRONIUM. 
CONTRIBUTIONS FROM THE BOTANICAL LABORA eroah OHIO a savy a VIII 
(WITH PLATES Iv-Ix). John H. Schaffner 


STUDIES ON CALIFORNIAN PLANTS. I (witH pLaTE x). AH. Hall - . - - =i, 36a 
CONTRIBUTIONS FROM THE ROCKY MOUNTAIN HERBARIUM. II. Aven Nelson - 394 


STUDIES ON THE Se here OF STEMS. II (wirH THREE FIGURES), £dwin 
Bingham Copeland ‘ 2 : : - - - : 


BRIEFER cnigileke: 
i 


SOM Teaver. Vi... David G. Bairchtid oo 3 ae ae 
E ayia PRINCIPLES OF MorpHoLtocy. W. -. Ganong - * 5 zi - 426 

Py aes dae & & et TORE. 
B WS . - - ; : : : . hi ae 


A i a aaa MANU 


THE CYCLOPEDIA OF cera ORE ne 
MINOR. NOTICES : ‘ . = - - ee i, 
NOTES FOR STUDENTS - : - - . : - - - 439 
OPEN LETTERS. 
UsE oF FERN Names. Willard N. Clute - - . - - - z - - 446 
NEWS : ‘s : , , 2 i ‘ fe f Z oe 


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VOLUME XXXI NUMBER 6 


BOLANICAL > GAwer Ee 
JUNE, tr901 ! 


A CONTRIBUTION TO THE LIFE HISTORY AND 
CYTOLOGY OF ERYTHRONIUM. 
CONTRIBUTIONS FROM THE BOTANICAL LABORATORY, OHIO 
STATE UNIVERSITY. VIII. 

Joun H. ScHAFFNER. 

(WITH PLATES IV—IX) 


SEVERAL years ago, while studying at the University of Chi- 
cago, the writer made a special investigation of the reduction 
nucleus in the ovule of Ziiium Philadelphicum, maintaining that a 
true reducing division occurs in this plant (29). Immediately 
after the completion of the investigation, search was made for 
another plant in which to continue the study of reduction, 
together with other points of interest in the life history. The 
type finally selected was Erythronium, and both £. alidum and 
E. Americanum have been studied. The work was carried on for 
some time at the University of Chicago, and has been continued 
for the past three years at the Ohio State University. The 
material was collected mainly near Chicago and in the vicinity 
of Columbus, but some was also obtained from Kansas. It is 
exceedingly difficult to procure the earlier stages of the flower, 
since bulbs with flower buds are very rare when compared with 
the numerous younger sterile ones. 

The usual methods of killing and staining were used; the 
killing fluids being chrom-acetic acid and Flemming’s weaker 
fluid; and the stains anilin-safranin and gentian-violet, Heiden- 
hain’s iron-alum-hematoxylin, Delafield’s haematoxylin, and 


369 


37° BOLTANICAL GAZETTE [ JUNE 


anilin-safranin and picro-nigrosin. The sections were mostly 
cut 10, 12, and 18, and stained on the slide. 


GENERAL CHARACTERS 


The deeply buried bulbs begin to develop the incept of the 
flower early in the summer. By the first of September the 
incipient flower bud is considerably advanced, and the carpels 
are developing the ovules. Usually before the first of October 
the single hypodermal archesporial cell can be distinguished, 
and the integuments are just beginning to make their appear- 
ance. The anther wall shows five layers and the pollen mother 
cells are enlarging. By December first the tetrads are formed, 
but separation of the four microspores may be delayed for some 
time later in certain flowers. The cells of the anther are filled 
with starch grains and the tapetum is still active, some of its 
nuclei being in stages of direct division. At this time there is 
no sign of the division of the nucleus of the microspore, but the 
exine of the wall is developing. The nucleus divides some time 
between December 1 and April 1, but the time was not ascer- 
tained. In the meantime, the archesporial cell in the ovule has. 
been increasing in size and activity, and has formed the continu- 
ous spirem from the chromatin network. In this condition it 
passes the winter. The cell in which the reduction takes place, 
therefore, has a period of development extending over SiX 
months. In some years it cannot be much less than eight 
months. It will also be observed that while the reduction divis- 
ion in the anther takes place in the fall, in the ovule it is delayed 
until early spring. 

The flowers are growing rapidly long before the frost is 
entirely out of the ground, and during this time the divisions in 
_the embryo sac occur, so that when the flowers come out of, the 
ground the divisions are usually completed. 

Very few flowers appear to develop ovules of any size, and 
ripe seed is very scarce. In fact I have seen very little during 
the past three years. Propagation is effected largely by means 
_of the multiplication of the bulbs. About the first of June, at 


haber “gr ay 


| 
; 
fi 


er 


Igor] THE LIFE HISTORY OF ERYTHRONIUM 371 


Columbus, most of the leaves have wilted away. The plant, 
therefore, is rarely much more than two months above ground. 

Erythronium is an ideal example of the retreating bulb. The 
retreating stems are axillary buds which are carried downward 
by growth and division of the cells above and beside the apex of 
the bud. zg. 7 is a section of such a young bulb, and the dotted 
region shows where active cell division is going on. The devel- 
opment of these offsets has been described several times recently, 
so that it is unnecessary to refer to the subject further. 

The deep burrowing is probably not only to place the plant 
in deep soil, but also to keep the flower protected in the warm 
earth during winter. The advantage of retreat for nine or ten 
months underground must be decided, and the causes for the 
habit complex. The leaves come out before there is any danger 
of shading from other plants, and before the leaves of the higher 


stratum of trees shut out the light. The plant is thus well 


adapted to forest conditions. 


KARYOKINESIS IN THE BULB. 


The division stages in the bulb were studied in order to trace 
out the development of the spindle. The resting nuclei usually 
have a rather dense chromatin network with numerous nucleoli 
imbedded in cavities (fig. 2). Often the nucleoli take on fan- 
tastic shapes, probably due to budding and division. Some of . 
these are shown in fig.3,a,6,c, etc. Farther up, beyond the 
division region, the nuclei elongate in the cells of the develop- 
ing vascular bundles. After the continuous spirem begins to 
form, two caps of fibers appear on opposite sides of the 
nucleus, which are the incepts of the future spindle. These 
are dome-shaped or cone-shaped in appearance, and often end 
in definite granules around which there is sometimes a system 
of radiations (fig. 5). Similar incipient spindles, but farther 
advanced, are shown in figs. 6, 7,and 8 In fig. 7 the chroma- 
tin granules are plainly visible in the spirem, forming a single 
chain. fig. 9 represents a loose mother skein some time before 
the formation of the mother star. The spindle at this stage is 


. 


372 BOTANICAL GAZETTE [JUNE 


sharply pointed and ends insmall centrosomes. After the daugh- 
ter star stage there are sometimes two bodies at each pole, which 
may represent divided centrosomes (fig.zo). The cells of the 


- bulb are often packed with starch. In some cases the starch is so 


abundant that the spindles are very much crowded by it (figs. 7z, 
z2). It will be seen from an examination of the figures that 
the development of the spindle proceeds as in the roots of Allium 
(30) and Pinus (12). It is never multipolar. This seems to be 
the normal course of division in vegetative cells, and represents 
the way in which the spindle is developed during the normal 
quantitative karyokinesis in the higher plants. 

Some time after the appearance of the Cytologische Studien, 
Némec, studying in the Bonn laboratory, announced that in 
vegetative cells the spindle is bipolar from the beginning. It is 
doubtful, however, whether the classification he made of bipolar 
and polycentric spindles will be found to hold good in general. 


THE MALE GAMETOPHYTE. 


The nucleus of the microspore probably divides early in the 
spring, for the pollen grain is well developed before the frost 
has left the ground. The tube nucleus takes a very light stain 
and is comparatively small, while the generative nucleus is large 
and is surrounded by dense-staining cytoplasm which is organ- 
ized into a cell amoeboid in form (figs. 76-78). These genera- 
tive cells are very striking in appearance. They can hardly 
have a cellulose wall because of the peculiar shapes they assume. 


The dense coat of cytoplasm is very different from that of Sagit- 


taria and Alisma, where there is very little cytoplasm, and it is 
difficult to see anything but the nucleus. 

Fig. 19 shows a germinated pollen grain on the stigma. The 
tube nucleus and generative cell are still side by side in the body 
of the grain, but are evidently preparing to pass into the tube. 
In the tip of the tube there is some dark-staining material which 
becomes very abundant as the tube elongates, forming dark 
masses or plugs (fig. 20). The tips of tubes in very favorable 
preparations showed no definite nuclei (fig. 27). They must be 


tgor | THE LIFE HISTORY OF ERYTHRONIUM oho 


farther up among the masses of dark-staining material and are 
thus difficult to identify. 

In the liliaceous types the division of the generative nucleus 
takes place in the tube; in many Helobiae and some other 
monocotyledons, in the grain before it is shed. In Typha the 
generative nucleus does not divide in the grain, while in Sil- 
phium °(22), one of the highest types, the division is in the grain 
and the sperm cells are elongated and even coiled like spermato- 
zoids. Such elongated sperm cells are also common in Alisma. 
It appears, therefore, that progressive reduction of the male 
gametophyte has not been uniform in the various lines of 
angiosperms. Search was made for the division of the tube 
nucleus, as is common in certain species of Lilium (8) and 
Hemerocallis (13), but nothing was found. Such a division 
probably never occurs in Erythronium. 

The style has a large continuous: canal, from the stigma to 
the cavities of the ovulary, for the conduction of the pollen tube. 
This canal is lined by a layer of glandular cells for the nourish- 
ment of the tubes (figs. 73, rg). The pollen tube does not grow 
through any tissue until after it passes into the micropyle. It is 
not difficult to see how such an angiosperm could develop from 
a Symnospermous condition. 


THE DEVELOPMENT OF THE MEGASPORANGIUM AND THE 
REDUCTION DIVISION 

As stated before, the archesporial cell begins to enlarge about 
the first of October, and by December first the chromatin net- 
Work is very distinct and is being transformed into the continu- 
ous spirem (jigs. 22-24). Inthe following stages the nucleus 
becomes very large, and the same is true of the chromosomes. 
This makes Erythronium a favorable subject for the study of 
these structures. After December the nucleus probably goes 
into a partial state of rest until early in the spring, at which 
time development and division continue. During this period it 
will be convenient to call the cell a megaspore. The spirem is 
at first very long and slender and the chromatin granules are 


374 BOTANICAL GAZETTE [JUNE 


never so prominent asin Lilium Philadelphicum (figs. 25-25a). In 
L. Philadelphicum the chromatin granules divide and the chroma- 
tin band undergoes longitudinal splitting before much shorten- 
ing and looping take place, but in Erythronium the division of 
the granules seems to be somewhat later, and they do not 
appear double until the band has twisted into the twelve loops 
(jigs. 26, 27, 28,37). The granules are large and more or less 
irregular in shape. A little later the chromosomes appear 
homogeneous throughout. This would certainly give support 
to the belief that the division of the chromatin granules is a 
mechanical contrivance for bringing about the longitudinal 
division of the linin band, although this might not at all interfere 
with their supposed function as bearers of hereditary tendencies. 

The breaking apart of the twelve loops to form the twelve 
individual chromosomes ( fig. 29) appears to be accomplished by 
the twisting and contracting movements of the band. The 
chromosomes are usually of various sizes, some being much 
larger than others. They often appear as single coiled loops 
(figs. 37, 38), but usually their double nature can be readily 
observed. The chromatin loops are not so closely coiled as in 
Lilium Philadelphicum, and when they are arranged in the mother 
star the twisted condition can still be distinguished. The 
chromosomes appear to be attached to the spindle threads near 
the two free ends of the loop, the closed end extending outward 
(see figs. 39-48, chromosomes before division, and figs. 49-57: 
chromosomes after division). During metakinesis the loops are 
uncoiled, and the two free ends are gradually pulled apart until 
each chromosome breaks in the middle, thus accomplishing a 
transverse division, one end of the original chromatin loop 
going to the one pole, and the other to the opposite one. This 
would be a true reducing division. It is exceedingly difficult 
to follow out the course of events at this point, and there is 
always room for doubt as to correct interpretation. There is a 
possibility that the loops are attached at the closed ends. 
However, some of the examples are quite convincing, and 
another than a transverse division seems out of the question (see 


a 


1901 | THE LIFE HISTORY OF ERVTHRONIUM 375 


especially figs. g6 and 47, also compare the undivided chromo- 
some, jig. 44, with a chromosome in the daughter star, fig. 57). 
There is not a single example which will not agree with the sup- 
position of a transverse division, while many of the figures 
could not be explained on the supposition of a longitudinal one. 
And while it may perhaps be granted that a transverse division 
has not been absolutely established, it may be said that there is 
much less evidence in favor of a longitudinal one. In the next 
division the chromosomes are V-shaped and the longitudinal 
splittings perfectly apparent (figs. 69, 70). 

Although there is no way known to the writer of tracing the 
origin of the reduction chromosomes in this nucleus to two 
previous ones, theoretically one might consider it possible that 
the reduction chromosome represents two normal chromosomes, 
and the closed loop the point where the usual transverse break | 
should have taken place. Were this the case, the points of 
attachment of the spindle fibers at or near the two free ends 
would represent the heads of the two simple chromosomes, and 
the break at the head of the loop during metakinesis simply the 
delayed division bringing about the usual number of pieces. 
But such a process would necessarily result in a qualitative 
division. 

The process here described is essentially the same as that 
reported for Lilium Philadelphicum, and the interpretation is 
similar, since it appears to the writer, after a long and careful 
Study of the objects, that no other interpretation seems possible. 

On account of the contradictory character of the investiga- 
tions so far published, it appears that one or the other set of 
observations has been wrongly interpreted, or else there is more 
variation in the phenomenon of chromatin reduction than is 
generally supposed. There may not be so much uniformity in 
the manner in which reduction is brought about as our present 
ideas in regard to the nature of chromatin seem to demand; 
and once the hypothesis is accepted that the chromatin organs 
are not the only bearers of heredity, there is no reason why 
a large amount of variation should not be present. There is 


‘ 


376 BOTANICAL GAZETTE [JUNE 


still room for entirely new hypotheses, and care should be taken 
lest newer and perhaps better suppositions be rejected by the 
too common appeal to authority. 

Shortly before the publication of my paper on Lilium, but 
not until the investigation had been completed, articles on the 
subject of reduction were published by Calkins (6), Mottier (23), 
and Strasburger (34). Each of these authors presented evi- 
dence favorable to the hypothesis that a transverse splitting of 
the chromosomes occurs during the reducing divisions of the 
plants studied. Miss Sargant (27) had also published a paper 
somewhat earlier, in which some facts were presented favorable 
to the supposition of a transverse division. Calkins, however, 
seems to be the only one of these investigators who has not 
reversed his published opinion. More recently Belajeff (2) has 


.asserted the transverse division, while Stevens (32) holds that 


in the ferns studied by him both divisions which go to form the 
spore tetrad are longitudinal. Guignard (14) has lately also 
published articles on the subject, maintaining that there is only a 
longitudinal division. Atkinson (1) has published the results 
of his investigation of sporogenesis in the anthers of Avzsaema 
triphyllum and Trillium grandifiorum. In the case of Arisaema 
he states that a qualitative division takes place in the first divis- 
ion, while in Trillium it occurs in the second. Duggar (11) also 
believes that a transverse division occurs in the first division in 
Symplocarpus fetidus. In studying the development of the 
microspores of Convallaria majalis and Potomogeton foliosus, Wie- 
gand (35) was unable to determine whether the division was 
longitudinal or transverse, but he inclines to the belief that it is 
transverse in the second division. Thus it appears to be very 
doubtful in which division the reduction normally occurs. | Here, 
as in many other problems of cytology, the personal element is 
still very large. 

The zoologists also report these variations. Paulmier (25), 
in his study of the spermatogenesis of Anasa tristis, says that the 
chromosomes have a twisted appearance, and that the first 
division is transverse and a true reduction division, while the 


* 


ashlee in taille nti a 


Igor] THE LIFE HISTORY OF ERYTHRONIUM Sit 


second is an equation division. Some zoologists have found 
that in certain animals the second is the reduction division. 

Some of the nuclei of Erythronium are of enormous size. 
Those in the walls of the ovule are usually from 15 to 20 in 
diameter, while the large reduction nucleus often measures from 
40 to 50h. In many cases, where the sections were cut 18m 
thick, the spindle was distributed through three sections. In 
such cases the spindle threads not only have their terminals cut 
but they are often more or less distorted. The same is true of 
the nuclei before the spindle is formed. If such a large spindle 
were cut into sections 5 thick it would be distributed through 
nine or ten sections ! 

In the sliced spindles of Erythronium multipolar figures are 
very common. Examples are shown in figs. 31, 32@, 33,34) 
and 35. In fig. 3z one pole is intact and ends in a dark body, 
while the other is cut off. Fig. 32a is a multipolar spindle rep- 
resenting a tangential section. The other part of the spidle is 
little injured and shows well-developed centrosomes at the poles 
(fig. 32). In fig. 34, a strand of spindle threads has been dis- 
placed, so that it projects beyond the limits of the cytoplasm. 
In this material no multipolar spindles were found which were 
not sectioned, and they are therefore not regarded as being the 
result either of normal or diseased conditions, but simply due to 
the method of preparation. Indeed, the nuclei and spindles 
were so large that it was difficult to obtain the chromosomes in 
their normal positions, as they were frequently displaced by the 
knife. 

‘Unfortunately, the stages were not at hand for tracing out 
the origin of the spindle. fig. 28 represents the general appear- 
ance of a section of the nucleus some time before the final 
looping takes place. The nucleus usually has an enlarged or 
expanded appearance, with the spirem lying free in the cavity. 
Fig. 27 shows a large number of false poles produced as the 
result of contraction. In fig. 26 the spirem has looped up into 
the twelve loops, but no sign of a spindle appears either on the 
inside or outside of the nuclear membrane. The loops have not 


378 BOTANICAL GAZETTE [JUNE 


broken apart, but were cut by the knife. It is probable that 
the spindle begins to form rapidly at about this stage, although 
it might already have passed its incipient stage and not be 
detected, if it lies closely applied to the nuclear membrane. 

In the study of Lilium Philadelphicum the writer was unable 
to discover the origin of the radiations which appear around the 
daughter nuclei, but subsequent study of Ranunculus demon- 
strated conclusively that they originate around the poles. Fig. 
456 in Dr. Coulter’s article on Ranunculus (10) was furnished by 
the writer as a good example of this. It is from the endosperm 
of R. multifidus, which is a very favorable object for the study of 
such radiations. In the root tips of Allium Cepa the same origin 
was traced step by step. A comparison of figs. 30, 32, 36, 58, 
59, and 60 will show conclusively the origin of the remarkable 
radiations to be seen in well-prepared material of Erythronium. 
The radiations have their origin from the poles, and only later do 
the daughter nuclei push outward and give to the radiations an 
apparent nuclear origin. The radiations at first appear to be very 
straight and regular ( fig. 36), while later they become more or less 
distorted before they begin to disappear (fig. 60). In favorable 
sections centrosomes are visible, as appear in figs. 32, 32, 36, 55: 
and 59. In the stage represented in fig. 76, the attraction sphere 
appears to form a rather indefinite area from which the radiations 
arise. As to whether these bodies are built up temporarily or 
are permanent, the present study gives no information. In either 
case it is proper to call them centrospheres. At least they are 
the centers for the spindle threads and polar radiations. 

The fate of the nucleolus was not discovered. It is still pres- 
ent at the time of the looped mother skein (fg. 26). In later 
stages, at the beginning of the daughter skein, spherical bodies 
were seen in the cytoplasm which may be extruded nucleoli 
(fig. 59). No figures were seen in the entire study which could 
be interpreted as a synapsis stage. The writer has maintained 
that what is usually called synapsis is a mere artifact which can 
be produced at will by using proper reagents. At the beginning 
of the formation of the spirem, however, the chromatin thread 


1901] THE LIFE HISTORY OF ERYTHRONIUM 379 


becomes free and continues to orient itself and contract until 
the looped mother skein is formed. There is a continuous 
shortening and thickening and often twisting up of the entire 
spirem, but the contraction is not one-sided, and it does not 
appear to have any special relation to the nucleolus. 


THE SECOND AND SUBSEQUENT DIVISIONS. 


The division of the reduction nucleus gives rise to the first 
two cells of the gametophyte. The daughter nuclei go into a 
resting stage and form a network from which a new spirem is 
developed (figs. 67-63). The network at first shows granules 
which are visible in a single chain in the spirem (ig. 66), but they 
are not visible after the mother star is formed (figs. 68, 69, 70). 
The chromosomes are distinctly V- and U-shaped, and the daugh- 
ter chromosomes are formed in the ordinary way by longitudinal 
splitting (figs. 69, 70). This is a normal quantitative karyo- 
kinesis, therefore, which is quite similar to the sporophytic quan- 
titative karyokinesis except that there are only half the number 
of chromosomes formed by the transverse breaking of the spirem. 
Several countings indicate about twelve chromosomes in the 
daughter star. In one case the chromosomes were all distinct 
and plainly twelve in number. 

The spindle in fig. 68 has been sectioned, and this may 


account for the lack of poles. The relation of the large vacuoles 


to the position of the poles of the incipient spindle should also 
be noted (figs. 65-67). There are often remarkable radiations 
around the mother nucleus. These have nothing to do directly 
with the formation of the spindle, however, and are the radia- 
tions normally present at this stage in both plant and animal 
karyokinesis. In some cases it appears that they may have 
their origin at the dome-shaped caps of the spindles (figs. 63, 
64). There are also numerous strands of the central spindle 
left between the daughter nuclei of the first division, and it is 
probable that some of the radiations around the daughter skeins 
may also be left and be preserved to the beginning of the fol- 
lowing division. The third division which gives rise to the 


380 BOTANICAL GAZETTE [JUNE 


eight-celled embryo sac appears to be of the same nature as the 
second. Fig. 71 shows the position of these spindles. The 
uppermost nucleus gives rise to the two synergids, the one below 
this to the egg and upper polar nucleus. A typical arrangement 
of these divisions is shown in fig. 72. The old spindle has sur- 
vived in this instance, and has separated into two limbs below. 
As is usual in many of the Liliaceae, the egg apparatus is not 
very definitely organized. A nearly mature sac is represented 
by fig. 73. In this case, however, the nuclei are larger than 
usual. 

Some interesting features were observed in the second divis- 
ion. Insome cases the cytoplasmic radiations around the nucleus. 
at the beginning of the formation of the spirem did not extend 
to the incept of the spindle, but ended in a rather dense cyto- 
plasmic zone surrounding this (fig. 62). This of course may 
not be of any special significance, but merely an individual 
peculiarity. The fate of the central spindle of the first division,. 
however, deserves special mention. This,.as stated, persists 
usually until the following division is well under way. Then it 
often appears to mass up into two very dense irregular bodies 
which stain very deeply (figs. 65, 66, 68). Whether these 
masses represent a special substance distinct from the general 
cytoplasm and that part of the cytoplasm which alone is used in 
the formation of spindle threads and radiations it would be diffi- 
cult, of course, to tell at present. There is little question as to 
the origin of the masses, and if there is a special substance for 
the formation of spindle threads and radiations, distinct from 
the cytoplasm proper, these masses must represent such a sub- 
stance. 

THE DEVELOPMENT OF THE EMBRYO. 

No stages of fertilization were discovered, nor any in which 
the polar nuclei were conjugating. When the pollen tube 
enters the micropyle it increases enormously in size and is 
exceedingly distinct. It is very different in appearance from 
the tubes in Alisma and Sagittaria. The definitive nucleus 
begins to divide about the time of the union of the male 


a 


1901 | THE LIFE HISTORY OF ERYTHRONIUM 381 


and female gametes. /zg. 74 shows an embryo sac with the 
three antipodals, the dividing definitive nucleus, the oospore, 
two synergids, with an extra nucleus which may be a sperm 
cell, and a nucleus in the pollen tube, probably the tube 
nucleus. During the first few divisions of the embryo, the for- 
mation of the endosperm proceeds very rapidly (jig.’76). The 
oospore divides first by a transverse wall ( figs. 75, 76), and then 
each resulting cell divides by a vertical wall, forming a four- 
celled embryo (figs. 77, 78). These divisions are almost simul- 
taneous, although the upper one usually leads. Sometimes, 
however, the divisions are more irregular (fig. 87)» 

The young embryo lies free in the endosperm some distance 
from the upper end of the sac, and the synergids disappear very 
early. After the first few divisions of the embryo the sac 
enlarges greatly below, while the upper part remains narrow 
and may even contract (jigs. 76, 77, 79). Fig. 79 represents a 
five-celled embryo, one cell being cut from the upper tier and 
one from the lower. Fig. 80 is a six-celled embryo in which 
the two lower cells have each divided by a transverse wall, 
while fig. 8r represents a six-celled embryo with two cells of the 
lowest tier cut away. fig. 82 is a twelve- to fifteen-celled 
embryo, and Jig. 83 about a twelve-celled one. These examples 
will show how very irregularly the development proceeds. Up 
to this time and later there is usually a distinct difference 
between the cells which came from the upper and lower cells of 
the first division. This difference is shown not only by a differ- 
ence in the contents of the cell, but especially by the staining 
reaction. Thus in jig. 83 the upper or suspensor cells have a 
bright yellowish cytoplasm, while the embryo cells are very 
granular and deep red. This is also present in Lelium Philadel- 
phicum, sometimes being very prominent in the older embryos 
(9). Figs. 84 and 85 represent later stages of the embryo of 
E. albidum. The suspensor region is rather large and often 
very irregular in shape and much lobed, but the whole structure 
ives rise to only a single embryo, as was verified by numerous 
examples. The embryos were very badly shrunken, however, 


382 BOTANICAL GAZETTE [JUNE 


because imbedded in the horny endosperm which is not easily 
penetrated by killing fluids. In 4. Americanum, as shown by 
Jeffrey, the suspensor is much larger. Figs. 87 and 88 represent 
sections of two embryos from this plant. These show well the 
large umbrella-like suspensor. The lowermost lobe is the one 
which develops the embryo, but if any of the other lobes of the 
suspensor should become separated from the main mass it would 
probably develop an independent embryo. This would be only 
an accidental case, however, as may frequently happen in any 
embryonic tissue. This is probably not to be regarded, there- 
fore, as an ordinary case of polyembryony, but the large su-- 
pensor is especially developed as an embryonic absorbent 
organ, as suggested by Coulter (9). 


COLUMBUS, OHIO. 


LIST OF PAPERS ESPECIALLY CONSIDERED, 


1. ATKINSON, GEORGE F.: Studies on reduction in plants.. Bot. Gaz, 28:1- 


> 


BELAJEFF, W.: Ueber die Reductionstheilung des Pflanzenkernes. (Vor- 
laufige Mittheilung.) Ber. d. deutsch. bot. Gesells. 16:27-24. 18098. 


3. ———: Ueber die Centrosome in den spermatogenen Zellen. Ber. d. 
deutsch. bot. Gesells. 17 : 199-205. 1899. 
4. BLopGetTtT, F. H.: Vegetative reproduction and multiplication in Ery- 


thronium. Bull. Torr. Bot. Club 27 : 305-315. 1900 

. CALDWELL, O. W.: On the life-history of Lemna minor. Bot. Gaz. 
27: 37-66. 1899. 

. CALKINS, G. N.: Chromatin-reduction and tetrad-formation in pterido- 
hytes. Bull. Torr. Bot. Club 24: 101-115. 18 

. CAMPBELL, D. H.: A morphological study of Naias and Zannichellia. 
roc. Cal. Acad. of Sci. Bot. HI. 1: 1-61. 1897 

. CHAMBERLAIN, C, J.: (Contribution to the life-history of Lz/iwm Phila- 
elphicum.) The pollen grain. Bot. Gaz. 23: 423-430. 1897. 

. CouLTeR, J. M.: Contribution to the life-history of Lz/zum Philadel- 


on 


oO 


“I 


oo 


9 
phicum,; the embryo sac and associated structures. Bot. Gaz. 
23: 412-422. re 

0, oe to the life-history of Ranunculus. Bot. Gaz. 25 :73— 


88, 

: ee, B. M.: Studies in the development of the pollen grain in Sym- 
plocarpus foetidus and Peltandra undulata. Bot. Gaz. 29:8! 98. 
1900. 


Lal 
al 


4 
é 


3 ag flatten og 


7 


Igor] THE LIFE HISTORY OF ERYTHRONIUM 383 


12. 


13. ——— 


ol 
> 


mH 
uw 


cI 
Oo 


Lal 
~sI 


al 
co 


Lal 
e 


Nv 
Lal 


Nv 
N 


bv 
w 


w 
- 


FuLLMER, E. L.: Cell division in pine seedlings. Bot. Gaz. 26 : 239-246. 
mage 


: The development of the microsporangia and microspores of 
in Julva. Bot. Gaz. 28: 81-88. 1899. 


. GUIGNARD, L.: Le développement t du pollen et la reduction chroma- 


tique dans le Nadas major. Arch, d’anat. Micr. 2: 455-509. 1899. 


. HAcKER, V.: The reduction of the chromosomes in the sexual cells as 


described by botanists. Ann. Bot. 9: g5-I0!. 1895. 


; ELIRASE, SS: Etudes sur la fécondation et l’embryogénie du Ginkgo 


btloba. Jour. Coll. Sci. Imp. Univ. Tokyo 12: 103-149. 1898. 


. IKENO, S.: Entwickelung der Geschlechtsorgane und den Vorgang der 


Befruchtung bei Cycas revoluta. Jahrb. f. wiss. Bot. 32: 557-602. 
1898. 


. JEFFREY, E. C.: Polyembryony in Evythronium Americanum. Ann. 


Bot. 9: 537-541. 1895. 
KNERR, E. B.: The propagation of Erythroniums, Trans. Kan. Acad. 
of Sci. 15 : 73-75. 1808. 


- Lawson, A. A.: Origin of the cones of the multipolar spindle in Gladio- 


Bot. Gaz. 30: 145-153. 


9 
- Means, M. E.- The range of variation in species of Erythronium. Bot. 


Gaz. 18 : 134-138. 1893. 


. MERRELL, W.D.: A cee ees: to the life-history of Silphium. Bot. 


29 99-133. Igoo. 


Gaz 
: Mowstee: D. M.: Beitrage zur Kenntniss der Kerntheilung in den Pollen- 


matterzellen einiger Dikotylen and Monokotylen. Jahrb. f. wiss. 
Bot. 30: 169-204. 1897 


- NEMEC, B.: Ueber die karyokinetische Kerntheilung in der Wurzelspitze 


von Allium Cepa. Jahrb. f. wiss. Bot. 33 : 313-336. I 


- PAULMIER, F.C.: The spermatogenesis of Amasa tristis. Jour. of Morph. 


15: 223-272. 1899. 


- RIDDLE, Miss L. C.: The embryology of Alyssum. Bot. Gaz, 26: 314- 


324. 18 
. SARGANT, Miss ETHEL: The formation of the sexual nuclei in Lilium 


citeaon cigs I. Oogenesis. Ann. Bot. 10: 445-477. 1896. 
: The formation of the sexual nuclei in Li/um Martagon. il. 
Sec avdiciepdilite: Ann. Bot. 11 : 187-224. 1877 


- SCHAFFNER, JOHN H.:: (Contribution to the life-history of Lz/ium Phila- 


aelphicum.) The division of the macrospore nucleus. Bot. Gaz. 
ci i wild be 1897. 

pe riers in the root tips of 4//ium Cepa. Bet. Gaz, 26:225- 
238. 189 


- SmiTH, R. W.: The structure and development of the sporophylls and 


sporangia of Isoetes. Bot. Gaz. 29: 225-258, 323-346. 1900. 


384 4... BOTANICAL GAZETTE [JUNE 
32. STEVENS, W..C.: Ueber Chromosomentheilung bei der Sporenbildung 
der Farne. Ber. d. deutsch. bot. Gesells. 16: 261-265. 189 
33. STRASBURGER, E. und MorttieEr, D. M.: Ueber den zweiten Theilungs- 

a schritt in Pollenmutterzellen. Ber. d. deutsch. bot. Gesells. 15 : 327- 


332. 1097. 
34. STRASBURGER, E.: Ueber Cytoplasmastructuren, Kern- und Zelltheilung. 
. f. wiss, Bot. 30: 375-405. 1897. 
35: Wacol, K. M.: The development of the microsporangium and micro- 
spores in Convallaria and Potamogeton. Bot. Gaz. 28 :328-359. 1899. 


EXPLANATION OF PLATES IV-IX. 

The figures are reduced to three eighths of their original size. All the 
figures not especially indicated are from Evythronium albidum. The combi- 
nation of objective and ocular is given for each case, the following being 
used: Zeiss compensating oculars, 4, 6, 12, 18; Zeiss 8.o™™ apochromatic 
objective; Leitz 7s oil immersion and 1 objective; Leitz ocular 8; Bausc 
and Lomb +; oil immersion and 3 objectives. The drawings were made with 
the aid of an Abbé camera lucida. 

Fic. 1. Section of young bulb showing region of cell division dotted. L. 


FiG, 2. Normal resting nucleus in growing tip of bulb. B. & L. dy Z. 12. 
Fic. 3. a, 6, ¢, etc. Nucleoli of various shapes from nuclei in growing 


Aas 4. Elongating nucleus in region of developing vascular bundle. B. 
& L. yy Z. 12. 

Fie, 5. Close mother ie with incipient spindle, from young bulb; 
fuchsin iodin- green. B.& b. a2) Fs. 

Fig. 6. Dome-shaped ae from bulb with granular areas at the tips of 
‘the domes; anilin-safranin gentian-violet. B. & L. yy Z. 4 

Fig. 7, Dome-shaped spindle with centrospheres, from bulb; chromatin 
granules distinct; early close mother skein; anilin-safranin gentian-violet. 
Be 1, fe 2. 12. 

Fig. 8. Early close mother skein ise dome-shaped spindle, from bulb; 
anilin-safranin gentian-violet. B. ee eee 

FiG. 9. Beginning of looped mother skein with bipolar spindle and cen- 
narroetg fin bulb; anilin-safranin gentian-violet. B. & L 

IG. 10. Spindle with centrospheres from nucellus; anilin-safranin gen- 

tian-violet. L, py Z. 12 

Fig. 11. Cell with serch: from bulb.- B. & L. yy Z. 4.. 

Fig. 12. Dividing cell, packed with starch, hots bulb. B. & L. +g Z. 12. 

Fig. 13. Cross section of style showing glandular cells lining the style 
canal. B. & L. # 


nena 


7 


BOTANICAL GAZETTE, XXX1 


\ 


SCHAFFNER on ERYTHRONIU! 


M 


PLATE 1V 


PLATE V 


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SCHAFFNER on ERYTHRONIUM 


BOTANICAL GAZETTE, XXXI PLALL VI 


‘eS 36 JHS 35 
36a 
SCHAFFNER on ERYTHRONIUM 


PLATE Vil 


XXX] 


BOTANICAL GAZETTE, 


SCHAFFNER on ERYTHRONIUM 


PLATE Vif 


XXXI 


AZETTIE, 


BOTANICAL G: 


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SCHAFFNER on ERYTHRONIUM 


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BOTANICAL GAZETTE, XXXT 


SCHAFFNER on ERYTHRONIUM 


PLATE IX 


1901 | THE LIFE HISTORY OF ERYTHRONIUM 385 


Fic. 14. Longitudinal section of style showing glandular cells. B. & L. 
#Z. 18 

Fic. 15. Microspore of &. Americanum, December 1. L. +y Z. 4 

Fic. 16. Pollen grain of £. Americanum, April 15; anilin-safranin picro- 
‘nigrosin. L. +g Z 

Fig. 17. Pollen grain; anilin-safranin gentian-violet. L. ~g Z. 4. 

Fig 18. Pollen grain; anilin-safranin gentian-violet. L. +5 Z. 4 

Fig. 19. Pollen grain on stigma with short pollen tube; aniJin-safranin 
gentian-violet. B.& L. 7; Z4 

Fig. 20. Pollen tube in the style canal showing plugs of dark-staining 
material. B. & L. yy L.8 

Fig. 21. Tip of pollen tube. B. & L. py L. 8. 

Fig. 22. Young ovule of &. Americanum with archesporial cell, October 
ee BLT 

Fig. 23. Young ovule of E. Americanum with archesporial cell, Decem- 
ber 1. Flemming’s triple stain. Z. 8 Z. 18 

Fic. 24. Young ovule of £. Americanum with archesporial cell developed 
directly into a megaspore, December 1; network ees to form the con- 
tinuous spirem; anilin-safranin gentian-violet. L. Wy Z. 12 

Fig. 25. Megaspore nucleus in April; continuous spirem and pale-stain- 
ing nucleolus; anilin-safranin gentian-violet. L. 

Fig. 25 a. Another section of the sa 

FIG. 26. Megaspore nucleus in tebe mother skein stage; anilin-safranin 
gentian-violet. L. yy Z 

* Fig. 27, Megaspore showing effect of contraction ; 

ing; anilin-safranin gentian violet. L. 5 Z. 

Fic. 28. Megaspore nucleus at close of clbee mother skein stage. 


may be due to freez- 
L. +6 


Fie G. 29. Megaspore showing twelve chromosomes after disappearance of 

nuclear membrane, L. + Z. 4. 
1G. 30. Megaspore spindle with peculiar poles; anilin-safranin gentian- 
violet, iron-haematoxylin. L. v5 L. 8. 

Fig. 31. Megaspore spindle with one pole cut away and a centrosome at 
the other; the other pole was in the following section; anilin- safranin gen- 
tian-violet, iron haematoxylin. L. +g Z. 12. 

Fig. 32. Megaspore spindle cut 8 1 thick; distinct centrospheres at the 
poles ; anilin-safranin gentian-violet. 

Fig. 32 a. The following section of the same mone showing multipolar 
spindle because of longitudinal sectioning. 

FIG. 33. Megaspore; multipolar spindle produced by diagonal cut; sev- 
eral of the chromosomes disturbed by the knife and some in following sec- 
tion; anilin-safranin gentian-violet, iron- -haematoxylin. L. yy Z. 6 

Fig. 34. Another multipolar spindle produced by diagonal cut; one set 


386 BOTANICAL GAZETTE [JUNE 


of spindle threads dragged out by the knife; anilin-safranin gentian-violet, 
iron-haematoxylin. L. +5 
Fic. 35. Spindle sectioned near the cath part of one pole remaining ; 
anilin-safranin gentian-violet. L. +g Z. 
6. Daughter star stage ns nace radiations around the 
poles ; eiilin: safranin-gentian-violet. L. 7g Z. 1 
36a. Another section of the same 
F1G. 37. Chromosome from reduction nucleus showing twisted condition 
and late transverse division of chromatin granules; loose mother skein ; 
anilin-safranin gentian-violét. L. ys Z. 12 
Fic. 38. Mature typical chromosome ; chromatin granules have dis- 
appeared ; anilin-safranin gentian-violet. L. jy Z. 12. 
Fics. 39-42. Chromosomes at beginning of metakinesis, showing method 
of untwisting ; anilin-safranin gentian-violet. L. +g Z. 12 
Fic. 43-47. Chromosomes in metakinesis stage showing method of 
untwisting and indicating transverse division ; anilin-safranin gentian violet, 
iron- Sipsoggt age Lis py Zs 18 
G. 48. Single chromosome showing apparent transverse division and 
nig character of the twisted chromosome; anilin-safranin gentian-violet. 


FIGs. bes Half chromosomes after division in early daughter star; the 
pole end is in igor case at the top; anilin-safranin gentian-violet, iron-haema- 
toxylin. L. yy Z. 18. Compare figs. 57 and ¢ 

Fic. 58. Bee star in megaspore ; oe cuaiianpbeves at the poles 
from which extend a series of polar radiations ; anilin-safranin gentian-violet. 
L. +e Z. 12. 

Fic. 59. Daughter skein showing further development of radiations from 
the pole; dark bodies are probably micronucleoli ; anilin-safranin gentian- 
violet. L.+4 L.8 

Fic. 60. Daughter skein woe final arrangement of polar radiations ; 
avilin-safranin gentian-violet. L. +4 

1G. 61. Close daughter skein ‘onus disappearance of radiations and 
beginning of fusion of chromosomes to form the network ; anilin-safranin 
gentian-violet. L. 34 L. 8. 

Fic. 62. Upper nucleus in a two-celled embryo-sac; beginning of divi- 
sion; remains of central spindle still prominent; anilin-safranin gentian- 
violet, iron-haematoxylin. L. +y Z 

Fic. 63. Upper nucleus of two-celled sac, showing radiations probably 
Seas from two poles; anilin-safranin gentian-violet, iron- -haematoxylin. 
L. yy 2. 6: 

Fic. 64. Lower nucleus of the same sac ; the sections are diagon 

Fic. 65. Upper nucleus of two-celled sac; incept of spindle and are 
$ Mons spindle ; anilin-safranin gentian-violet, iron-haematoxylin. L. Ts 


1901] THE LIFE HISTORY OF ERYTHRONIUM 387 


Fic. 66. Beginning of second division, showing incept of dome-shaped 
spindle ; also beginning of the collection of central spindle material; anilin- 
safranin gentian-violet, iron-haematoxylin. L. +5 Z. 6 

Fic. 67. Upper nucleus of two-celled embryo-sac, showing incipient 
spindle; anilin-safranin gentian-violet, iron-haematoxylin. L. +5 Z. 6 

Fic. 68. Sectioa of two-celled embryo-sac showing two peculiar bodies 
probably formed from the remains of the central spindle ; anilin-safranin 
gentian-violet, iron-haematoxylin. L. jy Z. 6 

1G. 69. Chromosome from metakinesis stage of the second division show- 
ing longitudinal splitting ; anilin-safranin gentian-violet. L. yy Z. 12. 

Fig. 70. Upper spindle of second division showing the characteristic 
V-shaped daughter chromosomes; anilin-safranin gentian-violet. L. +g Z. 12. 

Fie. 71. Embryo sac showing the third division ; anilin-safranin gentian- 
violet. L. -Jy Z. 4. 

FIG. 72. Upper two spindles of the third division ; anilin-safranin gentian- 
viet. (Le ZA. 

Fig. 73. Mature but somewhat abnormal embryo sac; Delafield’s haema- 
toxylin-erythrosin. L. + Z. 4. 

FG. 74. Embryo sac with fertilized egg and dividing definitive nucleus; 
anilin-safranin gentian-violet. Z. 8 Z.12 

Fic. 75. Two-celled embryo. Z. 8 Z. 12. 

FIG. 76. Two-celled embryo and dividing endosperm ; pollen tube in the 
micropyle. Z. 8 Z. 4. 

FG. 77, Four-celled embryo; pollen tube above. Z. 8 Z, 12. 

Fic. 78. Four-celled embryo. °Z2. 3 2.12, 

IG. 79. Section of five-celled embryo ; two cells in upper tier and three 


Fig. 80. Six-celled embryo. Z. 8 Z. 12. 
Fig. 81. Six-celled embryo ; two cells in the lowest tier cut away. Z. 8 
i2. 


Fig. 82. Section of twelve to fifteen-celled embryo. Z.8 Z. 12. 

FIG. 83. Section of about twelve-celled embryo, showing marked differ- 
€nce in staining reaction between the upper and lower cells. Z. a Ae © 

ie 84. Embryo considerably advanced, showing the large suspensor. 
Z.8 Z. 12. . 

Fig. 85. Restored embryo showing much lobed suspensor about the same 
age as the preceding embryo. Z, 8 Z. 12. : i 

Fic. 86. Daughter star from endosperm showing sharp pointed spindle 
ending in a centrosome; anilin-safranin gentian-violet, iron-haematoxylin ; 
£. Americanum. UL, yy Z. 12. 

Fic. $7: Section of advanced embryo showing lobed suspensor, Z.3 2,12, 

BIG. 88. Sectian of embryo somewhat older than the preceding, showing 
the massive suspensor; remains of pollen tube in the micropyle. Z. 8 Z. 12. 


< 


STUDIES ON CALIFORNIAN PLANTS. I. 
H. M. HALL. 
(WITH PLATE X) 
SPECIES FROM SOUTHERN CALIFORNIA. 


Frasera neglecta, n. sp.— Plant perennial, 3-6%" high, with 
several slender terete glabrous stems arising from a single per- 
pendicular tap root: leaves all opposite, thick and leathery, not 
at all succulent; the lowermost crowded, 13-18 long, 4.3- 
8.6™™" wide, interspersed with others only 5° or less long, taper- 
ing for nearly their whole length to the base which abruptly 
widens and unites with the opposite leaf to form a short sheath ; 
the upper merely sessile by a broad base, those of the inflores- 
cence gradually reduced to spreading or recurved linear-lanceo- 
late acute bracts: inflorescence an elongated interrupted thrysus, 
the lower cymes on ascending peduncles 5—10™ long, few- 
flowered, upper cymes short-peduncled or sessile and more 
densely flowered ; pedicels varying from 1™ in length to almost 
obsolete: sepals 4, lanceolate, cuspidate-acuminate, 6.5"™ long, 
the margins scarious: petals oblong, acute, 8.6-10.8"™ long, 
greenish-white, the veins purple; petaline gland replaced by a 
tubular nectary, which reaches from the base of the petal nearly 
to the middle where it appears on the inner surface as a circular 
cavity with an indenture on the upper side, the orifice surrounded 
by a raised membrane with finely divided margin forming @ 
fringed border: stamens 4; filaments nearly as long as the 
petals, tapering from the broad base to the tip; anthers oval: 
ovary ovate, attenuate into the slender style which is as long as 
the stamehs. : 

Collected at the head of Swarthout cafion, San Antonio mountains, alti- 
tude 2070", June 1900, 1495 Had/ (type); Holcomb valley, San Bernardino 
mountains, August 1882, Parish; Rock creek, San Gabriel mountains, July 

‘Contributions from the Botanical Laboratories of the University of California. 

388 [JUNE 


Igor] CALIFORNIAN PLANTS 389 


1893, Davidson. All these localities are on the north slope of the San Ber- 
nardino range. The specimens from Holcomb valley are not as thrifty 
as the type; the flowers are somewhat smaller and the leaves (all under 7°) 
are principally from the base, leaving the long internodes exposed. The 
type is in the Herbarium of the University of California. 

This plant has been confused with Frasera nitida Benth., but it differs 
from that more northern species in the more open inflorescence, the more 
slender and acuminate leaves and bracts, and above all by the absence of the 
petaline gland, which in F. nétida is narrowly oblong and open for its whole 
length. The nectary in the proposed species is a pocket-like organ inserted 
within the tissues of the lower part of the petal, the opening surrounded by a 
narrow flap with ciliate margin. The raised tissues over the lower part of 
the nectary are ordinarily brown in color. 

EXPLANATION OF PLATE X.— Fig. z. A plant froma single root; X 4. 
fig. 2,a flower; X 1. Fig. 37, petal showing nectary; 2. Fig. 4, longi- 
tudinal section through petal and nectary; X2. /zg.5,astamen; X 2. 

ASCLEPIAS CALiForNIcA Greene, Erythea 1:92. <Acerates 
tomentosa Torr. Two well-marked forms of this species are 
found in Southern California. In the more common form the 
leaves are broadly ovate and distinctly petioled, while the pedi- 
cels are short. The extreme of this form comes from Lytle 
creek caiion, altitude 1830", and has leaves 5-6.2™ wide by 
7-6™ long, while the pedicels are all under 2 in length. 

The second form has oblong-lanceolate slenderly acuminate 
sessile leaves and elongated pedicels. The extreme of this form 
was collected along the Temecula river and has leaves 5™ wide 
by 15°" long, while the spreading pedicels are 3™ in length. , 

That these forms do not deserve even varietal rank is shown 
by the fact that many intermediate forms exist, and one may 
pass from one extreme to the other by merely ascending a few 
thousand meters on some of the mountain slopes. AtgI4 a 
in Cajon pass the slender-leaved form prevails, while in passing 
up the mountains west of the pass various forms are encountered 
until the extreme of the broad-leaved form is met at 1830 meters. 

Gilia modesta, n. sp.—Plant annual, 15° high or less, divari- 
cately branching from near the base, the branches ascending, 
either glabrous or sparingly pubescent: leaves mostly oppo: 
below, the upper alternate, 1°™ long, palmately 3-7 -divided 


390 BOTANICAL GAZETTE [JUNE 


nearly to the base into rigid linear pungent-tipped lobes, these 
with strong midribs: flowers disposed singly or in pairs in the 
forks where they are short pedicellate, and clustered at the ends 
of the branches where they may be either sessile or short 
pediceled: bracts similar to the leaves but lobes usually only 
three, sparingly beset with loose cobwebby hairs: calyx pubes- 
cent, 8-11™" long, about equaling the bracts, cylindrical with 
acute base, hyaline membranous with green ribs prolonged 
beyond the hyaline portion into pungent-tipped nearly equal 
lobes which are 2™ or so long: corolla white, with the tube and 
throat yellow and with two purple lines at the base of each lobe; 
tube short; throat funnelform, 6-8™™ long, nearly equaled by 
the broadly obovate spreading lobes: stamens equally inserted 
near the base of the corolla tube, unequal in length, included: 
ovules in each cell 2 or 3; seeds oblong, developing spiracles 
only sparingly when wetted. 

Gravelly hillsides of Lytle creek cafion along the trail to San Antonio 
peak, altitude 1830™, June 1, 1900, 1443 Ha// (type); Swarthout cafion, San 
Antonio mountains, altitude 2000", June 3, 1900; near Bear valley, San 
Bernardino mountains, altitude 2000", July 16, 1899. The type is in the 
Herbarium of the University of California. 

This species is most nearly related to Gi/ia demissa Gray, but is easily 
distinguished from that by a number of important characters. G. modesta is 
a much larger plant, with a corolla fully 13™™ long, and nearly as wide when 
expanded, while G. demissa has a corolla only 6.5" long. The proposed 
species has a calyx of 5 narrow green ribs of equal width throughout, between 
which is stretched a hyaline membrane connecting them for three fourths their 
length, while G. demissa has a calyx of 5 lanceolate nearly distinct segments 
each with a scarious margin. The lobes of the leaves and bracts are more 
slender and rigid than in G. demissa, and the ovules are never more than 
three in each cell, as against seven in G. demissa. From G. Parry@ Gray it 
differs in many respects, principally in the absence of scales at the base of the 
corolla lobes. The general appearance suggests relationship with the mem- 
bers of the Linanthus section, but it differs from all those in the pubescence, 
in the funnelform corolla, in the small mumber of ovules, and in other 
characters. 

COLLINSIA CALLOSA Parish, Erythea 7:96, Richardson’s ranch 
near Manzana in Antelope valley, May 1896, 2503 J. Burtt Davy. 
This was probably the first collection of this rare plant, the 


1901 ] CALIFORNIAN PLANTS 391 


original description being drawn from specimens collected in the 
San Antonio mountains, June 1899. 

CHENACTIS XANTIANA Gray. This species is well distributed 
along the mountains surrounding the western part of the Mohave 
desert, often at altitudes of 1800-2000". An exceedingly robust 
form has been collected at Manzana in Antelope valley, altitude 
780", by Mr. J. Burtt Davy (no. 1512, May 1896). The stout 
fistulous stems are 3°" high and terminated by heads 2 high ; 
involucral bracts slenderly acuminate, 2™ long; the inner pappus 
when fully developed longer than the corolla. It is hoped that 
collectors visiting that region may secure more material in order 
that the relation between this and the typical form may be more 
clearly understood. 

CHA:NACTIS HETEROCARPHA CURTA Gray. Near Estrella in 
the Coast ranges of San Luis Obispo county, Z. Jared. In these 
specimens the paleae of the pappus are reduced to mere rounded 
scales, which form a crown about the summit of the akene. 


A TRIP TO THE SOUTHERN HIGH SIERRAS.. 

During the summer of tg00 Mr. H. P. Chandler and the 
writer spent some two months botanizing in the Sierras of Fresno 
county. Our first stop was at Ockenden, a small settlement on: 
the ridge between the Kings and San Joaquin rivers, and well 
within the coniferous forests. Reaching here on June 12 we 
found the season just opening, many of the species still being 
in too young a condition to make good herbarium specimens. 

After spending two weeks in that region we followed the 
trail to Tehipite valley, making side trips to the Dinkey grove 
of big trees and Bald mountain en route. Tehipite valley is a 
deep gorge several miles wide on the Middle fork of the Kings 
tiver. The floor of the valley is 1220™ in altitude, while the 
walls rise to over 2100". The flora was somewhat similar to 
that of the foothill region, and when we reached the valley 
(July 6) many of the annuals had already passed out of flower. 

The Mount Goddard region was also visited, the route lead- 
ing by Black mountain, Blaney meadows, and the South fork of 


392 BOTANICAL GAZETTE [JUNE 


the San Joaquin. On July 24 we camped at the very base of the 
mountain just above timber line. The only shrub seen here was 
Salix glauca villosa, but hardy thick-rooted perennials grew 
abundantly around the lakes, and fifty-seven species were 
gathered above 3350". Thirty-three of these were also found 
below timber line, leaving twenty-four strictly alpine species. 
Ascending the peak the vegetation grew more and more scanty, 
until at 4000™ only Draba Breweri, Erigeron compositus, and 
Polemonium confertum remained, while on the very summit, at 
4130”, the Polemonium was the sole representative of the whole 
series of phanogamous plants. 

The following species collected on this trip seem worthy of 
note, 

ERIGERON Bioomer! Parish, Erythea 6:87. Not rare in 
Tehipite valley, July 7, Hall & Chandler, no. 513. Determined 
by Mr. S. B. Parish. 

ER10GONUM NuDUM Dougl. An alpine form was collected 
near Mount Goddard at 3100". The slender stems are not 
over 2°" high and are nearly always simple and monocephalous, 
July 1900, Hall & Chandler, no. 662. 

AQUILEGIA PUBESCENS Coville, Contrib. U.S. Nat. Herb. 4: 5 
Among rocks on Mount Goddard at 3660™, July 1900, Bee & 
Chandler, no. 671. 

DoODECATHEON JEFFREYI redolens, n. ube 
robust, 4.5-6°" high, the herbage redolent with a strong odor: 
leaves erect, lanceolate or spatulate, 2.5-49" long including the 
slender winged petiole: inflorescence glandular pubescent: 
flowers invariably 5-merous: lower part of stamens and capsule 
included in the corolla tube, which is not closely reflexed. 

Along the lakes at the base of Mt. Goddard, 3400, July 25, 1900, Had/ 
& Chandler, no. 676. Type in the Herbarium of the University of California. 
This variety is easily distinguished in the field by the rank odor and by 
the shape of the corolla. The folding of the latter takes place well up on the 


tube, thus making the yellow band very prominent, while the purple ring at 


the base of the corolla in the species is lacking in the variety. 
ERIGERON SALsuGINosuS Gray. This plant, not rare at middle 


BOTANICAL GAZETTE, XXXI PLATE X. 


« 


Wye. 
VX fh) 
ne Nee 
cu . , <7} ‘S AVE 
: I 
, a 
On 
GY i ba 
SIS WA Ae 
a q f¢zz =i 


Ul 
} Zz 
LS Ve <= > 
SS 


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Wy SNY 


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7 
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Sa M, Y, 7 4 ES 
at \ 


qo 
<a 


HALL on FRASERA NEGLECTA, 2. sp. 


1901] CALIFORNIAN PLANTS 393 


altitudes in the Sierras of Fresno county, exhibits a wide varia- 
tion in the pubescence. Some specimens are covered with a 
close pubescence on the leaves and stems, while other are 
entirely glabrous except on the peduncles and involucres. These 
forms grow side by side and intergradations are plentiful. 

Cnicus TroGanus Congdon, Erythea 7: 186. Blaney meadows 
on the South fork of the San Joaquin at 2320" altitude, July 
1900, Hall & Chandler, no. 721. 

PHALACKOSERIS BOLANDERI coronata, n. var.—Akenes with an 
evident pappus consisting of a crown with a divided margin. 
Otherwise as in the type. 

Pine ridge, in the Sierras of Fresno county, altitude 1650", June 15-25, 
1900, Hall & Chandler, no. 187 (type). Specimens were also examined 
from Mariposa county (Coazgdon, October 1896), Yosemite (Mrs. Brandegee, 
July 1883), and Fresno county (Mrs. Peckinpah, 1890), and in each case 
the akenes were found to bear pappus. In old akenes the pappus is not so 
evilent as in the younger ones, as it is curved inward, due to its inability to 
keep pace with the growing akene. The type is in the Herbarium of the Uni- 
versity of California, 

The preparation of this paper has been greatly facilitated by 
the kindness of Miss Alice Eastwood of the California Academy 
of Sciences, Mr. and Mrs. T. S. Brandegee of San Diego, and 

r. S. B. Parish of San Bernardino, in sending specimens for 
examination; and of Mr.-Merritt L. Fernald, of the Gray Her- 
barium, for notes on the type specimen of Gilia demissa. To Pro- 
fessor W, L. Jepson, of the University of California, I am under 

- Special obligations for his continued interest and valuable sug- 
gestions without which the paper would not have been prepared. 


UNIVERSITY OF CALIFORNIA, 
Berkeley, Cal. 


CONTRIBUTIONS FROM THE ROCKY MOUNTAIN 
HERBARIUM. II. 
AVEN NELSON. 
SOME COLORADO SPECIES. 


AN interesting collection of Colorado plants, some 300 
numbers, was kindly submitted to me for examination by Pro- 
fessor Ramaley, of the University of Colorado. They were 
largely of his own collecting, but some of them were by F. Y. 
Moseley, and yet others by Jennie M. Archibald, of Berwind, 
Colorado. Among them were many of much interest and a few 
novelties, upon which the following notes and characterizations 
are offered. 

ZYGADENUS COLORADOENSIS Rydb. This interesting recently 
described ally of Z. elegans seems to be peculiarly a Colorado 
species, as I have not yet met with it in Wyoming. 

CHEIRANTHUS sp. A peculiarly branched form, with petals 
almost rose-colored, was represented by two numbers. It was 
evidently not far from C. asper Nutt., but fruiting specimens 
may show it to be distinct. 

Crematis Scorri Porter. This rare plant was represented by 
rather young specimens, but even these indicated the distinctness 
of the species as compared with C. Airsutissima Pursh. 

SAXIFRAGA RHOMBOIDEA austrina, n. var.—Size, habitat and 
foliage of the species: stems not rarely several from the short 
rootstock, pubescent with gland-tipped hairs as is also the inflo- 
rescence: inflorescence paniculately cymose, strict: calyx lobes 
obtuse, as long as the adnate base: petals elliptic, white, 3" 
long: carpels ovoid, the tips at length widely divaricate : seeds 
very numerous, brown, oblong, more or less subacute at both ends. 

That this is a good species seems quite possible. The paniculate inflo- 
rescence and conspicuously gland-tipped pubescence is in strong contast to 
the usually spherical terminal cluster (which at most becomes only branched- 
thyrsiform) and the glandless pubescence of S. rhomboidea Greene, Pitt. 

304 : [JUNE 


a I i a 


eupany atk 


1901] ROCKY MOUNTAIN PLANTS 395 


3:343- In the variety proposed the petals are not emarginate, the calyx 
lobes are longer than in the species; the presence of a rather conspicuous 
ciliate-pubescence on the margins of the leaf in the variety adds another 
point of contrast. 

Excellent specimens are in hand from Jennie M. Archibald, Berwind, 
Colorado, 1g00, no. 236. 

Parosela Porteri, n. sp.—Caudex woody, with a deep- 
set woody root: stems numerous from the enlarged crown, 
prostrate-assurgent, quite unequal in length, 10-25 long 
(including the oblong spike), strigose, appressed-pubescent : 
leaves trifoliate, densely appressed strigose-silky, cuneately 
obovate or narrower, mostly obtuse, 8-14™™ long, the terminal 
one longer than the lateral pair, barely petiolulate, generally 
Shorter than the slender petiole; stipules slenderly and stiffly 
setaceous, 5-6™" long: spike dense, 2~5™ long; bracts purple, 
lanceolate-acuminate, equaling or shorter than the calyx, silky- 
villous ; calyx tube campanulate, membranous with ten greenish 
nerves, the triangular base of its lobes abruptly narrowed into 
long slender plumose setae which are twice as long as the tube; 
corolla lemon-yellow; the standard very short, reniform-triangu- 
lar, acute, broader than long, shorter than the slender claw; the 
wings oval, short clawed; keel petals oval or even broader, sur- 
passing the wings, the blade about 6™ long and the claw about 
am; Ovary silky, included in the calyx tube, half as long as the 
slender plumose style which is much exceeded by the calyx lobes. 

The somewhat varying descriptions of Parosela Jamesii indicate that 
more than one species has been distributed under thisname. That the species 
now proposed is the Dalea Jamesti of Porter, F7. Colorado, seems probable, 
but from the original as described in T. & G. FZ. V. Am. 308, the following 
points seem to separate it. . Porteri is more prostrate spreading, the 
leaves are narrower and are shorter than the petioles, the pubescence is 
Coarser, it has purple bracts, the flowers are yellow not purple, and the wings 
and keel are oval not oblong as in P. Jamesii.* 

Collected at Berwind, Colorado, 1900, by Jennie M. Archibald, no. 244. 

Petalostemon pubescens, n. sp.— Roots woody; the crown 


_thick, simple or branched: stems numerous, the exterior ones 


*Dr. Gray in redrawing the description for Pl. ce 49, seems to have so modi- 
fied the original description as possibly to include P. Porter7. 


396 BOTANICAL GAZETTE [JUNE 


prostrate-assurgent, the central suberect, 1-2° long, whitish with 
a sublanate pubescence: leaves neither glandular nor dotted, 5- 
foliate, nearly glabrous above, the pubescence underneath and 
on the petioles resembling that on the stems; leaflets spatulate- 
linear, sessile, 1o-14™™ long; petioles about as long, often with 
a pair of diminutive trifoliate leaves in the axils; stipules fili- 
form: spike oblong, dense, 2-5 long, 1™ or less in diameter ; 
peduncles short (1-5°™): bracts scarcely equaling the calyx, 
obovate, abruptly acuminate, the dark purple tip obscured by the 
copious silvery lanate pubescence of the spike as a whole: calyx 
4-5™™ long, the teeth shorter than the tube: petals light purple; 
the limb of the standard broadly cordate, about 3™™ in diameter, 
exceeded by the filiform claw: ovary and base of style softly 
pubescent. 

Most nearly allied to P. tenuifolius Gray, from which it is at once to be 
distinguished by its very different habit, shorter broader leaflets, shorter 
peduncles and slenderer spikes, obovate short-acuminate purple tipped 
bracts, light purple petals, and the almost entire absence of glands or dots 
anywhere on the plant. _ 

The species is founded on no. 247 from Berwind, Colorado, by Jennte 
M. Archibald. 

Gentiana Moseleyi, n. sp.— Perennial from slender fleshy root- 
stocks, about 12" high, glabrous throughout, 1—2-flowered (pos- 
sibly more): leaves about 4 pairs, two of: which are basal and 
nearly verticillate, from narrowly oblanceolate and petioled 
below to broadly linear and sessile above, 2—-3°™ long: the ter- 
minal flower subtended by a pair of foliar linear-lanceolate bracts 
often nearly as long as the calyx; additional flowers (if any) 
axillary in the uppermost leaves and smaller: calyx green with 
purple streaks, about 15™™ long; the lobes triangular-lanceolate, 
equaling or exceeding the tube: corolla dark blue, about 25"” 
long; the lobes oblong, obtuse, exceeding the tube (15™™ long); 
5-7™™" broad, conspicuously long setaceous-lacerate on the sides 
and often nearly to the summit which is,obscurely dentate ; the 
tube arachnoid woolly within especially toward the base: fila- 
ments stout, flattened, equaling the tube: stipe half as long as 
the elliptic-oblong capsule: style almost wanting. 


- ewe I ee ee ee es ee a re eee ae NS Yo i 


1901] ROCKY MOUNTAIN PLANTS 397 


The habit and general appearance of this excellent species is much like 
G. elegans unicaulis Aven Nelson. In fact at first glance I thought of it as 
that, but no one really stopping to look at it can possibly mistake one for the 


‘other. G. Mose/eyi is certainly a perennial ; the conspicuous fringe upon the 


corolla lobes which exceed the tube and the sessile stigmas at once mark it as 
distinct from the other. 

The specimens were collected by, #. V. Moseley in Boulder county, Colo- 
rado, 1896. The exact locality is not known, but it probably was in some of 
the higher mountains. 

Polemonium Archibaldae, n. sp.—Stems tall (4-73), erect, 
slender, sparsely pubescent but not glandular except in the 
inflorescence: leaves glabrate, deeply pinnately divided, the 
midrib merely narrowly margined between the segments and on 
the very short petiole, 8-12°" long; the segments subpaired, 
13-19, oblong-lanceolate: inflorescence corymbose-cymose, 
minutely glandular-pubescent especially on the calyx; the small 
cymes (5—g-flowered) erect, congested (the pedicels very short) ; 
peduncles slender, the lower 5—10°™ long: calyx campanulate, 
6-7™™" long, the lanceolate lobes as long as the tube: corolla 
rotate-campanulate, Io-14™™ in diameter, the lobes suborbicular, 
as long as the tube, from deep purple to much lighter: filaments 
filiform, a little shorter than the corolla, glabrous except at the 
insertion where they are densely and finely pubescent; anthers 
oblong, large, 3™ long: stigmas scarcely exserted. 

This species is closely related to P. f/icitnum Greene, but differs from that 
in the somewhat narrower leaves, in the ampler inflorescence, in orbicular 
(not acute) corolla lobes, in the relatively shorter calyx lobes, in its erect (not 
declined) stamens, and in its shorter style. The inflorescence is probably also 
much less glandular, for Dr. Greene characterizes P. f/icinum as being densely 
glandular-viscid in the inflorescence. 

The species is dedicated to Jennie M. Archibald, who secured at Berwind, 
Colorada, in 1900, a most interesting collection. The type is in the Rocky 
Mountain Herbarium, no. 220 of the collection mentioned. 

Monarda Nuttallii, n. n.— Wonarda aristata Nutt., Trans. Am. 
Phil. Soc. n. ser. §: 186. 1837; not Monarda aristata Hook., Bot. 
Mag. t. 3526. 1836. This has been for some time included in 
Monarda citriodora Cerv., but any suite of specimens of the two 
Shows that they are distinct. M. citriodora is a larger, coarser 


398 BOTANICAL GAZETTE [JUNE 


plant, with broader leaves; the bracts more deeply colored, 
longer, more abruptly aristate and less strongly ciliate than in 
M. Nuttali. The latter has the mid-nerve of the bracts rela- 
tively much more conspicuous than the others, and the aristate 
calyx lobes strikingly ciliate-bearded. 

M. Nuttallit is well represented by the following specimens, all from Col- 
orado: 166, Professor Francis Ramaley, Boulder, 1900; C. S .Crandadl/, Fort 
Collins, 1893; Marie Holzinger, Boulder, 1892; 614, Baker, Earle, and 
Tracy, Durango, 1898 ; 428 of the Hall and Harbour plants. 

In this connection [ would inquire why Monarda aristata Hook. should 
not be restored as a substitute for 4/7. clinopodioides Gray? 

Monarda Ramaleyi, n. n.— Rootstocks horizontal, rather 
slender, abundantly and conspicuously rootbearing: stems erect, 
or sometimes decumbent and somewhat flexuous, mostly simple 
and monocephalous, 3—6"™ high, glabrate below, softly lanately 
white-pubescent upward, densely so at the nodes, more or less 
purplish: leaves lanceolate or ovate lanceolate, broadly rounded 
or subtruncate at base, sharply but rather remotely serrate, softly 
pubescent especially below; petioles very short, less than 1™, 
the uppermost almost wanting, lanately pubescent as are also the 
midribs of the leaves: involucral leaves about 6 or, counting the 
approximated uppermost stem pair, 8, ovate, acute, entire, 2-3™ 
long, obscurely tinged with purple: floral bracts slenderly linear- 
subulate, ciliate-hispid on the margins, about 1™ long: calyx 
tubular, minutely puberulent, 8—1o™™ long, the five equal subu- 
late teeth very short and somewhat hispid-pubescent at base: 
corolla lilac-purple, softly pubescent, strongly bilabiate; the 
tube within the calyx very narrowly linear, dilating gradually 
above, the exserted portion exceeding the calyx; upper lip lin- 
ear-lanceolate, but slightly curved, 10-12™" long; lower lip 
oblong-elliptic, as long at the upper, obtuse and broadly emargi- 
nate at apex, with a short, broadly linear, cucullate, pubescent 
appendage from the notch; the appendage bidentate: stamens 
2, exserted, exceeded by the pistil. 

This species at once presents a very different appearance from the only two 
species with which it can be compared, viz., MZ. menthacfolia Graham and M. 
stricta Wooton. On Suggesting to Professor Ramaley that it was probably 


eS ene a i pe 


Igor | ROCKY MOUNTAIN PLANTS 399 


new, he very kindly supplied me with a good series of specimens from which 
the characters are drawn. It may at once be distinguished by its rather low, 
strict, usually monocephalous stems, its unusual pubescence, its almost 
lavender corollas, and the peculiar appendage-like appearance of the middle 
lobe of the lower lip. 

I am informed that it occurs rather sparingly in sandy soil near Boulder 
creek, Boulder, Colorado, where it was collected by Professor Ramaley (to 
whom it is dedicated) June 27, 1goo, no. 92. 


PHACELIA HETEROPHYLLA Pursh. The Rocky mountain speci- 
mens of this species seem somewhat aberrant, but the characters 
so far as made out do not seem to warrant separation. 

Mimutus Lancsporriu Titinct (Regel) Greene? A pecu- 
liarly pubescent form was sparingly represented. It seems 
probable that fuller material may show this to be distinct. 

LITHOSPERMUM LINEARIFOLIUM Goldie was represented by 
several numbers and showed much diversity. However, since 
these numbers represented different localities and plants in dif- 
ferent stages of maturity no dependence can be placed on the 
diversity shown. In this species and its allies successive col- 
lections shouid be made during the season from what 1s 
undoubtedly the same community of plants. 

CHRYSOPSIS FULCRATA Greene. What seems to bea pretty 
good match for the type number of this species, originally 
described from New Mexican material, was collected near 
Boulder, 

Oonopsis monocephala, n. sp —Caudex woody, its branches 
Slender: stems one or more from each branch of the caudex, 
frect or ascending, 1-2™ high, simple, uniformly leafy from 
base to summit, monocephalous: leaves glabrous as are also the 
stems, linear-oblong, sessile, acute, quite uniform in size and 
Shape, 5—g™ long, about 1™ broad, the uppermost involucrating 
the large head: involucre hemispherical, about 15" high; the 
bracts oblong, rather abruptly short acuminate-cuspidate, glab- 
rous but for some ciliate pubescence on the margins: heads 
tayless; disk flowers very numerous: pappus nearly as long as 
the corolla tube, the bristles widely spreading. at maturity: 
akene oblong, about 3™™ long, sparsely and obscurely glandular. 


400 BOTANICAL GAZETTE [JUNE 


I am aware in pronouncing this plant an Oonopsis that I am putting into 
this genus a monocephalous species and another rayless one, but the habit is 
thoroughly characteristic. The hue of the leaves, which retain their foliar 
character up to the heads which they surpass, the appearance of the involu- 
cre, and the floral characters are in perfect accord with the genus. 

The specimens upon which the species is founded were submitted to me 
by Professor Ramaley, of the University of Colorado, and are the collection 
of Jennie M. Archibald, at Berwind, Colorado, in Igoo, no. 257. 


WESTERN EUPATORIEAE. 

Eupatorium atromontanum, n. sp.—Tall, 1-2™ high: stems 
with an obscure short white puberulence, striate, greenish or 
mottled with purplish-brown: leaves verticillate in fours, short 
petioled, ample, from ovate-acute to lanceolate-acuminate on 
the same plant, 12-25°™ long, 4-10™ broad, much exceeding the 
internodes, coarsely serrate, the teeth broadly triangular and 
apiculate, rather thin, prominently veined below, glabrate above 
and dark green, lighter and with a short fuscous puberulence 
underneath, sprinkled with minute, shining, resinous dots: inflo- 
rescence corymbose-cymose or more paniculate, equaled or 
more often overtopped by the uppermost leaves; the peduncles 
of the cymes arising from the axils of the leaves of the 2 or 3 
uppermost verticils cinerous-pubescent as are also the numerous 
short slender pedicels: involucral bracts in 4 or 5 series; the 
outer short, elliptic ; the second and third more broadly elliptic or 
oval, about 7™™ long, pubescent, with 3 greenish striae and scari- 
ous Margins; the inner and longest becoming oblong and almost 
wholly scarious: heads about 15-flowered: corollas tubular 
(hardly at all dilated upwards): akene obscurely angled, dotted 
with resin particles, linear, but slightly shorter than the some- 
what unequal pappus. 

This finds its nearest ally in the somewhat polymorphous £. maculatum 

The more typical form of that differs from this in the thicker, usually 
rugose leaves which at the summit become smaller or bract-like, leaving the 
inflorescence naked (surpassing the leaves), That also has a harsher, more 
copious pubescence, different involucral bracts, and fewer flowers in the head. 

E. atromontanum is represented in the Rocky Mountain Herbarium by 
two collections, as follows; 2251, Sheridan, July 10, 1896; 2553, Beaver 
creek, Black hills, Wyoming, July 30, 1896 (type). 


901] ROCKY MOUNTAIN PLANTS 4OI 


Coieosanthus congestus, n. sp.— Roots fascicled, large, semi- 
fleshy, with a woody crown: stems several to numerous, ascend- 
ing or nearly erect, simple, 3-6%" long, minutely pubescent, 
obscurely striate: leaves opposite below, only the uppermost 
alternate, green and nearly glabrous, or obscurely roughened- 
scabrous, especially on the veins below, lightly sprinkled with 
resin particles, deltoid-triangular, with truncate or cordate base, 
from sub-acute to acuminate, 3-6™ long, crenately-serrate, the 
apex entire; petioles variable, very short to equaling or sur- 
passing the blade: inflorescence congested at the summit of the 
simple stems, the crowded cymes of the few heads subtended by 
foliar bracts, sometimes one or more additional cymes terminal 
on widely divaricate or ascending peduncles from the axils of 
the uppermost leaves: heads 10-12™™ high, subtended by a few 
loose long-acuminate bracts; the involucral bracts in about 5 
Series, the outermost ovate, acuminate, pubescent and ciliate on 
the margins; the succeeding rows passing from broadly lanceo- 
late to broadly linear, green between the 3 or 4 raised white striae 
and ciliate on the scarious margin: flowers about 50: corolla 
narrow, tubular, about 7™ long: anthers wholly included within 
the tube proper: style branches spatulate-clavate, the exserted 
portion about 3™ long: akenes linear-cylindric, subglabrous, 
10-striate, 3-4™™ long. 

This is undoubtedly, in part, the Bricke//ia grandiflora of Gray, Syn. F7. 
105, but is certainly distinct from Eupatorium(?) grandifiorum Hook. F7. 
2:26, and probably from Brickellia grandiflora Nutt. Trans. Am. Phil, Soc. 
n. ser. 7:287. The original seems to have been a tall, probably single- 
stemmed plant, paniculately branched at the summit, with larger, fewer- 
flowered heads, and white corollas. This Oregon plant is probably in large 
part the Colcosanthus grandiflorus of the recent Flora of N. W. Amer ed 203, 
by Howell, who writes from near the type locality. The tufted habit, simple 
stems, congested inflorescence, and more numerous flowers of C. congestus 
seem to me to be specific characters. 

The collections at hand are: 423, Fairbanks, July 10, 1894; 1687, Cum- 
mins, July 16, 1895; 4212, Battle lake, August 17, 1897; 7506, Antelope 
basin, July 8, 1900 (type); also Palmer lake, Colorado, August 12, 1896, 
by Professor C. S. Crandall. 


402 BOTANICAL GAZETTE [JUNE 


Kuhnia Fitzpatricki, n. sp—Stems I-several, with a woody 
root, erect, 1™ (more or less) high: stems simple, rather slen- 
der, light green, obscurely striate, scarcely puberulent: leaves 
numerous, very uniformly distributed except near the base 
where they are early deciduous, green and appearing glabrous 
but puberulent or minutely scabrous especially on the margins 
or veins, thin, not impressed punctate and the resin particles 
microscopic, very uniform in shape and size, narrowly lanceolate, 
the cuneate base subpetiolate and with 1 or more linear bracts 
in the axils, 8-12 long, 8-15™™" wide, coarsely and somewhat 
uniformly toothed, upwardly becoming narrower, entire and in 
the short inflorescence somewhat reduced: cymes few and of few 
‘heads, crowded: pedicels and involucre softly pubescent; the 
outer bracts filiform; the inner broadly linear, acuminate: 
corolla purple(?), tube short, very slender, dilated at base, 
expanding into a much longer cylindrical throat: akene fusi- 
form, 5™ long, equaled by the very fine white pappus. 

This remarkably distinct species from the prairies of Iowa, I have from 
IT. J. and M. F. L. Fitzpatrick, who collected it in Decatur county, Septem- 
ber 25, 1899, where it is said to be common. It was distributed as KX. eupa- 
torioides L! (var. ?). 

Kuhnia Gooddingi, n. sp.— Perennial, from a densely caespi- 
tose-tufted caudex: stems very numerous, loosely spreading, 
5-8 high, simple and almost leafless below, striate and minutely 
puberulent: leaves numerous, crowded on the middle portion of 
the stem, sparsely and minutely puberulent especially on the 
margins and veins, linear to narrowly lanceolate, 2-4™ long, 
entire or nearly so, more or less revolute-margined, minutely 
impressed-punctate on both surfaces, those of the inflorescence 
gradually reduced and linear-bracteate: inflorescence loosely 
and widely paniculate, occupying the upper third of the plant: 
the pedicels and involucre finely pubescent: involucres turbinate, 
the bracts in about five series; the outer short, lanceolate; the 
inner broadly linear, with villous-pubescent setaceous tip, about 
4-striate, sparsely sprinkled with shining resinous particles as 
are also the corolla teeth: disk 10-12™™ high, I 5—25-flowered ; 


1901] ROCKY MOUNTAIN PLANTS 403 


corolla ochroleucous or shading to brown, slender, tubular; 
akene linear, finely about 2o0-striate, only a little shorter than 
the white finely plumose pappus. 

No western species of Kuhnia have heretofore been described. They 
occur but rarely, but at least two good species seem to belong to the eastern 
Rocky mountains. This species now proposed is one of the strongest of 
these, and differs so radically in habit, to say nothing of the other characters, 
that it seems unnecessary to call attention to the points of divergence from 
the well known species. 

It was secured by Mr. Leslie Goodding, to whom it is dedicated, in a 
mountain valley, West Dry creek, Larimer county, Colorado, where it 
occurred in the greatest abundance, usually in stony places, but with the 
tufted stems and woody caudex imbedded in low mounds of loose drifting 
soil. The type number is 

Kuhnia Hitchcocki, n. sp.— Tufted-suffrutescent, the lignes- 
cent stems decumbent-spreading, each divaricately branched 
from the base up, dark in hue but minutely puberulent as are 
also the leaves: stem leaves mostly wanting (or if but little 
branched, very numerous), nearly linear, with a few sharp teeth, 
3-4™ long; those of the rigid branches numerous, small, linear, 
aoa long: passing into the bractlets: involucre usually sub- 
tended by a few linear bractlets, its bracts in about 5 series, the 
short outer ones broadly lanceolate, the inner broadly linear 
with thin scarious margins and cuspidate apex: disk corollas 
20-30, the tube nearly uniform: akenes small, about Io-striate, 
Shorter than the dull-white to tawny distinctly plumose 
pappus. 

This species seems to belong to the arid plains east of the Rocky moun- 
tains. It has been frequently collected by Professor A. S. Hitchcock (to 
whom I am pleased to dedicate the species) in several of the counties of 
Kansas. Among the large series of specimens that his courtesy has enabled 
me to examine, the following may be named as typical: 211a, Meade county, | 
September 1897; Edwards county, September 1897; Stafford county, Sep~_ 
tember 1897; Clark county, August 1896; Kiowa county, August 1896; 
Barton county, August 1895. 

Kuhnia reticulata, n. sp.—Caudex woody: stems several or 
many, ascending, 3-4 high, simple, light green, finely puberu- 
lent: leaves light’ green, appearing glabrous but puberulent on 


« 


4 


404 BOTANICAL GAZETTE [JUNE 


the upper face and on the veins and margins, closely impressed- 
punctate on both faces, the veins on the lower surface raised- 
reticulate, oblong-lanceolate, 2-4°™ long, subacute, from entire 
to irregularly few toothed, usually some small fascicled linear 
leaves in the axils, smaller upward and passing into the bracts 
of the short inflorescence: cymes of few heads, congested- 
corymbose or becoming paniculate: involucre turbinate-cam- 
panulate; the bracts in 4 or 5 series, softly pubescent, strongly 
nervose, resin-sprinkled, shorter than the 12-15™™" high disk; 
the outer short, lanceolate; the inner broadly linear, acute: 
flowers about 25: the corollas linear-cylindric, but slightly 
dilated at base, about 8™ long: style tips compressed, linear, 
not dilated-clavate at summit: akenes 20-striate, some of the 
striae more prominent than the others, linear, about 5™™ long, 
exceeded by the white softly plumose pappus. 

This species is perhaps most nearly allied to A. g/utinosa Ell., but the 
points of distinction are so patent that to call attention to them seems 
unnecessary. 

The only collection of it is no. 2771, Platte cafion, Laramie county, 
Wyoming, August 27, 1896. Some specimens of it were distributed as A, 
eupatoriotdes corymbosa T. & G. 

Lacinaria alata, n. sp.—tTall, 8°" to probably twice as high: 
stem stout, striate, puberulent: leaves numerous and crowded 
(especially on the middle portion of the stem), quite similar 
throughout (root leaves not seen), glabrous, closely and uni- 
formly punctate, linear-oblong, acute; the lower 8-12 long, 
mostly less than 1™ broad, deflexed on the short margined 
petioles, upwardly becoming smaller sessile and erect, passing 
into the large foliar bracts of the spike: spike 3—5%™ long, 
crowded: heads sessile or nearly so, numerous (30-50), about 
15™™ high, equally broad, 35—45-flowered: the involucral bracts 
in about 6 series, minutely punctate, purplish; the outer very 
short, ovate, acute, green, rarely slightly scarious-winged near 
the summit; the succeeding 2 or 3 series similar as to the 
foliaceous center but conspicuously and broadly scarious-winged 
toward summit, becoming broadly obcordate in outline, the 
wings from entire to dentate or even lacerate, about 1™ long 


Igor] ROCKY MOUNTAIN PLANTS 405 


and broad; the inner series longer and narrower, scarious mar- 
gined all around: corolla resin-dotted throughout, 8—-1o™™ long, 
the broadly linear lobes half as long as the tube which is stiffly 
pubescent within near the base: style branches as long as the 
corolla; akene narrowly obovate, dark brown with short whitish 
spreading pubescence, about 5™™ long: receptacle nearly plane, 
with shallow alveolations. 

The plants upon which this species is founded were collected 2, ae ane 3 
Kimmons, August 1895, in the Creek Nation, I. T., and distributed by Pro- 
fessor J. W, Blankinship as Liéatris scariosa Willd. From this there are 
many Characters to separate it, but attention need only be called to the 
reflexed lower leaves, the long foliar-bracted spike, and the alate bracts of 
involucre. 

Lacinaria ligulistylis, n. sp.— Perennial, from an enlarged 
woody tuberous root: stems single, light green, glabrate below, 
lightly white pubescent above, striate, 4-5" high, uniformly 
leafy from the base up: leaves bright green, rather obscurely 
punctate, glabrous, usually minutely ciliate on the slightly thick- 
ened margin; the lower lance-oblong, 8—12™ long, tapering into 
a usually much shorter margined petiole; upwardly becoming 
more lanceolate, the winged base shorter, gradually smaller, 
passing into the lance-linear bracts of the inflorescence: heads 
from few to several, racemose: peduncles 1.3% long, tinged 


with purple as are also the peduncle and rachis: involucre 


broadly campanulate, often 20-25™" broad; bracts in about 6 
series, foliar-green with dark purple scarious dentate margins; 
the outer short, from nearly orbicular to oval; the middle rows 
broadly obovate; the inner elliptic or oblong and 15-18™™ long: 
flowers 50-70, purple: corolla tube slightly dilated upward; the 
teeth lanceolate, one third as long as the tube, with a delicate 
marginal vein: exserted style branches flattened, as long as the 
Corolla, light purple, conspicuous and superficially suggesting the 
Slender rays of some Erigeron; akene flattened, finely pubescent, 
Nearly as long as the corolla tube. 

The Rocky mountain forms of Lacinaria that have passed as L. scariosa 
(L.) Hill are clearly distinct from the eastern forms. That, even with the 
Rocky mountain form eliminated, L. scariosa, as applied to the eastern forms, 


406 BOTANICAL GAZETTE [JUNE 


is still an aggregate seems to me more than probable. In spite of the 
extended synonomy of this species, the various names all seem to be asso- 
ciated with the forms of the eastern or the Mississippi valley states. Some of 
the descriptions of Z. scariosa, however, probably cover or may even be 
founded upon the western or Rocky mountain form, notably that of Hooker, 
Ho ie Sele a) 6 

The species now proposed is at once distinct from the Serratuda scariosa L. 
and the Liatris scariosa Willd., and surely from the spicate forms of the 
Mississippi valley. The smooth leaves gradually diminishing upwards, the 
relatively short margined petioles also diminishing upward, the clearly 
racemiform purple inflorescence, the large heads, the appressed involucre, 
the very conspicuous style-branches, and the numerous flowers will serve to 
distinguish ZL. ZigulistyZis. 

The following collections are cited as typical: 1651, Laramie peak, 
August 8, 1895 (type); 691, by C. F. Baker, Pagosa springs, Colorado, 
August 30, 1899; 4554, by &. H. Snow, Colorado; G. £. Osterhout, Steam- 
boat springs, Colorado, August 24, 1899; less typical my numbers from 
Inyan Cara divide, August 20, 1892; and Buffalo, July 26, 1896. 


STUDIES IN ARNICAS. 

The collector and the student of the Rocky mountain Arnicas 
has in the past found many difficulties with which to contend. 
So many of the species attributed to this region were originally 
described from such widely different geographical ranges that it 
ought to have been expected that no close agreement would be 
found between our species and those of the earlier investigated 
localities. The recent activity in taxonomic botany is bring- 
ing to light many real novelties and separating many of the 
heretofore uncertain but well-known forms. It is perhaps 
inevitable that at such a time some confusion should result, 
but perhaps the sooner synonyms and homonyms are cleared 
up the better. The following have come under my obser 
vation : 

Arnica multiflora Greene, Pitt. 4:162, evidently is A. 
Columbiana Aven Nelson, Bot. Gaz. 30: 200, since both are, in 
part, founded on the same collections and the same numbers are 
cited. The latter name is the earlier by two or three months. 
Dr. Greene’s A. Columbiana (Pitt. 4:159) having thus become 
a homonym it may become Arnica Greenei, n. n. 


1901 | ROCKY MOUNTAIN PLANTS 407 


Arnica platyphylla, n. sp.— Perennial from woody rhizomes, 
3-6" high, rather slender, the internodes subequal, nearly glab- 
rous throughout, the leaves very thin and delicate in texture: 
the leaves that spring in fascicles of 3-5 from the rhizome oval 
to oval-subcordate, sinuately dentate as are also the stem leaves, 
obtuse or acute, ciliolate on the margins, 4-6™ long, equaled 
by the slender petioles: stem leaves 2~4 pairs and a pair of 
sheathing scales at base, all closely sessile by broadly rounded 
or truncate base; if only 2 pairs, similar in size and shape; if 4 
pairs, the upper and the lower smaller than the two middle pairs ; 
from nearly orbicular to broadly ovate and subacute, 5-10™ 
long: heads 3—s; if three, on slender nearly equal peduncles, 
subtended by ovate-acuminate bracts 2-3°" long; if five, the 
lowest pair from the uppermost leaves and borne on elongated 
slender petioles: involucres somewhat turbinate, slightly gland- 
ular-pubescent as are also the peduncles; scales thin, narrowly 
obovate-acuminate, in one series: rays few, long, bright yellow: 
tube of disk corollas pubescent, very slender, abruptly dilated 
into a wide cylindrical glabrous throat with slender erect teeth: 
akenes narrowly linear, 5™™ long, equaling the fine white hardly 
scabrous pappus. 

This is seemingly a well marked species, occurring in the moist woods of 
the northern Rocky mountains. I have seen four collections of it as follows: 
flenderson, Cascade mountains, Oregon, July 18, 1896, distributed as A. /att- 
Jolia,;, Sandberg and Leiberg, Stevens pass, Washington, distributed as A. 
Chamissonis, August 1893; R. S. Williams (no. 443), Columbia falls, June 
18, 1894; Belt mountains, Montana, July 24, 1899. Of the latter two I cannot - 
be very certain, as they are not now at hand. This species seems nearest to 
the recently described 4. ventorum Greene (based upon one of my numbers), 
a species of which I now have several collections. 

Arnica stricta, n. sp.—Very erect with gradually tapering 
stem, 2-34" high, tomentose pubescent throughout: leaves about 5 
pairs, the lower pairs exceeding the internodes, gradually smaller 
upward, the upper half of the stem nearly naked; the lowest 
pair lanceolate, acute, 3-5 ™ long, tapering into a slender petiole 
usually shorter than the blade; the next 1 or 2 pairs similar 
but with shorter petioles; the upper and smaller pairs sessile by 


408 BOTANICAL GAZETTE [JUNE 


a broad base: inflorescence of about 5 heads subtended bya 
pair of ovate-lanceolate bracts, 1-2°™ long, in a close rounded 
corymb, the terminal head and the first pair very short peduncled: 
terminal head much the largest, 1o-12™" high and about as 
broad: involucral bracts in 2 series, shorter than the disk, nar- 
rowly oblong, obtusish, the tip tinged with purple and tomentose- 
ciliate: rays few, short, broad, slightly narrowed at apex: tube 
of disk corollas pubescent, the glabrous throat scarcely dilated: 
akenes very minutely pubescent; pappus dirty-white, minutely 
scabrous. 

The specimens on which this species is based were collected in 1892 by 
Mr. J. D. Parker, somewhere in southern Wyoming. Specimens were sent to 
the Gray Herbarium for determination. The name returned was A. foliosa 
near var. incana Gray. Now that Dr. Greene has helped us to a more defi- 
nite knowledge of A. incana (Pitt. 4: 169), this may with safety be designated 
by a name. 

Arnica celsa, n. sp.—Perennial from horizontal rootstocks 
and fleshy roots, erect, tall, 4-64™: the strict stems lightly 
striate, sparsely pubescent with flat spreading hairs, upwardly 
becoming also obscurely glandular-viscid: leaves (exclusive of 
the foliar bracts) 6-8 pairs, puberulent and sprinkled with micro- 
scopic shining resin particles, narrowly oblong or oblanceolate, 
tapering to both ends, many of them 3-nerved; the lower small, 
2-3°" long, equaled by the slender petioles which abruptly 
dilate into the equally long ocreae; ocreae and leaves gradually 
shorter upward; middle stem leaves longest (8-10), sessile or 
with short petioles and ocreae; the uppermost sessile and partly 
clasping: lower internodes short, much exceeded by the leaves, 
gradually longer upwards and above much exceeding the leaves: 
heads 3-5, on rather slender leafy-bracted peduncles; involucre 
and peduncles moderately glandular-viscid; the bracts approxi- 
mately uniserial, oblong, much shorter than the 12-15™” 
high disk: rays 12-20, about 7-nerved, 2 or 3-toothed, 1o-I14™™ 
long: tube of disk corollas slender and pubescent below, grad- 
ually dilated into an equally long glabrous sub-cylindric upper 
portion with short spreading glabrous teeth: akenes sub-glabrous, 


4 


1901 | ROCKY MOUNTAIN PLANTS 4°99 


nearly linear, tapering downward, the scabrous pappus equaling 
the corolla tube. 

This excellent species was found growing in abundance on the wet, 
meadow-like banks of a mountain streamlet at an altitude of 8000 feet. The 
nearest ally known to me is A. ocreata Aven Nelson, from which it differs in 
its pubescence and wholly different disposition of its leaves. The habit of 
the inflorescence, too, is very unlike, which in the species now proposed con- 
sists of subequal heads always corymbosely disposed. 

No. 7643, Tie city, Albany county, July 20, 1goo, is the type. 

Arnica rhizomata, n. sp.—Conspicuously rhizomatous, the 
creeping rootstocks slender, sheathing-bracteate at the nodes, 
giving rise at intervals to the erect leafy stems and a few fleshy 
roots: pubescence whitish, soft, almost arachnoid, minutely 
granular-glutinous underneath the pubescence on the inflores- 
cence: stems 2-5“ high: leaves 5—7 pairs, rather uniformly dis- 
tributed, oblong-lanceolate, from obtuse to acute; the lowest 
pair with short scarious-margined petioles, early deciduous; the 
next I or 2 pairs 6-g™ long, about 2™ broad, on short mar- 
gined petioles which dilate at base to form the short ocreae; 
the upper pairs sessile, shorter: heads 3-5, 10-12™™" high, or 
Sometimes more numerous and then smaller: peduncles mostly 
short, erect, the lateral often exceeding the terminal : involucre 
campanulate, the bracts much shorter than the disk, narrowly 
oblong, obtusish: corolla long-pubescent on the tube, sometimes 
a few straggling hairs on the lobes: akenes linear, almost glab- 
Tous, the pubescence short and scattering, half as long as the 
8™" corolla: pappus fulvous or dirty-white. 

This species has been collected several times and has been held as a 
form of 4. foliosa, or rather as variety zacana. Such a disposition is 
no longer tenable, neither am I able to refer it to the species (A. foliosa 

utt.). It seems rather to belong to a section of which A. ocreata, A. celsa, 
and this are the principal members. In so far as this region is concerned, 
the following collections répresent it: 8012, Lincoln gulch, Albany county, 
August 8, 1900 (type); 1417, B. C. Buffum, Pine creek, 1892; Green moun- 
tain, July 6, 1896; 3587, North Vermillion creek, July 17, 1897. 
UNIVERSITY oF WYOMING, 
Laramie, Wyo. 


STUDIES ON THE GEOTROPISM OF STEMS. II." 
EDWIN BINGHAM COPELAND. 
(WITH THREE FIGURES) 


THE review of my earlier papers on this subject, written by 
Czapek for the Botanische Zeitung, is in error on a point of 
priority and on the interpretation of the results. Sachs never 
showed that if a stem be split and the halves placed horizontal the 
under one would grow faster. In Sachs’ experiment referred to in 
the review the average elongation of the two faces of the upper 
half stem was 9.7™™, that of the lower half only 9.0™™. If the 
experiment had been continued longer (see footnote Bort. GAZ. 
29: 189. 1900) the relation would have been reversed. The 
review concludes: 

Insbesondere kann der Ref. sich der Ansicht des Verf. dass diese 
Erscheinung alle Theorien, welche den Sitz der geotropischen Reactionen 
in beiden Langshalften annehmen, ausser Kraft setze, nicht anschliessen. 

I-do not hold and have never expressed such a view. The 
phenomenon in question proves that both halves do share in the 
reaction, which therefore does not take ‘‘place entirely in either 

alf.””. As Czapek construed my conclusion, it is no wonder 
that he disagreed. 


3. Positive geotropism in the hypocotyl or cotyledon. 


The descending axis of a plant must grow and become fast in 
the ground before the negative geotropism of the ascending ax!s 
can result in the erection of the growing tip. That the former 
does break first from the seed is a matter of universal observa- 
tion. In the popular conception this first outgrowth is simply 
the “‘root;” but that this idea is not exact is recognized, for 

* The first number appeared in Bor. Gaz. 29: 185-196. 1900. 

? Bot. Zeit. 58: 200. 1900. 

410 [JUNE 


/ Seer 


1901] GEOTROPISM OF STEMS 4iI 


instance, by the disposition in many texts to substitute the word 
caulicle for radicle for the part of the embryo below the inser- 
tion of the cotyledons. In all Dicotyledons a part of the stem 
is distinguishable below the cotyledons, and in a majority it 
elongates to carry the cotyledons into the air. The elongation 
of the caulicle is the first visible growth. When the caulicle is 
largely a stem structure, and only the tip represents the future 
root, the beginning of growth is in the stem. In all cases the 
caulicle promptly bends downward. We have here apparently, 
then, positive geotropism of the hypocotyl, which so far as I 
know has not hitherto been recognized or investigated. 

Lupinus albus, in the size and uniformity of the seedlings and 
their rapid growth, is the best material I have found for this work. 
Both in the seed and on the seedling there is usually a well- 
defined line where the root merges into the stem. In the first 
cultures I marked this line with India ink on the swollen embryos, 
to be certain that it was the same one separating root and shoot 
in the seedling, and found that such was constantly the case. In 
order to have the entire descending axis free to curve, I usually 
removed the seedcoats, after swelling, but before any consider- 
able growth. This was not necessary for the mere demonstra- 
tion that a part of the hypocotyl bends downward, but when 
germination took place within the coats a part of the hypocotyl 
was often constrained by them. Their removal was probably 
without effect on germination or growth, though in an experi- 
ment to ascertain its influence the peeled plants grew slightly 
faster. 

One culture of plants is reported upon here in detail, to 
show the constancy of the results, and the range of variation 
that may be expected. The seeds were put into water 
‘October 1. On October 2, 9:00 A.M., the coats were removed. 
Fifteen very uniform embryos wefe selected and pinned with the 
axis horizontal. The length of the hypocotyl was 6™™, of the 
root 1.8™", The table shows the result after 24 hours. The 


_ last column, showing how far above the base of the hypocotyl 


the curve of least radius occurred, is the significant one. 


412 BOTANICAL GAZETTE [JUNE 


ay SRSA, ONES NS aS In all except three plants, in 
Hypocotr! | ootlength|" tescol” which it fell at the middle, the 
hypocotyl ’ 
——— sharpest curve was above the 
a, . me — middle of the hypocotyl (fg. 7). 
ee 9 4. In several of the plants the curve 
: 2. om began immediately at the inser- 
: * : tion of the cotyledons. Two of 
8 | to. 10. 7. the plants of this series were put 
ni : : into water culture vessels, and 
pes : is when the elongation of the hypo- 
13 | 9. 4 5. cotyl was finished, in a south win- 
- ~ oe a dow, the points where the radius of 
Ay.| 3.3mm 72mm | 5.6mm curvature had been least were 20 


and 22™ above the foot of the 
hypocotyl. The curve had disappeared In three plants of another 
experiment the part of the hypocotyl below the sharpest curve 
elongated to 17.5, 18.0, and 19.0", the curve being eliminated. 

With somewhat older seedlings the result is less uniform. 
With twelve plants whose hypocotyl was ca. 10™™ and the root 
ca. 2.0™™" long, placed horizontal for twenty-one hours, the result 
was that the sharpest curve in three hours was in the hypo- 
cotyl, in six hours at the line of separation, and in three hours 
in the root. Before the experiment the plants had been in 
imperfectly saturated air, which shortens the growing region, as 
compared with plants grown in water or saturated air.‘ 

The first attempt at an explanation of the behavior of the 
hypocotyl was by a study of its anatomy. If the transition from 
stem to root character in the internal tissues or their arrange- 
ment occurred higher up than the external line of demarcation, 
there would bea fine prospect of demonstrating the relation of 

3From clinostat experiments I know that both negative geotropism and recti- 
petality are concerned in this straightening. 

4Sacus, J.: Ueber das Wachsthum der Haupt- und Nebenwurzeln. -Gesammelte 
Abhandlungen 807. From Arb. bot. Inst. Wiirzbur re 1. 1874. 

Popovicl, A. P. (Bot. Centralbl. 81 : 33. 1900) ies that several but not all 


factors checking growth shorten the growing region. For influence of insufficient 
water see p. 95 


1901 | GEOTROPISM OF STEMS 413 


the particular tissues concerned in it tothe geotropic irritability. 
With this idea in view, I made a careful study of the histology 
of the root and hypocotyl. Seedlings which have just curved 
are not satisfactory material because of the immaturity of the 
tissues and the amount of plastic material present, and the 
observations on maturer stems which had formerly 
shown themselves positively geotropicmake much , 
study of very young plants superfluous. The @ NS 
apparent line of separation between root and 
hypocotyl is the real one, as to the epidermal 
tissues; the stomatas descend to it, and root 
hairs occur only below it. In the cortical paren- 
chyma there is no sharp distinction between stem 
and root, nor between different parts of the hypo- 
cotyl, not even when, as it matures, chlorophyll 
becomes more abundant in the normally better 
lighted upper part. In the stele® the rotation in 
the arrangement of phloem and xylem is gradual, 
but usually begins below the transition line in the 
epidermis, and is completed at some distance (in 
one very extreme case 17™™) down in the root. ’ 
There is then nothing at all in the histology of £o- Voune 
the hypocotyl that can give a clue to its geotropic Lae of Lupi- 
variability. This does not prove, however, that 2us adbus, showing 
the explanation is to be sought in the finer struct- raaebabe 3 a 
e hypocotyl. 
ure of individual cells. 

The truth is rather that the downward curve occurring in the 
hypocotyl is a response to,a stimulus perceived by the root tip. 
This is not easy of absolute proof, but the evidence seems to 
warrant fully the conclusion. The ‘bent tube”? method of 
demonstration is inapplicable because it is during the very first 
growth that the phenomenon in question occurs. Decapitation 


5 Near the base of the hypocotyl most of the stomata are considerably elevated 
above the general surface. 

SI use the word as a matter of convenience for the bundles and pith. It is not 
Sharply delimited from the cortex. 


414 BOTANICAL GAZETTE [JUNE 


| experiments are at best unsatisfactory, or they would have defi- 
nitely determined the functions of the root tip without contro- 
versy decades ago. Mine were as consistent as they are likely 
to be, or as can be expected from a method which does great 
violence to the health of the plant, and in which any unsteadi- 
ness in the operation may entirely alter the result. When no 
traumatic bending occurred, the growth of horizontal decapitated 
plants was straight for more than twenty-four hours, and nearly 
always for more than forty-eight. In the end there was usually 
a downward curving in the basal part of the hypocotyl, but 
never farther up, where the curve, if any, was in the other direc- 
tion. The downward curve usually occurred before a new grow- 
ing point became evident, as is the case too when the response 
is by the root.?. In every respect the part of the hypocotyl which 
would execute a positively geotropic curvature is influenced by 
removal of the root tip, just as the responsive zone of the root 
would be in older plants. The interval during which a wound 
suspends the geotropic perceptivity in older hypocotyls is a very 
few hours at most, instead of about two days. 

Better evidence than decapitation can yield is obtained from 
a careful study of the distribution of growth in length along the 
young root and hypocotyl, and its relation to the location of the 
curve. For a basis of comparison this relation is first shown with 
older plants, in which the root alone curves. The plants had 
been germinated two days, and were marked off in 1™ zones 
and placed horizontal at 10:30 a.m., November 19. The result 
at 4:00 P.M. the same day is tabulated. The first column is 
the length of the roots at the beginning of the experiment. 

After twenty-nine and a half hours the average growth had 
been 20™" and the curve was go°, at about the same place. 
After five and a half hours, at the first measurement, the most 
growth had been in zones 3 and 4, but I could not measure it 
accurately enough to include it in the table. The experiment, 
like all in which the contrary is not stated, had been carried on 
in saturated air. 


7 Literature in CZAPEK: Ueber den Nachweis der geotropischen Sensibilitat 
der Wurzelspitze. Jahrb. f. wiss. Bot. 35 : 314-5. 1900. 


i 


.” a 


Igor} GEOTROPISM OF STEMS 415 
Reve lencth |. .c h Zone most Cc Growing 
gt rowt cheved urve eaies 

I 20,7" 2.9™n 3-4 7 oe 
2 19. 1.5 5 70 
3 5. 1.8 4 go 
4 | 14 1.4 4 90 
5 24 1.8 4 60 
6 19 1.0 3-4 65 
ih 15 2.6 4 60 
8 22. 1.6 3-4 70 
9 | 14. 2.4 3-4 50 

Io 33- 3. 3-4 50 

Av | 18.52" | 2.0% 4 70° 5 


The subjects of the experiment whose result is next tabulated 
had been soaked twenty-four hours. The coats were removed 
and the “caulicle’” was marked off into 1™ zones, and the 
embryos were fastened horizontal. The first column gives the 
total length of the descending axis; of this the rudimentary root 
was I.g-2.5™™. Figures in italics are the zones of sharpest 
curvature. All zones of the hypocotyl grew in length, but the 
growth reached a minimum at some distance from the tip which 
I have construed as limiting the growing region of the “root.” 
The experiment began at 10:30 A. M., November Ig. At 4:00 
P.M., November 20, all seedlings were curved go”. 


4p. m. November 19 4p. m., November 20 
Length : 
Growth Curve - Growth sepia 
: Oe Coc = 20° 17mm 5 6(+) 
2 9 0.8 5 40 16 ¢ ‘ 
3 7 0.8 d 30 16 4 rs 
4 7 a2 3 20 16 5 5 
5 8 0.4 4-5 30 17 ee liis 
6 7 ? 2-3 10 II 4 4 
7 7 ? 3-4 10 14 5 5 
8 a ? 15 18 Z 4 
9 9 0.4 ¢ 15 15 4 4 
bdo) 8 0.5 40 15 4 4 
- 7 J 2-4 15 13 ba 
= 9 0.3 4-5 16 13 os | S 


416 BOTANICAL GAZETTE [JUNE 


The older plants, subjects of the preceding experiment, were 
growing so rapidly that the first observation (after five and a 
half hours) was too late to show the gradual retreat from the 
tip of the region of curvature. But in roots it always happens,® 
as in the base of the stem in these very young plants, that the 
curve first appears near the tip and gradually moves farther away. 
Of course if the growth is rapid so will be the development of 
the curve. 

In the base of the hypocotyl, then, the downward curve 
occurs only in a zone not separated from the root tip by any 
mature tissue, and not at most as remote from the tip as the 
extreme limit of what may fairly be considered the apical grow- 
ing region. In these points, and in all other visible details of 
the execution of the response, it agrees with the familiar mani- 
festations of positive geotropism in roots. From this and the 
decapitation experiments, and from the subsequent elimination 
of the curve by later growth, the conclusion 
| seems obvious that the response is to a 
stimulus received at the root tip. 

Similar but less complete observations 
have been made on Robinia Pseudacacia, 
Helianthus annuus, Cucurbita Pepo, Eucglyptus 
globulus, and Pinus ‘Sabian (fig. 2): The 
embryos of Robinia, with radicles 0.5™™, and 
hypocotyls 2.0™ in length, were fastened 
horizontal. In one day the hypocotyl 
became 5.4™™" (average of ten plants) long, 
and the root 5™™, the curve of least radius 

Fic. 2.—Young seedling being 3™" above the line of separation, that 
of Pinus Sabiniana, show- is, it was above the middle of the hypocotyl. 
ing downward curve in Two plants were placed in water culture and 
wiih the curve grew out. My observations 0? 
little to form the arch the anatomy have no value here, for the 
ses the cotyle- reason explained in the treatment of Lupi- 

eae nus. Decapitation was tried on Helianthus 


®Sacus, /. c., pp. 831, seg. 


. 
a 


1901] GEOTROPISM OF STEMS 417 


and Cucurbita, with results unsatisfactory, but, so far as they 
showed anything, agreeing with those on Lupinus. 

Uninjured Cucurbita freed from its coats behaves at first 
exactly like Lupinus, but later the hypocotyl sometimes becomes 
concave below, even when the root tip has grown some centi- 
meters away from it. The behavior of these older Cucurbita 
hypocotyls is exceedingly various, but can in no instance be 
closely related to the positive geotropism of very young hypo- 
cotyls. Sometimes the hypocotyl becomes concave on the side 
where the peg forms, apparently under the peg’s influence.? 
This is not geotropism at all, though the concave side is nor- 
mally beneath, because the peg grows there. Later the stem 
may become concave below, next to the cotyledons, in an 
attempt to form the arch which in nature breaks through the 
ground, pulling the cotyledons after.*° This is real and fixed 
geotropism, not epinasty or nutation (unfixed geotropism). 
Frank, Véchting, and others have described similar behavior in 
the peduncles or pedicels of various plants. That it is not 
dorsiventrality in Cucurbita is shown by its occurring inde- 
pendently of the plane in which the cotyledons are placed. But 
they subsequently become erect on the clinostat. The curve in 
the other direction (negatively geotropic), described for Cucur- 
bita in the first of these studies, is also early evident. Double 
curves are very common, often about equaling each other and 
leaving the direction of the root unchanged. The whole subject 
is not really relevant here, and I leave it in the hope that prom- 
ised papers by Pollock * and Noll” will explain my diversity of 
results. 

The phenomenon which this paper describes shows anew 
that the structure of stems does not demand that the geotropic 
response they execute shall be negative, and that root structure 


° DARWIN and Acton: Physiology of plants, 193 fig. 38A. 

7? DARWIN and ACTON: /.c. fig. 388. 

™ Prelim. paper before Soc. Plant Morph. and Physiol. Dec. 1899. 

%2 Sitz. ber. Niederrhein Ges. f. Naturf. u. Heilkunde zu Bonn (S. A.) p. 3. 1900; 
also Jahrb. £. wiss. Bot. 34: 461. 1900 


418 BOTANICAL GAZETTE [JUNE 


is not essential to the positive response; but that the more 
characteristic feature in both organs is the perception. The 
clearness of the distinction between perception and response 
demands that we discriminate in the use of words to name the 
two processes. Tropism itself is the disposition to respond by 
turning or bending, and has no necessary reference to the place 
or manner of the perception of the stimulus, more than that the 
two processes be some way connected in space and time. The 
young hypocotyl bends downward, and is therefore prosgeotro- 
pic. For the act of perception Czapek proposes the word 
‘‘aesthesis.’’*3 The root tip is prosgeoesthetic ; it cannot pos- 
sibly be geotropic because it cannot actively turn. The very 
young hypocotyl seems not to be directly irritable at all by 
gravity, but when it becomes so it always bends upward, that is, 
it is apogeoesthetic. With perception and execution, and 
transmission when the two other processes do not occur in the 
same place, the performance is complete. Czapek’s introduc- 
tion of a ‘“Reflex-centrum™ as a potential link between the 
organ of perception and that of response, and as a seat of inter- 
pretation and decision, seems to me unnecessary,’ and is with- 
out any empirical support. It is simpler and hence preferable 
to suppose that the organ of perception itself in the act of perceiv- 
ing determines the direction of the response, whether or not it 
itself executes it. The manifestations of irritability in plants, 
like those of instinct in animals, are very short-cut psychic 
processes, and constantly dispense with steps which would be 
necessary to the attainment of the same end by the exercise of 
intelligence. On many grounds, the perception should be 
regarded as the characteristic and decisive though invisible 
feature of the entire phenomenon in plant irritability. 

There is supposed to be a considerable number of instances 

13 Weitere ce zur Kenntniss der geotropischen Reizbewegungen. Jahrb. 
f. wiss. Bot. 32: se 1898. 


Zc. 294. ropisms were called “ Redccieacugen” by Oltmanns (Ueber 
die RE ah Bewegungen der Pflanzen, Flora 75: 265. 1892), but the elabo- 
ration of the idea is Czapek’s 


4S NOLL: Ueber Gunaieas Jahrb. f. wiss. Bot. 34: 492-496. 1900. 


1901] GEOTROPISM OF STEMS 419 


of alteration or reversal of irritability during the development of 
plants or their parts, independent of any changes in external 
conditions. At first sight the young hypocotyl of Lupinus 
seemed to present such a case, but investigation shows that 
there is no change in the nature of the perception in any 
esthetic tract, but merely change in the zone controlled. Very 
possibly, renewed study of some other cases, such as the erect 
tip of runners, or the tips of many climbers—at first seemingly 
positively heliotropic, but later appressed to the support —will 
show that no esthetic center undergoes any essential change; but 
rather that different parts of the growing region are under differ- 
ent control, or that the development of new members introduces 
automatic curvatures (nastie), or that the tendency to symmetry 
(autotropism, rectipetality) eliminates curves in parts which 
grow out of the zone of influence of an esthetic center. Nuta- 
tions too suggest themselves here. 

Most intimately related to the positive geotropism of the 
hypocotyl is that of the cotyledons of numerous monocotyle- 
dons, serving at the apex as organs of absorption in the endo- 
sperm, but elongating at the base and carrying the remainder of 
the embryo out of the seed. These have long been known to be 
positively geotropic.%* I have made the same experiments on these 
as on Lupinus, and so far as the initial downward curve is con- 
cerned with strictly analogous results. Several seeds were 
tried, of which Phoenix and Yucca angustifolia were found most 
suitable subjects. 

The growing region of Phoenix is rather long, though elonga- 
tion is considerable throughout most of it. The root begins to 
develop after the elongation of the cotyledon is about complete. 
The first table of results is from six plants of very different size, 
on four of which the root was still only a rudiment. The 

*©SacHs: Ueber die Keimung des Samens von Allium Cepa. Gesamml. Abh. 
oe from Bot. Zeit. February 1862. 

NK: Lehrbuch 1: 465. 
—. isa paper, which I know only by a very brief notice in Bot. Jahresb. 


15 : 622, by ee Spa: (Bidrag till hjertbladets anatomi hos Monocotyledonerna, in Sv. 
V. Ak. Bih, III. 12: —~. 1887), giving a list of plants with such cotyledons. 


420 


BOTANICAL GAZETTE [JUNE 


experiment began November 30, at 12 noon, the plants being 
marked into 1™™ zones and placed horizontal. In the first plant 


the growth was in the root; 


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“ A Ch + IUNO 


in the second, in the cotyledon and 
root; and in the others in the 
cotyledon. In the larger plants, 
with more rapid growth, the execu- 
tion of the curvature was prompter, 
and with the longer growing region 
the curve was farther from the tip; 
but it very evidently made no differ- 
ence whether the curve was in the 
root or the cotyledon. In either 
case it was obviously in the apical 
growing region, in immedidiate con- 
nection with the sensitive tip. It is 
interesting to note that in the 
youngest plants, when the cells of 
the apical meristem are probably 
not yet dividing actively, they 
already exercise their function in 
irritability. 

The next table contains nothing 
new, but is introduced to empha- 
size by numerous instances the fact 
developed in the preceding experi- 
ment that the curve in the cotyle- 
don is in exactly the same position 
in the apical growing region that 
it would be in older plants where 
the growing region is all in the 
root. In most of these plants the 
rudimentary root was less than 1™ 
long. The experiment began Janu- 


ary 16,9:00 A.M., when sixteen seedlings were marked and 


fastened horizontal; one proved unsound. 


The curve is a 


little farther from ]the tip than the most elongated zone, just 


1901] GEOTROPISM OF STEMS 421 


as occurs in roots when the experiment lasts more than a 


few hours. 
| 
| January 16 | January 17, 9 p.m, 
| Length | Growth | ges piensa Curve 
I yym™m 3mm 3 | 4 go° 
2 12 7 3-4 3 75 
3 6 4 2 3 90 
4 II 7 2-3 3-4 80 
5 14 z 3 3-4 go 
6 14 6 3 4 go 
7 19 E) 3 4 95 
8 7 5 2 3 70 
9 2+ 3 2 a+ 50 
10 15 5 3 4 70 
It 9 ‘i 2-3 4 990 
12 9 I I 2-3 tea 
13 5 3 4 a2 go 
4 4 3 2 3 85 
15 10 6 3 4 bezier => 


The root of Yucca begins to grow earlier than that of Phoenix, 
and a considerable part of the growth of the cotyledon takes 
place afterward in the development of the elbow from whose 
‘inside the plumule breaks, as shown in fig. 3. Yucca is more 
like Allium than is Phoenix, but in neither does the cotyledon 
develop visible chlorophyll. Experiments with a considerable 
number of Yucca plants were perfectly harmonious during the 
first growth, and it will suffice to report on three typical plants. 
The first two seedlings were marked and placed horizontal 
November 9, 5:00 P.M. 

I.. Lengtho™", of whichrootis4.5™™. November 10, 11:00 A.M.: growth 
in zones 1-3, 2.5", mostly in zone 2; curve 85° in zone 2; in zone 6-9 growth 
0.8™™ without curve. November 12,11 :00A.M.: growth in zones 6-9 is 1.4™™, 
still straight. 

2. Length 5 ™, of which root is 2™™. November Io, II :00 A.M.: no appre- 
ciable change. November 11, 12:00 A.M.: growth in zone 1, 0.4™™; curve 
15°; trace of growth throughout; growth above zone 5, 2.0™™ without curve. 
November 12, 11:00 A.M.: growth'r ™™ in zones I-3 ; curve 40° ; growth above 
5, 2.8™™, still straight. 

3. Began November 22, 9:00 A.M. Length 3™", of which root is 1™™. 


a ce sna aa 


Fic. 3.— Older 
seedling of Yucca 


angustifolia. 


BOTANICAL GAZETTE [JUNE 


November 23, 3:00 P.M.: growth 6™", mostly in zone 1; 
but curve of 90° mostly in zone 3, doubtless made before 
most of the growth in the root. 

In Yucca, as in Phoenix, the downward curve 
takes place in root or cotyledon, in whichever the 
most rapidly elongating part of the apical growing 
region falls. But the elongation of the more 
remote, parts of the cotyledon is not at first 
accompanied by any curve at all. Afterward, in 
the formation of the elbow, a combination of 
curves develops. These are in part probably 
geotropic, but are in part not under the direct 
influence of gravity, being executed in the develop- 
ment of structures whose initial position gravity 
determines. For the same reason as in the case 
of Cucurbita, a further treatment of these later 


curves is unnecessary here. 


The common statement is literally true, that 
these cotyledons are positively geotropic. But 
the meaning that has been conveyed by the words 
is only half correct. They do curve downward, 
but cannot perceive the gravity stimulus in a 
way that would make them curve so. As in the 
hypocotyl of Lupinus, any positive geotropism 
they may exhibit is under the control of the 
punctum vegetationis of the root, which alone is 
positively geoesthetic. 


WEST VIRGINIA UNIVERSITY. 
Morgantown, W. Va. 


ae 


BRIEFER AR Tig, 


NOTES OF TRAVEL. VI." 
THE BOTANICAL INSTITUTE OF NETHERLANDS INDIA. 

THE gardens of Buitenzorg, Java, have been described so often that 
their name should by this time be familiar to every American botanist, 
The institution has grown so rapidly in the last few years, however, that 
a short description of the improvements made may be of interest to 
any one who expects to visit Java, and to all who are interested in the 
development of tropical botany. Although the Institute bears the name 
“botanical,” it has many decided agricultural features, and on account 
of its various divisions and the nature of the subjects under investiga- 
tion would be called by many Americans a most unusually well 
equipped station. 

The impression made upon the writer in 1896, during an eight 
months stay in these gardens, was that they offered unrivaled oppor- 
tunities for study, a wealth of interesting material, and surroundings 
full of the most interesting oriental sights. After three years’ absence, 
much of which time has been spent in a study of other botanical gar- 
dens and institutions, I find that Buitenzorg still possesses the charm 
and offers even greater opportunities than in 1896. During the last three 
years, under Dr. Treub’s excellent management, five new buildings have 
been erected. They consist of a double laboratory in which tobacco and 
coffee are investigated, a very comfortable pharmaceutical laboratory, 
a new library building which now holds the most complete collection of 
botanical works in the tropics, a luxuriously appointed office building, 
and a very pretty tropical bungalo for Dr. Treub. The censtruction of 
the laboratories is very substantial. They are all one story buildings 
with concrete floors, projecting tile roofs, and numerous large windows 
which in this latitude let in a flood of light and require white shades. 

Tables, desks, cases, and shelves are kept scrupulously clean and neat, 
and in this regard are in strong contrast with most tropical institutes, 


in which a musty smell pervades everything. Dr. Treub has given in 


the first number of his Bulletin de l’ Institut Botanique de Buttenzorg, 

published in December of 1898, an excellent sketch of the gardens and 
*No, 5 of this series was lost in transit.—ED. 

Igor] 423 


424 BOTANICAL GAZETTE [JUNE 


their personnel. He has included also a description of the publications 
of the Institute, an estimate of the necessary expenses of a stay at the 
gardens, and the cost of the sea voyage from Europe. This first num- 
ber of the Bud/etin will be sent to any botanist who writes for it with 
the idea of preparing for a stay at the gardens. 

The new quarter, to which Mr. Wigman, the head gardener, had 
transferred nearly all of the climbing plants, was in 1896, like all such 
newly planted ground, unsatisfactory to look at. It has now grown 
until it is an attractive portion of the gardens, and the new avenue of 
canary trees in it, which was planted to rival the old avenue (for which 
the gardens are famous in the eyes of travelers), is already very hand- 
some, with its regularly rounded tree tops and light gray trunks. A 
water-garden which had just been laid out in 1896 is now indistin- 
guishable from older parts of the garden, and the fern quarter and collec- 
tion of Pandanus, of which photographs have been so often published, 
have grown more interesting with their coatings of epiphytic algae. 
The small nursery, which was large enough three years ago to repro- 
duce all the plants needed in the gardens, has been more than trebled 
in size, and packages of seeds and cases of plants are being sent all 
over the archipelago (2294 packages and cases in 1897), as well as in 
exchange with all the principal botanic gardens of the world. 

Native labor is being utilized in a new printing office in the gardens, 
where all small forms and even scientific periodicals are printed, the 
compositors being Javanese who do not understand a word of what they 
set up. The work is done very slowly, and the proof reader’s patience 
is taxed to the utmost, but because of the low price of labor and the 
inconvenience of having the work done in Batavia or Amsterdam is 
very great, the office is a great convenience. 

I had the pleasure of accompanying Dr. Treub and Mr. Wigman in 
one of their early morning strolls, in the course of which they bar- 
gained with two neatly dressed Javanese land owners for some paddy 
fields of which to construct experimental plats. In response to Dr. 
Treub’s and Mr. Wigman’s inquiries the Javanese replied with respect- 
ful salaams and remained sitting on their heels as they would before a 
raja. These new plats which were purchased are to be under the 
supervision of a newly appointed specialist, whose acquaintance with 
Javanese vegetables and other native food plants will enable him to 
select and improve them, and to distribute information among the 
natives regarding the best methods of their culture. 


=x 


———— 


ae es eee ee 


1901 | BRIEFER ARTICLES 425 


The new laboratory for pharmaceutical research is a model of com- 
pactness and convenience. The library building, which was the gener- 
ous gift of Mr. Janse, of Amsterdam, has now not only the increased 
library of the gardens but the considerable collections of books for- 
merly belonging to the scientific society of Batavia. The removal of 
the library from the herbarium building gives Dr. Boerlage greater 
space for his rapidly growing collection, and the old building, for- 
merly occupied as the pharmaceutical laboratory, is to be utilized for 
an exhibition of economic plant products. 

A lively interest in the fruits of the island has been awakened 
among the planters, and a horticultural society with more than 300 
members has been formed. The first exhibition of fruits, which was 
held in December last, was a great success, and the garden authorities 
hope through cooperation with members of the society to secure a col- 
lection of the best varieties of fruits, and by distributing grafts from 
these, to replace the inferior seedling kinds, which now furnish the 
fruit for the tables of the Europeans. A seedless doekoe (Zansium 
domesticum) has already been found, and other superior varieties are 
known to exist in the island. Almost everywhere in the tropics fruit 
trees are wild, and it is one of the curious observations which a traveler 
makes that little is done to improve fruits which are evidently capa- 
ble of very great improvement. There are mangosteens which without 
selection are nearly or quite seedless, and yet Europeans choose to 
plant seeds instead of grafts, and still have a strange fear that a 
grafted tree will be a short-lived sickly thing and not repay for the 
extra trouble taken with it. The new horticultural society should do 
much to enlighten the planters and enable them to plant and breed 
better fruits, even in a country noted for its delicious pineapples and 
incomparable mangosteens. 

There are doubtless many other lines along which the gardens at 
Buitenzorg have improved. The selection of sugar cane seedlings, the 
hybridizing of coffee, the establishment of a new zoological museum to 
be under the management of Dr. Konigsberger and to contain mounted 
specimens of all the many interesting animals and cases of the injurious 
and biologically interesting insects of the archipelago, were all propo- 
sitions under consideration at the time of my short visit in January. 
I am thoroughly convinced that to any one who expects to make a 
thorough study of tropical plants a visit to this Botanical Institute will 
be of the greatest advantage. Its opportunities surpass those of any 
other in the world.— Davin G. FaircHILD, Department of Agriculture. 


420 BOTANICAL GAZETTE [JUNE 


THE CARDINAL PRINCIPLES OF MORPHOLOGY.’ 


ALTHOUGH botany has made remarkable advances in America 
during the past few years, there is still one phase in which it remains 
singularly backward, namely, in its treatment of the morphology of 
the higher plants. There still prevails among us, with little modifica- 
tion, the old formal idealistic morphology, whose founder was Goethe, 
and whose great exponent in this country was Gray; while we give 
scant consideration to the newer natural realistic system, now more or 
less fully accepted elsewhere, and recently given greater extension by 
its leading advocate, Goebel. We have, it is true, some literature of 
the newer morphology, of which an example is Professor Barnes’ dis- 
cussion of the Flower in the Cyclopedia of American Horticulture, while 
the treatment of the homologies of the higher with the lower plants is 
good in most of our recent text-books; but from these there is every 
gradation backward. Happily the newer standpoint is becoming gen- 
erally accessible to American students through the publication of 
Goebel’s Organographie der Pflanzen (Jena, Fischer, 1898-1900), now 
being translated into English under the title Organography of Plants 
(Part I, Oxford, Clarendon Press, 1900). In the present paper I 
propose to summarize what seem to me the principles upon which 
the newer morphology is based. 

The difference between the idealistic and the realistic morphology, 
while partly one of fact, is mainly one of point of view. The idealistic 
system is based principally upon comparative anatomy ; it concentrates 
attention upon the steps, or stages, in morphological changes, or meta- 
morphoses, but is largely indifferent as to the processes, or mechanics, 
by which the metamorphoses have been brought about ; metamorphosis 
is therefore to it chiefly a phylogenetic operation, whose exact ontoge- 
netic basis is of secondary consequence. The realistic system, while 
giving great weight to comparative anatomy, lays especial emphasis 
upon the testimony of embryology, particularly seeking the actual onto- 
genetic origin and development, the mechanics, of metamorphoses, only 
through which, it maintains, can. the true nature of metamorphosis be 
understood ; metamorphosis is, therefore, to it primarily an ontogenetic 
process which later and secondarily becomes fixed in the phylogeny. 


The former, the idealistic or phylogenetic systein, predisposes one to _ 


generalized and abstract conceptions, while the latter, the realistic or 


? Read before the Society for Plant Morphology and Physiology, Johns Hopkins 
meeting, December 28, 1900. 


- 


1gor | BRIEFER ARTICLES 427 


ontogenetic, leads to more definite and concrete conclusions. Both 
views equally assume the fact of evolution in the phylogenetic fixation 
of metamorphoses, but both are equally independent of the exact 
method (the dynamics) by which that evolution is brought about, 
whether this be through natural selection involving the whole organ- 
ism, or through germinal selection, or through organic selection, or 
through the accumulation of transmitted effects of individual irritable 
responses, or through some other method still unknown. The realistic 
system, however, brings us more nearly face to face with the problems 
of the dynamics of evolution than does the idealistic system. 

We pass next to an attempt to deduce the fundamental or cardi- 
nal principles of morphology. Of these, some are not peculiar to 
morphology alone, but belong equally to other phases of evolution 
and adaptation, while others are especially characteristic of mor- 
phology. 

First, the eeible ¢ of continuity of origin, that is, no functional 
Structure ever arises de novo, but only from the modification of a pre- 
existing structure, which in turn arose from a still earlier, and so on 
backward through a longer or shorter chain ending only in the original 
protoplasmic variation, or in whatever it is which does lie at the begin- 
ning of specialization. This aia is axiomatic for both systems 
of morphology. 

SECOND, the principle of spiortiaia that is, the direction taken 
in metamorphosis is not determined by obedience to any pre-formed 
plan, but, except for the influence of the inertia of the heredity of the 
particular part, follows the factors potent at the moment. Heredity of 
itself cannot impose any plan, for it is but the summation of the inter- 
action of past experiences with original properties. ‘Theoretically this 
Principle should be as acceptable to the idealistic as to the realistic 
‘morphology. In practice, however, the idealistic conception of meta- 
morphosis as a whole is that of a play of a very few highly plastic 
“members,” which, however much they may vary and combine, retain 
a sort of fundamental immutability of nature, as witness the efforts to 
explain all parts of the flower in terms of “leaves” and “stem,” and 
the use of the expression “disguised” often applied to metamorphosed 
parts. There is thus imposed upon metamorphosis a sort of ideal plan, 
aplan implying that modification keeps within certain limits, deter- 
mined by the possibilities of permutations and combinations of those 
members. The realistic morphology is bound by no such conception, 


428 BOTANICAL GAZETTE [JUNE 


for the members are unlimited in number and have no ideal nature or 
limitations to be consulted. 

THIRD, the principal of fwsctional domination in metamorphosis, 
that is, it is function which takes the lead and structure follows. It 
is of course true that function and structure are reciprocally related ; 
there are cases where structure determines function; there are other 
cases in which non-functional factors, especially an aggregation of 
them, may outweigh a functional factor ; ‘nevertheless, it must be true 
that in a broad way it is function which determines structure, function 
often hampered, and even sometimes thwarted by other influences it is 
true, but function dominant in the long run. If this is not true, 
adaptation is but an accident if not a myth, and our whole idea of it 
but a vain vaporing of the imagination. This principle in general is 
necessary to both systems of morphology, but it is more prominent in 
realistic than in idealistic discussions. 

Fourth, the principle of indeterminate anatomical plasticity, that is, 
in all anatomical characters (size, shape, number, position, color, cellu- 
lar texture) plant-organs, or, if one pleases, plant-members, are not 
limited by anything in their morphological nature, but, under proper 
influence, may be led to wax and wane indefinitely in any of these 
respects. Of course this plasticity is hampered by innumerable prac- 
tical considerations, and by many hereditary “tendencies,” and many 
generations may be (though they are by no means always) necessary 
to produce a marked and permanent result; but the point is that lim- 
itations to anatomical plasticity do not come from the morphological 
nature of the part concerned. The principle may also be stated thus, 
that there is no causal relationship whatever between morphological 
nature and anatomical structure; stipules are usually smaller than the 
leaves they accompany and markedly different in form; yet they may 
become as large as the leaves and indistinguishable from them (as in 
Galium), or much larger, as in some Leguminosae ; there is obviously 
therefore nothing in the nature of a stipule as such to limit its size or 
shape; that it is usually smaller than its leaf is a purely functional and 
non-morphological matter. So, stems may lengthen immensely as in 
climbers or shorten to apparent disappearance as in rosette-plants, and 
so on with all other parts. This principle is equally true for both 
systems of morphology, and is perhaps the most clearly recognized of 
them all. 

Firtu, the principle of metamorphosis along lines of least resistance, 


Igor] BRIEFER ARTICLES 429 


closely related to opportunism mentioned above. This means that 
when, through a change in some condition of the environment, the 
necessity arises for the performance of a new function, it will be 
assumed by the part which happens at the moment to be most avail- 
able for that purpose, regardless of its morphological nature, either 
because that part happens to have already a structure most nearly 
answering to the demands of the new function, or because it happens 
to be set free from its former function by change of habit, or because of 
some other non-morphological reason. It is due to the operation of 
this principle that structures of the most different morphological origin 
may come to serve the same function, and correlatively, structures of 
the same morphological origin may come to serve the most different 
uses. The genus Pereskia, in the Cactaceae, includes mesophytic 
climbers with true broad leaves, from which all gradations in reduction 
of leaves and condensation of stem may be traced even to the typical 
desert forms of Cereus. Now, one division of Cereus returns to a 
life in the woods, where the demand for an increase of green surface 
is felt; no attempt, however, is made to restore the old leaves (now 
reduced to tiny scales), but the stem enlarges and branches, while the 
vertical ribs, developed during the desert habit, are expanded farther- 
and made to function as leaves. As the mesophytic habit becomes 
more extreme, the ribs become larger in size and fewer fn number 
until finally, in Phyllocactus, but two remain, and these become so 
flattened and arranged in such a manner on the branch that they form 
a physiological and anatomical leaf. Here we have a case, indubitable 
because abundant intermediate steps persist, in which a physiological 
leaf has been developed from a morphological stem, purely by follow- 
ing the line of easiest accomplishment, or least resistance, at the 
moment; no single step is in itself remarkable, but the sum total 
yields a very remarkable result. This principle is, of course, equally 
applicable to both systems of morphology. 

SIXTH, the principle of metamorphosis by transformation, as con- 
trasted with metamorphosis by differentiation, which means that when, 
in response to any influence, a new function and hence structure 
(function-structure) is assumed by any part, this always comes about, 
both ontogenetically and phylogenetically, through the transformation 
or alteration of a previously existing function-structure in that part, 
and never through the differentiation of a new function-structure out 
_ Of a previously functionally-indifferent or unspecialized structure. In 


430 BOTANICAL GAZETTE | JUNE 


other words, since all parts of the plant actually are organs (that is 
have some meaning in the life of the plant), new organs can arise 
only by the transformation of previously existing ones. We face here a 
sharp contrast in the two systems of morphology. ‘The idealistic sys- 
tem, laying as it does great stress upon members in distinction from 
organs, comes to regard these members as if they had a real existence, 
forgetting that the conception of the member is a pure abstraction of 
the mind, a sort of mental composite photograph, with no objective 
equivalent, and that members apart from organs do not really exist. 
The conception of the member as an entity having once been formed, 
metamorphosis is naturally regarded as the differentiation of an organ 
out of a member, and this not only phylogenetically but also onto- 
genetically; so that by those who carry the idealistic system into 
ontogeny at all, the ontogenetic unfolding of any organ is viewed asa 
differentiation from primordia (Anlagen) which, after the analogy of 
the members, are supposed to be indifferent in their nature. Not only, 
however, is this view untenable upon philosophical grounds, but it is 


negatived by the fact that in cases where metamorphoses are experi-_ 


mentally brought about, embryology shows that the process is actually 
one of transformation of one function-structure into another, and not 
of differentiation of a function-structure out of a neutral or indifferent 


primordiui (Anlage). But this subject is so clearly treated by Goebel _ 


in the Introduction to his Organography that it needs no further con- 
sideration here. How these, in their origin purely ontogenetic, meta- 
morphoses become fixed in the phylogeny, is an entirely separate 
question, the solution’ or non-solution of which does not in the least 
affect the truth or non-truth of this principle. The idealistic concep: 
tion, that an organ is formed by differentiation from a member, 
implies as a corollary that each organ is but one step, so to speak, 
from a member, and should be readily reducible to it; hence arise the 
attempts to explain all parts of such complex and specialized structures 
as epigynous flowers in terms of leaf and stem, necessitating the 
adnate calyx theory with its requirement of extraordinary assumptions 
as to growing together of parts, etc., entirely unsupported by the facts 
of development. 
SEVENTH, the principle of gradation in morphological membership, 


involving the existence of degrees of morphological independence, 


culminating in the attainment of full morphological membership with 
full independence. Or, it may be expressed thus: in the progressive 


901] BRIEFER ARTICLES 431 


development of metamorphoses, difference of degree passes over 
gradually into difference of kind. This principle, in my opinion 
the most fundamental in morphology, marks far more sharply than 
any other the difference between the two systems, for it is fundamental 
to the realistic, but inconsistent with the idealistic conception. It 
means that, as an organ gradually acquires a new function-structure, 
and the old function-structure is gradually lost, new powers of varia- 
tion, adaptation, etc., are acquired which become more and more inde- 
pendent of those formerly possessed by the organ, until finally the 
change may become so complete that the new organ not only acts 
itself quite independently of its old nature, but becomes a new start- 
ing point or center of metamorphosis, that is, it becomes a new mor- 
phological member. New metamorphoses, however, are not confined 
for their starting-point to the full members, but may originate from 
any of the points along the lines of gradation. Hence, not only may 
any Organ become a member, but the members grade into one another 
indefinitely and any of the gradations may act as members. This is 
in great contrast with the conception of the idealistic system as applied 
to the higher plants, for that conception not only limits the number 
of the members to a very few (at the extreme root, stem, leaf, plant 
hair, exclusive of the sporangia), but practically views these as sharply 
distinct, not recognizing intermediate transition from which new meta- 
morphoses may originate. In fact the idealistic morphology, while 
admitting the original evolutionary origin of its members, ignores 
evolution in their subsequent interrelationships ; it views its members 
much as species were viewed in pre-evolutionary times, while the real- 
istic system applies the idea of evolution throughout. The idealistic 
System views the morphological members much as a chemist does his 
elements, which may combine in many ways, but retain their identity 
throughout; the realistic system regards them more as a physicist does 
the colors of a spectrum, as a series of stages in a graded sequence of 
phenomena. 

As an example of an organ which has attained to full morphologi- 
cal membership and independence we may consider the spines in the 
Cactaceae. Research has shown much evidence for the belief that these 
structures have arisen by the metamorphosis of leaves; the only com- 
peting theory is that they are a form of ‘‘emergences;” no investiga- 
tor has ever seriously supposed they were anything else. Now, despite 
much long-continued observation and special search, no one has ever 


432 BOTANICAL GAZETTE [JUNE 


been able to find transitions between these spines and either leaves or 
emergences (except in the case of a few monstrosities which may have 
another meaning), nor have repeated experiments succeeded in making 
the spines return to leaves or emergences. Now, the cactus spines are 
immensely variable, becoming very big and hard on the one hand, or 
weak and small even to disappearance on the other, cylindrical and 
erect to ribbon-like and contorted, plain or variously ornamented, 
smooth or beautifully plumed or fringed, curved into hooks useful for 
climbing, or altered entirely into nectaries. But, throughout all of 
these variations it is distinctly and unquestionably a spine, an anatomi- 
cal spine that is varying, and not a disguised leaf or emergence. We 
must conclude from all these grounds that the cactus spine has attained 
to full morphological membership, is itself a member, a center of modi- 
fication and metamorphosis. The mamillae, or tubercles, in the same 
family, originate by a union of the leaf-base and its axillary bud, but 
the identity of these two parts becomes completely lost in the new 
identity of the tubercle, which becomes a member and acts as such 
through many genera. The ribs in Cactaceae arise by the running 
together of vertical lines of tubercles; once formed, however, they pay 
little attention to their mode of origin but proceed to act as independ- 
ent members, as one may clearly see when he considers their perfor- 
mances (particularly their independent increase or reduction in 
number) in the development of the cladophylls of Phyllocactus 
already cited in this paper. In some genera, however, particularly 
Echinocactus, the ribs have not attained to full independence, for they 
occasionally revert to lines of tubercles. But we need not go so far afield 
for our illustrations of the attainment of independent membership, for 
the members commonly accepted by the idealistic system (root, stem, 
and leaf) illustrate it perfectly. Most of us no doubt believe that the 
present-day foliage leaf and stem arose through the sterilization of 
sporogenous tissue in a primitive very simple sporophyte ; but whether 
we believe it or not does not matter for our present purpose, for we must 
believe, if we accept evolution at all, that leaf and stem have become 
specialized out of a simpler structure which did not show those dis- 
tinct parts? All morphologists accept the foliage leaf and its stem as 
of full morphological membership and independence, to such a degree 
indeed that they stand in most minds as the very types of morphologi- 
cal members. Now, in their case, even the idealists never attempt to 
interpret their morphological behavior in the light of the nature they 


Igor] BRIEFER ARTICLES 433 


had before they were leaf and stem. Why, then, does the idealistic 
morphology insist, for example, upon reducing everything (excepting 
the sporangia) in a highly specialized flower to the categories of leaf 
and stem? In fact, the flower has been so long an independent organ 
that it has had time to progress far toward independent morphological 
membership, as witness its ability to suppress circles, to alter the num- 
ber of their parts, and to rearrange their phyllotaxy quite independently 
of any actions performed by leaves on a stem. Moreover, various 
parts of the flower (in some flowers, not in all) have become more or 
less independent members, as we may clearly see in those which are 
epigynous. The ovary of such a flower, for example, unquestionably 
originated in sporophyllary leaves standing upon a conical receptacle, 
precisely as in numberless flowers today; gradually, however, as 
embryology proves, the formation of the ovarian cavity was given up 
by the carpels, and assumed by the receptacle, which grew up in the 
form of a cup carrying the other parts upon its rim, while the carpels 
finally came to form simply a roof over the cavity. But, and here is a 
crucial point, it must not be supposed that during this process the 
receptacle and carpels retained their old carpel and receptacle nature 
(much less their “stem” and their “leaf”? nature); on the contrary, the 
new kind of ovary acquired an identity and a character of its own, and 
in that new identity and character the old identity and character of 
receptacle and carpel gradually melted away, and lost their distinctness, 
so that such an ovary has become a new member and is not simply a 
compound of receptacle and carpel. It is useless, therefore, to expect 
that such an ovary will build placentae, partitions, style, or stigma 
according to the rules in vogue with ordinary receptacle and carpel, 
and useless also to discuss whether in such an ovary the cavity is lined 
with carpel or not, for the ovarian wall is no longer either receptacle 
or carpel or both, it is ovarian wall; carpel and receptacle have not 
fused to form it; their tissue has melted away, so to speak; into the tis- 
sue which does form it. For simplicity I here treat this attainment 
of membership by such an ovary as if it were complete, though in fact 
it is not so in any ovary known to me, for in all of them some features 
of both carpel and receptacle may be traced, especially at top and bot- 
tom of the ovary. So also, with other parts of the flower; the placenta, 
which originated in the manner still shown by many flowers, as swollen 
edges of carpels, has become independent of its carpellary origin in 
many flowers, as in those with free central placenta, where no trace of 


434 BOTANICAL GAZETTE [JUNE 


the old leaf nature may be found in its entire ontogeny. The tube of 
gamopetalous flowers is another structure which in some cases has 
attained to approximate independence, and the style and stigma yet 
another. All of these parts may become centers of independent 
(though usually limited) metamorphosis, altering in size, shape, num- 
ber, building new structures (as the corona) all independently of any-: 
thing they did before they acquired their present more or less 
completely independent membership. It is so with the ovule, orig!- 
nating in a macrosporangium, but now an independent member. The 
same is true of the embryo-sac, which, originating unquestionably as a 
germinated macrospore, has, after a long history, become a new member 
with a high, though not acomplete independence. Some of its features. 
are to be explained as a persistence of its ancient nature, but it is use- 
less to attempt to interpret all its actions upon that basis, for it does 
some new things upon its own account—as a new member, the 
embryo-sac. To return for a moment to the flower as a whole, it is 
important to notice that the study of its morphology is in one sense 
complex and difficult, partly because it is a composite structure with 
various degrees of independence in its component parts, partly because 
its development in different families has been so independent that it 
has given rise to homoblastic rather than homogenetic homologies. 
Hence it is impossible to make distinct categories of members apply- 
ing to all flowers, but each group must be considered by itself, a feature 
indeed which applies not to the flower alone but to the vegetative parts 
as well. Hence we must in theory recognize as potential members all 
organs, one may even say all recognizable parts, although in practice 
it is needful to take account for the most part only of those most con- 
spicuous and distinct. 

The realistic system, with its infinite gradations and limitless pos- 
sibilities, is much harder to grasp and to apply, and is less pleasing to 
teach than ‘the idealistic system, with its few distinct categories and 
their involved limitations. But it is truer to nature, more stimulating 
to research, and more replete with promise of great results.— W. F. 
GANONG, Smith College, Northampton, Mass. 


— 


CURRENT LER RA PORE. 
BOOK REVIEWS: 
A Californian manual.* 


THIS manual by Professor Jepson deserves more than a passing mention, 
for in a certain sense it is constructed on new lines. The average manual is 
chiefly a compilation of scattered literature checked by more or less herba- 
rium study. The result is merely an approximation to the facts and never 
quite satisfactory to the field student. Professor Jepson has met his plants in 
the field and has described them as they live. He has recognized literature 
so far as it fits his material, but has not allowed it to bias or handicap him. 
As a consequence, the descriptions are kably fresh and telling, and have 
no flavor of stereotyped diagnoses. Not only is the wonderfully diverse 
Californian flora set forth, but numerous ecological notes suggest the factors 
that lie behind the diversities. Repeated attention is called to the variations 
in vegetative characters which a single species may undergo in different situ- 
ations. The following statements from the preface are worthy of quotation, 
since they are of general application : 


Near the ocean a species is often more depressed or condensed than in the 
an and more fleshy. 

mps or wet soils the plant tends to become succulent and of ranker 
ek a ora gla 

3. In valley soils ae ore is commonly much more rank than elsewhere. 

4. On hilltops plants tend to become dwarf and acaulescent; often far more 
pubescent also. 

5. In saline or subsaline soil the stems and foliage in many species are far more 
vigorous and the flowers larger than on stiff clays or adobes. 
In the shady woods leaves become thinner and larger, often conspicuously so. 

7. At high altitudes the flowers are larger in proportion to stature and brighter 
in color. 

Such facts are known to the ecologist, but it seems hard to get the tax- 
onomist to give them due weight. . The region covered lies west of the Sacra- 
mento and San Joaquin rivers, south of the counties of Mendocino and 
Colusa, and north of the Pajare river and Pacheo pass. Several such manuals 
will be needed to present the flora of California, and it is daily becoming 
more evident that no one is competent to prepare them who has not lived 
among the plants. The numerous analytical keys are prepared with special 


*JEPSON, WILLIS LiNN: A flora of western middle California. 8vo. pp. iv-+- 
625. Berkeley, California: Encina Publishing Company. April 16, 1901. $2.50. 
1901] 435 


436 BOTANICAL GAZETTE [JUNE 


care, and the verbiage of pedantry has been eliminated. A number of new 
species are described, but from the statements made the field seems to be 
white for the harvest. Professor Jepson is to be congratulated upon pro- 
ducing a useful, rational, and modern manual.— Cc 


The Cyclopedia of American Horticulture. 


The third volume of Bailey’s Cyclopedia of American Hortieulture has 
just come from the press of the Macmillan Company. It includes the letters 
N-Q.?_ Aside from the articles on important genera, among which the most 
noteworthy are those on Nepenthes, Nymphaea, Odontoglossum, Oncidium, 
Opuntia, Pinus, Prunus, Pyrus, and Quercus, the following articles are of 
special interest. The botanical treatment of Orchids is by H. Hasselbring, 
the culture of orchids by Robert M. Grey, with a general introduction by the 
editors. On the Peach, the general article is written by Professor Bailey, 
peach culture in the south is treated by J. H. Hale, peach culture in the far 
north (northern Vermont) by J. T. Macomber, the Michigan peach industry by 
R. Morrill, peach culture in Delaware by Charles Wright, the peach in California 
by H. Culbertson. Professors Bailey and T. T. Lyon contribute a general 
article on Pear, C. L. Watrous writes about pears on the northern plains, E. 
J. Wickson on the pear in California, John S. Collins on the Kieffer pear, 
L,. T. Yeomans on dwarf pears, and George T. Powell on pears for export. 
The horticultural capabilities of the PAz/ippines are described by F. E. 
Gannett. A somewhat unexpected article is the one on Horticultural 
photography by J. Horace McFarland. It ought to be very helpful and to 
result in the great improvement of the illustrations in our experiment station 
reports and other similar publications. A compact article upon the general 
Physiology of plants is contributed by Dr. B. M. Duggar. The article on 
Plant breeding by Mr. H. J. Webber will attract attention. The principles of 
the art are well set forth, but it is unfortunate that he unnecessarily insists on 
calling stamens and pistils the male and female organs of plants. On the 
Plum Professor Bailey writes the general article, Professor F. A. Waugh 
discusses the cultivation of native plums, and Mr. A. V. Stubenrauch gives an 
account of the plum in California. The article on Pomo/ogy is naturally 
handled by the editor, an appendix on Russian fruits being contributed by 
John Craig.- The work maintains the high standard of the preceding volumes.’ 

R 


* BaILEy, L. H.: Cyclopedia of American Horticulture, comprising suggestions 
for cultivation of horticultural plants, descriptions of the species of fruits, vegetables, 
flowers, and ornamental plants sold in the United States ~ peprinay together with 
geographical aut Seiecapeial sketches. Vol. III. N-Q. » PP- ee 
Sigs. 1454-2059, pls. 20-29. New York: The Macmillan gies 1900. $5- 

3 Bot. Gaz.2g :282; 30:277. 1900. 


{901 ] CURRENT LITERATURE 437 


MINOR NOTICES. 

Dr. G. N. BEstT has revised the North American species of Heterocladium 4 
recognizing four species, of which two, 1. acount and 7. heteropterioides are 
new. Three Kindbergian species are in the ‘doubtful or excluded” list. 
—C.R. B. 


Emma J. COLES has published a catalogue of the native vascular plants 
growing in the vicinity of Grand Rapids, Michigan. The ntroduction pre- 
sents the general physiographic and ecologic features of the region, and in 


the list all known stations are carefully recorded. The number of native 


plants recorded is 1111, of introduced plants 176.—J. M. C. 

V. K. CHEsnut and E. V. WILcox have published a valuable preliminary 
report (Bull. 26, Div. of Bot., U.S. Dept. Agric.) of their investigation of the 
stock-poisoning plants of Montana. It contains not only a detailed account 
of their observations and experiments, but also summaries of symptoms 
and remedies. The most important plants poisonous to stock in Montana are 
the death camas (Zygadenus venenosus), tall larkspur(Delphinium glaucum), 
purple larkspur (D. dzcolor), Wyoming water hemlock (Cicuta occidentalis), white 
loco weed (Aragallus spicatus), and various lupines (Lufinus spp.).—J. M. C. 

THE FIRST FASCICLE of what promises to be a very excellent treatise on 
botany by Bonnier and DuSablon has just been published.© The work is 
intended specially for advanced students in the universities and schools of 
medicine, pharmacy and agriculture. The first section discusses such general 
matters as the characters of plants, the members of the plant body, the large 
divisions of plants, the cell and the tissues. The second section, entitled the 
morphology of angiosperms, concerns itself with the primary and secondary 
Structure of stamens, leaves, and roots. The illustrations are fresh and in the 
main excellent, though the anatomical ones are better than those represent- 
ing external features. We reserve further notice of this work until its 
completion.—C. R. B 


WILLIAM R. Maxon’ has published a very useful list of the pteridophytes 
of North America north of Mexico. Eaton’s Ferns of North America and 
4Bull. Torr. Bot. Club 28 : 123-131. pls. 77, 74. 1901. 
rand Rapids flora, a catalogue of the flowering plants and ferns growing with- 
out pe eens in the vicinity of Grand Rapids, Michigan. 8vo. pp. xxii-+ 170. 
Grand care Lyon, Kymer & Palmer Company. I90I. $1.00. * 
NNIER, GASTON, and DuSABLON, LECLERC: Cours de botanique ; anatomie ; 
physiologic ; ‘daasiicaia: applications agricoles, industrielles, médicales ; morpholo- 
le expérimentale ; géographie botanique ; art éontologie ; historique + a usage des 
éléves des universités, des écoles de médecin et de pharmacie, et des écoles d’agri- 
culture. Vol. I, part 1. 8vo, pp. 384, figs. oe Paris: Paul Dupont. 1901. Com- 
plete in 6 parts. 25 fr. Each part singly 6 
7A list of the ferns and fern allies of North America north of eee wae prin- 
cipal synonyms and distribution. Proc. U. S. Nat. Museum 23 : 619-651. 


438 BOTANICAL GAZETTE [JUNE 


Underwood's Our native ferns and their allies have been the only works of suffi- 
cient scope to include this territory. The former is more than twenty years old, 
and the latter is essentially a popular treatise in which there are no citations, 
As a consequence, a list of the known ferns and their synonymy has been a 
desideratum for some time, and this Mr. Maxon has supplied. Besides his 
careful bibliographic work, the author has included the recorded geographic 
range of each species, thus adding very materially to the value of the list. 
What may be called the Underwood nomenclature and sequence are fol- 
lowed.—J. M. C 

A LABORATORY MANUAL has recently been published by F. E. Clements 
and I. S. Cutter. It is of special interest as being “(an authoritative expres- 
sion from the Department of Botany of the University of Nebraska upon the 
kind and amount of elementary botany that should be taught in the accredited 
schools and colleges of the state.” The directions for work are clear and 
compact, and based upon long experience in handling the material. Granted 
that such material is best suited to high-school courses, no exception can be 
taken to the way in which it is presented. A question might be raised, how- 
ever, as to the ‘‘kind of elementary botany” that this book calls for. To start 
high-school pupils with a short course in histology is probably not commended 
now as much as formeyly. The part devoted to structure and classi- 
fication would seem to be a more fitting introduction to the use of plant 
material. A good set of physiological experiments is also included, and it is 
interesting to note that a certain amount of work in ecology is called for. The 
book must be of great service to the schools of Nebraska in so far as it relates 
them to the work of the University.—J. M. C 

THE PROCEEDINGS of the twenty-first meeting of the Society for Promo- 
tion of Agricultural Science, recently distributed, form a volume of 183 pages, 
containing articles of botanical interest. The president's chair was occu- 
pied by a botanist, W. J. Beal, of Michigan, but the annual address dealt 
chiefly with matters of general interest to the society. A ‘syllabus for a 
short course on grasses and other forage plants” by the same person is 0 
decided pedagogical value. “The development of a tomato hybrid” by W. 
M. Munson is an account of the production of a desirable hybrid variety by 
crossing the common and the currant tomato. “The chemical function of cer- 
tain soil bacteria’? by Frederick D. Chester, “Seven years of experiments 
with bush beans” by Byron D. Halsted, ‘‘ The value of willows in retaining 
the banks of streams” by W. W. Rowlee, “ The course in cryptogamic 
botany’’ by L. H. Pammel, “The weedy plants of Iowa,” also by Mr. Pam- 
mel, contain botanical matter with practical applications of the facts. H. L. 
= corrects some errors of microscopic interpretation occurring in his 

A laboratory manual of high school botany. 8vo. pp- 123. Lincoin, Nebraska: 

The Peron Publishing Company. 1900. 


ee ee 


 ™~ «i 


1901] CURRENT LITERATURE 439 


bulletin no. 27 from the N. Dakota Experiment Station. Two very interest- 
ing articles are “‘ Twenty years progress in pathology”’ by B. T. Galloway, 
and “ The botanic garden as an aid to agriculture” by William Trelease.— 
J. C. ARTHUR. 

A NEW LABORATORY GUIDE for bacteriology by Mr. W. D. Frost, of the 
University of Wisconsin, is a convenient combination of directions for exper- 
iments, blank pages, charts, and outline drawings in which results may be 
recorded, together with some general information.9 

e first part of the book is devoted to general bacteriology with the 
usual description of technique and laboratory methods. The medical part of 
the book is more complete than usual for an elementary text book. 

General information is given in the form of brief notes wherever a germ 
is mentioned for study, relating its source, time of isolation and discoverer, 
with references to the original and other descriptions. A chapter on tax- 
onomy includes Chester’s scheme of nomenclature of cultural characters and 
Migula’s valuable classification of bacteria. 

he most striking features of the book are the well-outlined and system- 
atic arrangement of material, and the unusual and deserved attention given 
to one of the most important problems in the study of bacteria today, the 
effect of environmental action. The chart blanks are excellently designed for 
the record of the numerous physiological and morphological characters of a 
culture under study. A useful supplement to these detailed record blanks 
would be a few charts such as those recommended by Fuller for recording 
definite position and negation reactions in comparative form. The book as 
it stands, however, is’very complete and will be welcomed in many labora- 
tories— Mary HEFFERAN. 

NOTES FOR STUDENTS: 

THE great prevalence of potato blight (PAylophthora infestans) in the state 
of Washington for the past two years has led to the publication of a bulletin” 
on the subject by the Agricultural Experiment Station, based on experiments 
made at the Puyallup station. Bordeaux mixture applied before the disease 
began to be apparent and again after about two weeks was found to decrease 
materially the injury from the disease.— ERNsT A. BESSEY. 

PROFESSOR Conway MacMILLAN has been studying the growth perio- 
dicity of the potato tuber™ by the method described in this journal (16: 149. 
1891) which he has improved upon by employing a second wheel to magnify 

9FROsT, WILLIAM DopceE: A laboratory guide in elementary bacteriology. 4to 
PP. vili+-200, Published by the author. Madison, Wis. 1901. 

* Bropig, Davip A.: Potato blight and its treatment. Bull. 46, eee 
Agricultural Experiment Station, pp. 15, fgs.5. 1901. Pullman, Washingt 

* Bull. Minn. Acad. Nat. Sci., 3: 355-362, 1901. 


440 BOTANICAL GAZETTE [JUNE 


more the tracing of the auxanometer. He finds the increase in diameter 
rhythmic, with maxima once or twice or oftener in 24 hours, followed by 
minima when growth ceases or nearly so. There are indications of a grand 
period, and some connection between the periodic growth of the aerial parts 
and tubers.— C. R. B 


Miss MATHILDE GOLDFLUS® finds that chlorophyll not only persists under 
the cork of many branches, but that it is present in the medullary rays, in 
the pith, and is even formed in the cortex of trunks of trees at the bottom of 
the crevices in a ridgy bark, as in Quercus and Robinia, Indeed it may 
develop in any living tissues. Photosynthesis by this chlorophyll is active, 
more than counterbalancing the production of CO, in respiration, though no 
quantitative determinations seem to have been made. Inasmuch as. the area 
exposed by branches is considerable, and as photosynthesis continues in 
winter, it is evident that it is of considerable importance to the plant. Miss 
Goldflus and Mr, Miyake* have thus called attention to hitherto overlooked 
activity of the chloroplasts.—C, R. B 


S. Kusano (Bot. Mag. Tokyo 15:42-46. 1901) has published a short 
account of the parasitism of Buckleya Quadriala, one of the Santalaceae of 
Japan. Its haustoria apparently attack the roots of almost any plant, as 
witnessed by the artificial cultures made. As a result of the attack the 
activity of the cambium of the host-root seems to be much increased, since 
a transverse section of that part shows the greatest diameter of wood and 
cortex. ‘“If the age of a haustorium may be estimated from the annual 
rings of the host-root which overlie the sucking process, it is safe to conclude 
that the same organ of Buckleya can maintain its activity during fifteen 
years, and probably longer.”” The author found no difficulty in cultivating 
these green parasites by sowing the seeds in the same pots with the seeds of 
other plants which might serve as hosts.—J, M. C 


IN A RECENT PAPER Noll" takes up again the much discussed subject of 
polarity among the marine algae. Beginning with the statement that in 
Bryopsis muscosa, on which he worked, the polarity is as pronounced as in 
Pinus, he states as a result of his experiments that very few of his plants 
reversed their root and shoot poles when inverted. By measurements and 
dates he ascertained that the young and actively growing plants were SO 
strongly polarized as to resume the original manner of growth ; that only the 
older more slowly growing forms succumbed to the force of external condi- 
tions and turn root into shoot and shoot into root. Hence the inherited polar- 

*2 Revue gén. de Bot. 13: 49-92. 1901. 

*3 Bot. Mag. Tokyo. 14: 44. 1900. See Bot. Gaz. 30: 141. 1900. 


‘4On reversion experiments on Bryopsis, with remarks on energids. Ber. d. deut. 
bot. Gesell. 18 : 444-451. I900. 


t90T]} CURRENT LITERATURE 441 


ization is lost with age. These results agree with those of Winkler of an 
earlier date. To the definition of an “energid,” as given by Sachs, Noll 
takes exception, and calls the Siphoneae “single but multinucleate energids,”’ 
laying stress rather on the Hautschicht than on the nucleus with its dominated 
mass of protoplasm. He therefore defines the energid as a “one or many- 
nucleate plasmatic body enclosed in a definite wall.""— PHtL1p GRAEME 
WRIGHTSON 

‘‘A rhizomorphic root-rot of fruit trees’’*5 is the title of a recent bulletin 
of the Oklahoma Agricultural Experiment Station, For a number of years 
a serious root trouble, especially of the apple, has been attracting the atten- 
tion of fruit growers and botanists in Missouri, Illinois, Oklahoma, and Texas, 
as well as in other regions. This has been shown by von Schrenk and others 
to be caused, in all probability, by a hitherto unidentified rhizomorph-produc- 
ing fungus. In the present bulletin, based upon work taken up since last 
June, the disease is shown to attack other trees also than merely fruit trees, 
and is ascribed to a species of Clitocybe which is described as new under the 
name C. parasitica. This was found at the base of many diseased trees and 
was accordingly considered to be the cause of the disease. Most of the bul- 
letin is given up to a discussion of previous work on diseases of trees caused 
by agarics, to a host index of these fungi as they affect trees, and to a long 
bibliography. The discussion of the disease under consideration is really 
limited to four or five pages.— ERNsT A. BESSEY. 

THE suBjEcT of asparagus rust is one that continues to attract attention. 
A recent bulletin by F. A. Sirrine*® discusses the disease and its treatment in 
New York. All the stages of the rust are found to occur in the state. The 
variations in the distribution of the disease ascribed by some investigators to 
soil conditions seem rather to be due to the occurrence or non-occurrence of 
dews or dense fogs which furnish the moisture necessary for the germination 
of the spores. The measures usually recommended in combating this disease, 
viz., cutting and burning the affected fields early in the fall, are shown to be 
injurious to the plants and not of appreciable value in reducing the amount 
of rust next year. Using a specially devised spraying machine, which enabled 
the operators to wet all parts of the plants thoroughly, was found in two years 
trials under unfavorable conditions to effect a gain in the crop of 69.5 per cent. 
for the first year and of 47.8 per cent. for the second year. The best results 
were obtained with a Bordeaux mixture to which was added a solution of 

, E. MEAD: A rhizomorphic root-rot of fruit trees. Bulletin 49, 

car Agree Fxperiment Station. p. 32. f/s. zz. February 1901. Still- 
water, Oklahom 

*6SIRRINE, F. A.: Spraying for asparagus rust. I. Tests with resin- Bordeaux 
mixture. II. The Downs power asparagus sprayer. N. Y. Agric. Exper. St. Bull. 
no. 188, pp. 233-276. December 1900. 


442 BOTANICAL GAZETTE [JUNE 


resin. Spraying was not found to be profitable where the disease had so 
weakened the plants that they gave less than one third of the normal crop.— 
Ernst A. BESSEY. 

ATTENTION is called to a group of fungus diseases hitherto almost 
unnoticed in this country in a joint bulletin by B. M. Duggar and F. C. Stewart 
entitled “ The sterile fungus Rhizoctonia as a cause of plant diseases in Amer- 
ica.”*7_ The genus Rhizoctonia is a so-called form genus, Z. ¢., it consists of 
fungi whose spore forms are unknown but which agree generally in their 
vegetative characters. The fungus is characterized by a peculiar method of 
branching of the hyphae, by the formation of sclerotia, and by its usually par- 
asitic habit on roots and underground parts of plants. The authors find that 
this fungus is present on many different hosts, including among the most 
important the potato, beet, bean, carnation, lettuce, and radish, Whether 
the same species affects all these plants still remains to be determined by 
cross-inoculations. In most if not all of the plants mentioned, the fungus 
causes a severe root-rot, which often results in the death of the host. That 
the fungus is the cause of the disease was proved in some of the cases by 
repeated inoculations with pure cultures. As the paper is in a measure only 
preliminary the discussion of the special morphology and physiology as well 
as of the taxonomy and relation to European forms is reserved for a later 
paper, in which will also be taken up the results of inoculation experiments. 
— Ernst A. BESSEY. 

PROFESSOR G, J. PEIRCE” has published some interesting studies on 
Sequoia sempervirens, chiefly in connection with the development of suckers, 
a very unusual habit among Gymnosperms. The claim is made that reforest- 
ing is entirely possible if suckers and seedlings are not destroyed by fire. In 
the tendency of the suckers to fasciation the author confirms Frank’s view 
that it is in consequence of an excess of food substances in available form. 
The most prominent feature of the paper, however, is the discussion of the 
albinism of certain suckers. These are by no means uncommon, and they 
differ decidedly in rate of growth, in resistance to cold, in leaf structure, etc., 
from green suckers. The author thinks that the albinism is due to the fact 
that the leaves form and attain nearly or quite full size when there is insuf- 
ficient warmth for the formation of the chromoplasts and. the chlorophyll, 
though enough for growth. One interesting feature of the albinism is that 
when a sucker begins white it always remains so, no matter how favorable 

‘7 DuaGar, B. M. and Stewart, F. C.: The sterile fungus Rhizoctonia as a 
cause of plant diseases in sven Bull. 186, Cornell University Agricultural 
Experiment Station, Ithaca, N. Y. Bull. 186, a York Agricultural Experiment 
Station, Geneva, N., Y., p. 28, figs. 9. psi 19 

*8 Studies on the coast orken edna sempervirens Endl.). Proc. Calif. Acad. 
Sci. III. Bot. 2: 83-106. pi. rg. 


aibit? 


rece tereegrgrser see Wo 


rgor] CURRENT LITERATURE 443 


the later conditions may be for the development of a green sucker. The 
author suggests as an explanation that the sucker is in unbroken connection 
with an abundant food supply, and that there is not the stimulus of need for 
a later production of chromoplasts and chlorophyll, The contribution closes 
with an interesting discussion of parasitism and heredity from the stand- 
point of white suckers. Attention is called to the fact that in this case 
absolute parasitism in habit and structure is developed by the environment 
in a single generation from a long line of independent plants. It would 
seem to the author, therefore, that the influence of heredity is less powerful 
than the power of reaction to certain immediate stimuli, and he closes as 
follows: ‘May not this always be the case? May it not be that what we 
call heredity is really the response to similar stimuli and combinations of 
stimuli occurring in orderly succession in the course of nature?””—J. M. C 


BOTANICAL ARTICLES in annual reports of Agricultural Experiment 
Stations, not heretofore noticed in these pages, are as follows: Report of the 
New York station for 1900 contains reprints, with admirable plates, of 
bulletins 161, 162, 163, 164, 167, 169, and 170, all of much interest to students 
of plant diseases and of some other topics. Report of the Wisconsin station 
for 1900 contains a number of original articles. S, M. Babcock and H. L. 
Russell discuss the ‘‘ Causes operative in the production of silage,” and arrive 
at the conclusions that the presence of bacteria is unessential and even dele- 
terious, that the chief action is intramolecular respiration of the plant cells, 
producing carbon dioxid and organic acids in proportion to the length of 
time the cells remain alive and active, and that the aroma is due to the action 
of enzyms. These are unexpected and important results. The same authors 
describe with illustrations an excellent “Closed circuit respiration apparatus,” 
which would be most useful in a laboratory of physiology. E.S. Goff reports 
on “ Investigations of flower buds’’ and concludes that “in favorable seasons 
of flower formation, many of the buds formed that season, and nearly all 
those formed the preceding two seasons, that have not already flowered, will 
become flower-buds; an excessive apple crop results, which is necessarily 
followed by a light one, because the supply of reserve buds is exhausted.” 
The same writer treats of “The resumption of root growth in spring” and 
“The effects of continued use of immature seed ;” while F. Cranefield writes 
upon “Duration of the growth period in fruit trees.” In the 7th Report 
{1899) of the Wyoming station are excellent “ Alkali studies,” dealing with 
the germination and growth of seeds, by B. C. Buffum and E. E, Slosson, 
and in the roth Report (1900) Aven Nelson gives a list of “ The cryptogams 
of Wyoming,” including the algae, fungi, and mosses. Other botanical 
articles in these reports have previously been published as bulletins. The 
report of the botanists, George E. Stone and Ralph E. Smith, in the 11th 
Report of the Hatch (Mass.) station deals with a variety of fungal and 


444 BOTANICAL GAZETTE [JUNE 


physiological disorders of plants, In the 12th Report the same writers dis- 
cuss a number of fungous diseases, especially tracing a relation between the 
abundance of asparagus rust and the physical character of the soil in con- 
serving water. In the 1gth Report of the New Jersey station, B. D. Halsted 
presents a varied annual report (80 pp.), similar to those of a number of 
years back for which he is so justly famous, Among the numerous topics 
taken up probably the most interesting are “Experiments with asparagus. 
rust’’ and “ Fungi as related to weather.” In the 11th Report of the Rhode 
Island station (1898) H. J. Webber and J. A. Tillinghast present a large 
amount of interesting data upon the feeding of plants with chemical fertil- 
izers, especially the use of lime on acid soils, the articles being well illus- 
trated. The third part of the 22d Report of the Connecticut station (18¢8) 
opens with four articles by Wm. C. Sturgis on diseases of melons, lima beans, 
tobacco, peach, and use of fungicides. In the 14th Report of the Maine 
station W. M. Munson has interesting articles on ‘“‘ The blueberry in Maine,” 
“A comparison of large and small radish seed,” “ The effect of sub-watering 
radishes,” and “ Pollination and fertilization of flowers.’”’ The botanists of 
the Vermont station, L. R. Jones and W. A. Orton, give in the 12th Report 
(1899) a valuable account of potato and apple diseases and their remedies, of 
killing weeds with chemicals, and of parasitic fungi of Vermont, bee the 
second list.— J. C. ARTHUR. 

-WE ARE TO HAVE from Dr. Blackman, of the University of Cambridge, 
a series of papers on the algae, that will aim to give a critical account of the 
work that has accumulated since Wille’s treatment in ‘“ Die natiirlichen 
Pflanzenfamilien”’ of Engler and Prantl in 1890. The first paper” has 
recently appeared, and, excluding the Cyanophyceae, takes up the simpler 
types of algae, discussing the possible relationships and lines of evolution. 
The author starts his lines of development from the level of the Protococ- 
coideae, and lays special stress on the Chlamydomonas-like forms, finding 
in their varied life histories several possibilities of evolution, worked out in 
different groups of the Chlorophyceae. There are three conspicuous tend- 
encies, resulting in the well-known types of vegetative organization called by 
him the volvocine, tetrasporine, and endosphaerine. 

The volvocine tendency lays emphasis upon the motile phases in the life 
history, and leads to the specialization and segregation of the ciliated cells. 
in the complex coenobia of the Volvocales. A tendency on the part of rest- 
ing cells to form filaments, cell plates, and membranes is termed tetra- 
sporine, and results in forms like Tetraspora, Hormidium, etc., and through 
them leads to the thallus of the Confervales. In the endosphaerine condition 
vegetative division is reduced to a minimum, and in its place is found the 

‘BLACKMAN, V. H.: The primitive Algae and Flagellata. An account of 
modern work bearing on the evolution of the Algae. Ann. Bot. 14: 647. 1900. 


Igor ] CURRENT LITERATURE 445 


peculiar form of multiplication illustrated by Pediastrum and Hydrodictyon. 
This is essentially a method of spore-formation with the peculiarity that the 
zoospores, unable to escape from the sporangium, settle down to form a new 
individual within the mother cell. These three conditions have been recog- 
nized by several phycologists, and probably express the history of events fairly 
correctly for this region of the algae, where the original conditions and later 
modifying factors seem relatively clear. 

The Conjugales, Siphonales, and Diatomales form natural groups that 
cannot be closely related to the Protococcales or Confervales. They have 
not been treated in this paper, which deals chiefly with the Chlamydomonas- 
like organisms, flagellates, and a certain assemblage of forms termed the 
“‘Heterokontae.” It is doubtful whether most botanists realize that the 
immense group of the Flagellata resembles the lower Volvocales so closely 
that no hard and fast line can be drawn between them and the algae. This 
relationship is treated at some length, and should be welcomed. In his 
remarks upon the primitive Phaeophyceae Dr. Blackman enters a field of 
acknowledged difficulty, and while he presents certain possibilities the 
reader cannot but realize that he is speculating in a region where there is a 
conspicuous paucity of knowledge. 

The author of a paper of this character naturally lays himself open to 
criticism, for his is an attempt to connect and relate groups as definitely as 
possible with confessedly fragmentary evidence at hand. Certain difficulties 
are presented in this paper which cannot be easily overcome. Is not 
Chlamydomonas much too high a type on which to pivot so many evolutionary 
lines? What is to be done with the simplest of the Pleurococcaceae? There 
is little or no real evidence that they are degenerate forms, and this view 
seems to have its chief ~azson d’étre as an assumption to dispose of a trouble- 
some difficulty. Chlamydomonas is very complex as a cell, with its differ- 
entiated activities, sex, etc. There were, of course, immense stretches of 
simpler forms below this level, perhaps rising out of the Cyanophyceae. 
We know almost nothing of the cytology of the Pleurococcaceae, the Crypto- 
monadineae, and other border groups. It seems to the writer that we may 
hope for important results from this field of investigation. We shall probably 
have to go much farther back than Chlamydomonas before we can hope to 
clear the maze of relationships in the Pleurococcaceae.— B. M. Davis. 


SCE MSLETITERS. 


USE OF FERN NAMES. 


WHILE Messrs. Fernald and Pollard are discussing the names by which 
our American ferns should be known, a few observations upon another side 
of the subject may be inseason. I refer to the practice of fern students in 
the use of these names. There are in America about two thousand persons 
at present engaged in the study of our ferns and fern allies. Of this number 
I am confident that fully nine tenths are committed to what may be called a 
conservative nomenclature. Granted that the names given in the sixth edi- 
tion of Professor Underwood’s Our Native Ferns are correct according to the 
Rochester code, the mass of fern students see in this no argument for adopting 
them. The reason why they do not is easily found. In adopting most of the 
names proposed, the student would be departing from almost world-wide 
usage and bringing confusion into a part of the study that has thus far prac- 
tically escaped. A single instance will illustrate. Cystopteris fragilis is a 
plant of almost world-wide distribution. It is found in the West Indies, 
South Africa, India, and Alaska, as well as the United States, and is every- 
where known by the name I have here given it. The question may be fairly 
asked, then, what it would avail American students to unite with Professor 
Underwood in calling it Filix fragilis. They would only succeed in making 
themselves misunderstood to fern students in other parts of the world. On 
the other hand, certain changes, such as Athyrium for part of Asplenium, 
and Polystichum for part of Aspidium, have been readily adopted, because 
these names are in common use abroad, and by accepting them the American 
student comes more into harmony with universal usage. In thus rejecting 
some names and accepting others, the fern student is really consistent, 
although at first glance he may not seem to be. 

The fact that many of our ferns are common to the old world, also, puts 
a slightly different aspect on the subject of their nomenclature in the opinion 
of the student; we cannot “go it alone” in the matter of names. And, after 
all, those interested in the plants themselves care very little for improve- 
ments in their names. At the same time, the value of knowing what names 
have been proposed for our ferns in the past is not underestimated ; these 
names are a part of the plant's history. But the proposal to adopt them is 
quite another matter. Just as we have agreed not to go beyond Linnaeus for 
specific names, although there were such names before his day, so the fern 

446 [JUNE 


1901] OPEN LETTERS 447 


student prefers not to go back further for his fern names than, say, the fourth 
edition of Professor Underwood's book or the sixth edition of Gray’s Manual. 
In doing this he is essentially adopting the brand of nomenclature advocated 
by Mr. Fernald, without attempting it. 

I am aware that in adhering to certain names regardless of anything but 
common usage the student is open to the charge of being unscientific, but 
this does not alter the fact that he does adhere to them. Moreover, by a 
very few changes in his nomenclature, he can come into harmony with the 
great majority of students who happen to be following practically the rules 
suggested by Mr. Fernald, and so avoid being unscientific. The whole 
question resolves itself into which style of nomenclature will give us stability 
and uniformity throughout the world, with the least change. Some indica- 
tion of how the tide is setting may be found in the fact that no less than 
three books on ferns to be issued in the United States this year will use the 
conservative fern names.— WILLARD N. CLUTE. 


NEWS. 


Dr. Wm. BENECKE, privat docent in botany in the University of Kiel, 
has been appointed to an associate professorship. 

Dr. B. NEMEC has been appointed Director of the new institute for 
Plant Piidiuny of the Imperial Bohemian University of Prag. 

ACCORDING to Science, the extensive herbarium of the late Dr. T. Bernard 
Brinton has been presented to the Botanical Garden of the University of 
Pennsylvania. 

EpwarD W. BERRY a member of the Torrey Botanical Club has been 
awarded the Walker prize of fifty dollars by the Boston Society of Natural 
History for a memoir on Liriodendron. 

THE TRANSFER of the late Dr. J. G. Agardh’s herbarium of algae to the 
University of Lund is made on the express condition that no specimens be 
loaned. While this will doubtless tend to preserve a valuable collection of 
types, it seriously restricts its usefulness. 

A RECENT NUMBER of Plant World announces that the herbarium of the 
veteran collector, Mr. A. H. Curtiss, containing about 16,000 sheets, was 
destroyed in the recent great fire at Jacksonville, Florida. His early collec- 
tions for this year’s distribution were saved. 

A SUMMER SCHOOL for nature study is to be held at the Rhode Island 
College of Agriculture and Mechanic Arts, Kingston, from July 5th to 2oth. 
A course on trees and garden plants will be given by Professor Fred W. 
Card and Mr. George E. Adams; on flowering plants by Professors W. W. 

Bailey and H. L. Merrow; on seaweeds and fungi by Dr. A. B. Seymour. 

Ir IS WITH great regret that the BOTANICAL GAZETTE records the 
death of Professor Thomas Conrad Porter of Easton, Pa. He died April 27, 
Ig0I, at the age of seventy-nine. Since 1866 he has been identified with 
Lafayette College, and has long been one of the best known American tax- 
onomists. A biographical sketch will be published in an early number of 
the GAZETTE. 

THE BIOLOGICAL station of the University of Montana, established in 
1899 on Flathead lake, will be opened this year on July 22. The University 
supplies books, chemicals, glassware and microscopes for use free; students 
are charged only for material used and breakage. The botanical work will 
be in charge of Dr. D. T. MacDougal and Mr, R. S. Williams, of the New 
York Botanical Garden. 

448 [JUNE, 1901 


BA Royce has yore nara 


ee PE aaNet eee 


GENERAL INDEX. 


The most important classified entries will be found under Contributors, Personals, 


and Reviews. ew names and name 


s of new genera, species, and varieties, are 


printed in bold-face type; synonyms in ¢adics. 


A 


Abdra, Greene on 212 
Sion crispum 330 
Acerates sence 3 
Adams, George E., personal 4 
Adiantum eapilus Feneas Bek Under- 
ood on 281 


Median Ane e@ 331 
egopogo ep higenoniiery a 

Agardh, J. G., personal 367 

rested hea ho n malioration of 71 

Agriculture, Department of 2 

Ceerciers science, Eee a of So- 
ciety we r promotion of 43 

Agrimonia, Robinson on 21 

Agrostis, Scribner and saa on 213 
Airosperma, new gop ag 

Aldenella, Gre eene 0 

Algae, B lackman on or ; Noll on 440 

Allocarya, Greene o 

eet cag cLesnuatie 117; macranthus 


Aistcas School of Natural “aad 287 
Amphilophium oxylo Lge i 
Andropogon, Scribner d Merrill on ri 
Anguria, longipeduncolats 112; ovata I 
— canthus 

moe acerifolia 330; hastata 
sae preg on 280; sieuiaeinad: 


kane Kien n 363 

Aplopappus, inerio eibee on 212; 
linearifolius, Rydberg o 

Apogamy 347 

pponpory 3 47 
Aquilegia pubescens a7 
Arabis, Gre 

Arbutus. diversifl ae 

ave liella, Cavara on 135 

A — tapes iflora 

Ari macrospa sea m 32 

pening  Gcibad rand Merrill on 213 
rnica, Greene on 212; celsa 408, Cha- 


Greenei 406; incana 408; 
latifolia ie: sieaaieeont 406; wae 


JUNE, Igor] 


ao Leh agile 4075 rhizomata 409; 
7; vento 407 

A on i G 439, 

Asclepias Caliiocnica #9 

Ascobolus pena ming 

Ascocarp of Pyronema, bahes on 139 

Asia, Lipsk yon ‘Hore of 364 

As yaaa — Halsted on 444; Sirrine 


n 44 n 
Aspergillus avis 126; germination of 
Spores:44 
Augie hemionitis 349 
Asplenium ebeneum, Davenport on 213 
Aster, Greene on 212 
Ayers, H. B., work of 210 


B 


Babcock, S. M., work o 

Baccharis hirtella 331; multiflora 331 

Bacteria, Jordan on dis Sappearance of 
sewag e 279; of the air, Weinzirl on 
2 


Bacteriosis of walnut 272 

= dhamia utrienlarts ier 198 
yeas 5" Botiny 1 “Cyclopedia of 
American Heicaitare 436. We W., 

8 


pamone ai oe. new genus, Rydberg on 


Bamlers, new genus 297 

Barnes, GC: '-Ri, 60, 70,.,.71, oc = 208, 
275, 277, 358, 361, 436, 437, 

biel g sabulige set 

Benecke, Wm pc 448 

seman Bs: Worsdel. a ak 

ial trifolia 

Bergen’s “ Foundations w wisi os 

M., pers 


ard W., ses ae 
Bessey, C. ie ork of 133 
Bessey Be A. 207, 439, 441, 442; personal 


Best, G. N., “North American species of 
Heterocladium” 437; work of 364 


449 


450 


Rea Alaskana 236; alba 236; fontin- 
alis 239; occidentalis 236; papyrifera 


23 
Bidens, pro _ 280 
Bigelow, F., personal 367 
Biltmore oda Studies 360 
ika 288 


Biometr 
Bissell, C. H., work of 
Blackman, V. H., wor 


k 
Blakea, grandiflora FI2; rf Ale III 
Blueberry, Muns n 444 

Boletus, germination of spore 

pa ran nd Du Sablon’s “Coan de bo- 


nique ” 437 
Hisackes eheereig eh 143 
Borys vulgaris 126, germination of 
spore 
Brainerd: Jones, and Eggleston’s “ Flora 


Brassica nigra, Rob 
ae got growing in 352 
Il 


ork 
Brickellia grandiflora il 
36; veronicaefolia 
Be a sional 448 
of 363 


tomentella 


e, Davi ork of 439 
Bronghiarta, rg ; he casa 336; seri- 


iota heteroclitum 1 

Brown, Ste pees ewbaatal 142 
Brown, H. C., of 281 
Bepuiie muscosa, "Noll o on 440 
et bi cig Na a Terras on 133 
Buff B. C. work of 443 


Bulbils, soral $5 
Bu i aes 2 281 


Bur a P ork o 
Buitensore: oemical carlin ‘of 423 
. 


Caeoma or pilin Bi ce 331 


Calathea, dasycarpa 123; ao 
Oo rapax I 


24; Petersonii 125; Ver. 
Calea, hypoleuca oa saretacivcht 333 
negenatims Inlet s from, Eastwood 


n 135; Parish te 135 plants of 388 
Callisia ee Sree 


Card, Fred W., eit 448 
Cardamine, HAA si eon 212 
Carex, Fernald o n 364; arp on walla 6 
ent of pollen grain 284; firm 
hispidula 18; supina 


216 
nia, new genus, Greene on 212 
Cassia Absus 336; multiflora 336 


BOTANICAL GAZETTE 


[JUNE 


Caucasus, Sommier and Levier on plants 
o 

hata pe work of 135 

Cel w genus, Greene on 212 

Gelosia ‘latifolia 26 

Central America, undescribed plants from 
10 


Cestrum nitidum 326 

9 ome ctis heterocarpha curta 391; Xan- 
a 391 

rr oh a Jonesii, germination of 


spores 44 

Chamberlain, C. J. 205, 210, 284, 363 

Cheiranthus, 394; asper 394; Greene on 
212 

Chemotaxis in fern sperms, Buller on 281 


ee and Wilcox’ s “Stock poisoning ~ 
nts of sini sn Du 
Ches ae ede dD. Per 


China, Diels on Pig a cnt 364 
Chlorophyll, Goldflus on 
Chomelia microloba 1 
Chrysopsis fulcrata 399 
Circinella umbellata, 

spores 44 
nen Pies M., 


Seales of 
and Jones’ “Trees of 


met 2:2 
Ca eues Petri on 135 
Cle iene oe sai Scottii 394 
Clemen d Cutter’s “Manual of high 
Zi 


Brazil 352 
Rapid’s flora” 437 
Colecsanths, Me as on 213; congestus 
1} grandifioru 
Rcccriem anceps f wh paraphysatum 
a sae Verbesinae 33 


MS, Fo Ses Shoes ae 287 
Sc callos 
Colors of flower: 


oshi 
Columnea, mieroealyx microphylla 118; 


sulfurea 
Conifers, Von Schrenk on diseased: of 


ew England 361 
Conoclinium, Greene on 280 
Contributors: Arthur, J. C. 439, 4445 
Barnes, C. R. 69, 70, 71, 132: 133, 208, 


275,277,359) vias 436, 437, 440; aii ey; 


205, 2 206, 207, 213, 278, 
360, 436, 437, 438, 440, ee Cowles 


ots a is 5 ie Ce Be > ae 


1901 ] 


H. C. 73, 133, 137, 145, 208, 209, 210, 
212, 278, 283, 361,.362;. Davis, B. M. 


Duggar, B. M. oh et ong DG, + 35 

4253 ie 183; 

281; Gan ow. 4345 Hall, a Ms 

388: Hanititwarer ay . 203; Hef- 

rg Mary 439; a aca G. M. 3395 
Theo. 17; way, E. . 

32 Tefizey, E. oF ss, eles 


sey j geneae ty “ 4 yon, 
Floren 15; Nelson, Aven 394; 
Pierce, Newton B. 273; Pollard, C. L 
286; i hes HK. 728; bee Sere 


Cha 217; Schaffne 
ee Smith, Toha Donnell cone Smith, 
Mary H. 127; Timberlake, H. G. 203; 
O. 241; Trelease, Wil- 

M. 366; 


U 
Whitford, H.-N. 133, 135, 211, 289; 
i ; I 


Copelan 44105 phe 216 
Ce lag 

oe germina ation of spores 44 
Corbett, L. C., personal 287 

Corr rrens, Cw ork of ir 

Costus sanguineus 122; spicatus 123 
Cotyledon, as geotropism in 410 
Coulter, John 68 


rs) 
Q 
Ps 
-o 
mn 
o 


Sperm rmatophytes”’ 
‘Monograph of nt North American 


Re 
3, 133, 137, ig 208, 209, 
210, 212, 278, 283, 361, 362 
Cranefield, F., work of 443 


Crataegus r on 280; acutifolia 
217; : oldiana 221; 
paptoweiart: 230; apogee! ate i 


33 mplainensis 223; coccinea Io, 
peP imniah aoa 14, mollis 7, ceriea folia 
14, swbvillosa 7; collina 221; Colum 

biana 229; Crus-galli 3, 218, 220, 2333 
dilatata 9; Engelmanni 2; erecta 218, 


227; moilis 8, 224, 226; sci 7; 
nitens 233; nitida 2 a3t: 

226; Peoriensis 5; pra ratensis gee 
sili oe submollis 7. 0, 222: poss 
7 10% 225; tomentosa 221; ‘omen- 
tosa 7; goers is: nitida 231 


INDEX TO VOLUME XXXI 451 


Crawford, James D., personal 743 
Cucurbita Pepo, geotropism o 416 
niana 239, pea 239; 


3 239 

- ji ean 448 
Cutter, see Cleme 
Cycas ‘revoluta, cana of 265 
Cyperus, pro ophyllum ob 2 
Cytology 5 gee 369 
Cza of 71 


apek, wo 
D 
Dale, El lizabeth, work of 362 
Dalea Jamesit 395 


Delame w genus, Le Moore on 213 
Denn ay “ Plant life a structure” 277 
ae de soiaeny 350 

: nthodium ova 


34 
Detabley’s 4 Flora or ‘Cheshire” 360 
Diels, L., 
Digestion of food by Fuligo 
s, Dugg nore Stewat 0 on fungu 


n 439; root- rot of forest trees, Wilcox 
n 44 
Dicée: H. H., work of 362 
Dodecatheon Jeffreyi redolens 392 
Draba, Greene on 212; Breweri 392 
Druery, Charles T. 351 
Difooterss | Maxon on 
Duggar, B. M. 38; work o 442 
Du Sablon, see Bonnie 


E 
Eastwood, Alice, work of I 


Ecology, Nilsson on 209; of Northern 
Michigan 289; of oe 93 1455 
36. 


Merrill on 213 
Soyer natans 339 
ss dr cacao Burns on 209 
6 


lien” 


452 BOTANICAL GAZETTE [JUNE 


Enzymes in Fuligo 20 

Eremosemium, new pence Greene on 213 

Erigeron 392 

Eriobotrya Japonica, sens : 273 

Eriocaulaceae, ro a = ure of 

Eriocaulon dec 

Erythronium, life oem and cytology of 
6 


Escombe, work of 281 

Eucalyptus globulus, at Site of 416 
Euglena viridis, Wager on 140 
Eupatorium, atrom Gielen 460 ; brevi- 


pes 332; per pata 333; grandi- 
rum 401; 


maculatum 400; Greene 
on 280 

Exoascus, germination of spores 44 

F 


-lemsialaanee DG: 354, 425 
ea trinerv rvia 115, Suerrensis 11 5 
ala uson’s “ Soba of the U.S. species 


Pe 
Fern, names ee variations in Great 
Britain 
Fernald, M. L. 183; personal 287; work 
I 


of 2 ‘hg 364 
Fern B. E., personal 367 
Eertilisation, Munson on 444; Sargant 
on double 280 
Ficus Carica, bacteriosis 7 273 
Fitting, Hans, work of 2 
Flowers, Goff on = of 4433 a on 
colors of 278; 
rest, S 


» Hollick on 135; 
ale sei oni a Schowl of ‘21 
re on 212; neglecta 388; 


Frost’s “ Laboratory guide in elementary 


bacteriology ” 439 
haa trees, Wilcox on root-rot of 441 
F 


Fuligo, ceptica, Leagan of 19 
Fungi, ger on of spores 


and wi 
ag terse South b 
Fungus diseases, Dheae and Stewart on 
442 


G 
Galagania, — genus, Lepsky on 364 
Galium uncinulatum 


Galphimia Humboltiana 328 


Ganong, W. F., 434; personal 216 

Garden, Botani cal of Buitenzorg 423; 
Missouri Botanical, scholarship 142; 
12th Report of 144 

Gases, Brown and Escombe on static 
diffusion of 28 

Gaucher, work of 210 

Gen waoase) oe On 212: ; Moseleyi 396; 
unica 

Goebel’ Ce anapiaplite der Pflanzen” 


ie. Kennedy on plant 20 
Geology and forestry, Hollick on we 
Geotropism, Czapek on 71; of stems 410 


241; of poles a08 . spores 38 
Gertrudia, hg cs dette and 


rb ae 
a ae Osu. «Les problémes de la vie” 
Gildermeister and Hoffmann’s “‘ The vola- 


oils” 278 
Gilis gdhicus 390; modesta 389; Par- 
Gleditsia. aquatica 2; Texana 1; tria- 
canthos 
Gnaphalium plantaginifolium, Robinson 
on 21 
Goff, E. S., work of 443 : 
Goldflus, Mathilde, work of 440 
Graebner, work of 304 
seats ve ct of hydrocyanic acid gas 


Giand Gary, work of 283 
Grasses, Scribner on 364 
Graves, Henry S., cence 216 
Gray Bulletin 287 

Grayta 


Greene, Edwar we th Plantae Bakeri- 
: oe work of 212, 28 
Greenm M., work o 


nman, J. 
Growth, in fruit trees, Cranefield on 443; 
e potato tuber, Mac- 


4 

Grouts Sines es of bbs it OOS 
““Mosses with a hand len 132 

Gurana, Makoyana 113; eee 3343 
Tonduziana 112 

Guatemala, undescribed plants from 109 

Gymnolomia, Ghiesbreghtii 333; sub- 
flexuosa 333 


H 


Hall, H. M., 388 
Halerpestes, new genus, Greene on 212 


rete a Nee A ON Lo can aoe 


1901 ] 


Halsted, B. D., work of 4 
Harding’ s “Schwarze Faulniss des Kohls” 


are r, R. A., work of 139 
Pee John W., 203, personal 143 


Hegler, R., 

Helianthus, Recah Ft on 135; an- 
nuus, geotropism of 4 

Baiiicar nc, Americans Fae te 110; 
110 


EA) Or hu iG, 
urtis 448}; of 
- Bernard Brinton 448 Rocky moun- 
B20. 
Beibann M. J., personal 
Hernandia, didymantha ners Guianensi 
21 


at eterocladium, Best on 364 

Hibiscus vitifolius, Dale on 362 

Hiern’s*"! ‘Catalogue of Welwitsch’s Afri- 
> 69 


a, new genus eck eons on 364 
H aa see Gilder meis 
Holferty, G. M. i 330 
Hollick, C. A., work of 135 
Holm, Thee: I 


ci Sir Joseph, personal 72 


Howe’s “ Hepaticae of Vermont ” 206 
He. Abbé, simoiieras 142 
Ld 3 One Be rk of 1: 
Hydrocharis ‘Morsus- -ranae, Terras on 
winter buds of I 


eee. ee swarm spore 
formation 
Byucisrappes. : new genus, Rydberg on 


21 
Hypocotyl, positive geotropism in 410 
I 


Indigofera, ros caer 336; Mexicana 
28; Palmer 

Insects in the Sonest Smith on 135 

International Botanical Maasen 367 

Iostephane heterophys 334 

[pomoea, Rendle on 213 

jpchinnsiphon Moras leiostachya 123 
ystrix, Se and Hill on 136; 

“Fi ra on megaspore of 21 


J 


Jack, John G., work of 2 
Jackson's «A glossary of Motenié terms” 
9 


INDEX TO VOLUME XXXI 


453 


Japan, agian publications 216 
Jatropha uren 
Jeffrey, E. 387 

epson's ’s “Flora of western middle Cali- 
fornia” 435 
Jequitb trees 352 
Jon ed So 


work of 444; see Brainerd, 


aa ate 
ie EO. 280 ; work of 279 
ring Asa Gray Bulletin 287; Bilt- 
ore en, Studie s 360; Biomet- 
Botan ‘chet Centralblatt 


oa =e World 287 5 ; Torreya 216 
Juel, H. O., work of 2 
Juglans regia, sashes of 272 

K 


Kacyobigeets in Erythronium 371 
Kearney, T. H., personal 216; work of 
208 


Keller, Ida A., personal 143 
Kentrochrosia, new genus, Schumann and 


lett, A., work of 363 
Knowlton, F. H., ini 
Kolkwitzia, new gen nein on 364 
Korshinskia, » ew ete eed on 364 
Korshin S. J., personal 1 
uhnia euparivite 402; corymbosa 404; 
Fitzpatricki 402; glutin 4043 


Seerierpl 402: Hitchcocki ee re- 
ticulata 
Kusano, S., Tonk of 440 


Laboratory, Marine uinereag. 288 

Lachnocaulon Michauxii 33 

Lacinaria ~ 404; " eatiatyiie 405; 
scario 

Lamson- Sateen F Cokie of 364 

pela priaes eb 

‘egpdiny formation in the 


euie of 
tates ilkets 349; npc -mas 347 
Latex, Gaucher on fun of 210; in 
rubber plants, pesian on ee 
Lauterbachia, new genus, Schumann on 


retina Miss oi bans on 134 

Law Asi. 

Leaf of Erigcalon once 28 

on anatomy of varie- 


Leiberg, John B., work of pi 
oe and roots, Terras on 141 
Lepi on, new genus, Scott on 279 


¥ coscalllos, Scribner and Merrill on 21 3 


454 BOTANICAL 


ihc ae work of 364 

Lew ork of 2 

ewan taicise 337 

aye scariosa 406 3 scartosa 405 
Life, ‘A. C. 26 

Li ht a nd germination, Terras on 133 
Lippia Pringlei 33 


sky, W., wo 64 
Literature, reviews of botanical 143 
hi lage 9 hi ee eg um 399 


n, B. E 

Seca oie soa 6 
papnemschys Guatemelensis 119 

anthus Eur S 39 
Lot isy, iP Of I 137 
Lupinus albus, hae Ee of 4 
Lycopodium pg pho oben on 141 
Lyon, Florence 215 
Lyon, Harold L., work of ae 


M 
MacDougal, D. T., personal 448 


Macronema 21 

pert ede Api ang: ou poe and Lau- 
terbach on new gen 07 

Macroseepis convection 116; pleistan- 
tha 


MacMil lan, Conway, work of 439 

Ma peeie gee ache bacteriosis of 273 

Makino, T., personal 21 

Mulipemation. Davis on 362; of Agaricus, 
Smith on 71 

Markea leucantha 116 

Marsdenia Mexicana 330 

Marshall’s “The mushroom book” 358 

Martellia, Mattirolo on 135 

Mattirolo, O., work of 135 

Maxon, William R., work of 212; “ Ferns 
and fern allies of North America er or 

Meehan, Thomas, Selptege 142, 287 


280 
Megasporangium of Enytheonium 373 
Melilotus, Schulz on 364 
Mentzelia hispida 337 
Merrill, El] aor 19 ~~ of 135, 213 
errow, Ve Bem ae nal 448 
m 


a 

mbryanthem cot bBieiiek on 211 
etalstelma seg ssifoliem 330 

exico, fun 

ichigan, euie 

imulus nape Tilingi 399 
inks, ae ko 

innesota, eler on ecology of 361; 
seaside “botania station 

Mitosis, Dixo 62 

Miyake, work of 133 


4-4-8 
: o 


O 
wor 
Ov 


GAZETTE [ JUNE: 


oo . work of 278 
Moisture, effect of on plant ar ees 


397; clinopodioide s 398; menthaefolia 

308) Nuttallii 397; Ramaleyi 398; 
a 398; fist ulosa, Fernald on 213 

Moni fructigena, germination of spores. 


pcan saprogena 109 
Montana, ey ae Station of the Uni- 
he rsity of “ay 
oore, Spe 
ete. yo ieicilen of 426 
is, E. L., work of 364 
ucor, germination of spor 
Sanne ge eee “i “oF Merl on 213. 
he De of 4 
yee 0a, he tins anise if: 198 
Myrodia Guatemalteca I10 


N 


Napeanthus repens 118 
ali e study, summer —— for 448 
aska, = ie trees in 133 

ce. Aven, 3043 Were tf 443 

eg sage Lyon on 278 
B ie eal 446; work of 133 
Reese. ew genus, Greene on 213 
aahaaed A. personal 73 work of 208 
be, F. C., personal 216 

te w Jersey fo orests, Vermeule on 134 
Ni ge e Moore on 213 


) 
Nitrates as a source of nitrogen 126: 
Nomenclature 285, 365; in oiee geogra- 
se ag re dag a on ale 
s Byxb 134 
ela neal Webber ond ‘Tillinghast on 444 


O 


yaa apmplae albidum, germination of 
Spor Mead 


hygen 
Gegoneis csbeheebre ala 399 
bi joven ee non-sexual propaga- 


Ore sont emma, new genus, Greene on 212 
Oreastrum, Greene 0 on 212 
oO eopanax capitatum 114; macrocepha- 
lum 114; pycnocarpum I13 
Oroban chacese, seeds of 39 
rton’s “Parasitic fungi of Vermont” 2063. 
k of 


ularia primulina, germination of spores 


Ovule of Potomageton natans 339 


1901 | 


P 


leh Ward on work in a 
, ocribner and Merrill on 213 
Parsi of Buckleya alin, Kusano 


Parathesis eae 115 
Parish, S. B., pat 2g 135 
Paikic. wc ork of 2 
Paro Sie famesii 395: oer 395 
Parrasia, Greene 

lu , Series abil Merrill on 213 
59 s a ye ort of 
” 359 
Penhallow, D. P., personal a work of 

ou 


e, G. J., work of 422 

Penn glaucum, germination of 

spor 
Percival’s <aggicultral beg dl 67 
Peritonia, Greene on 2 
Personals: Adams, G, "r, 448; Cigar 

ie 307). a epespaed W.., 448; Bai 

S. M enecke W. 438; pete 
G. ne E. W - 448; Pint 
E. A. 287; Brinton, T. B. 448; Brow 
5.142; Card, F. W. , 448; Carleton, M. 
A. 216; Chester, F. D. 2 16; Collins, F. 
3.287; Copeland, EB. 2165 Corbett, 
LC. 287%) Coulter, S, M. 287; Craw- 


ford, J. D. 143; Curtiss, A. H. 448; 
Fernald ; Fernow, ae : 
367; Ganong, W. F, 


216; Graves H.S. 
R. 4 287; Harshiberger, 


VY IH "216: payer Ida A. 143; 
Rorttineky. S. J. 142; MacDougal, D. 
aa ; i 


448; Makino, T 216; Meehan, T 
142, 287; Merrow, H. L. 448; Nemec, 
B., 448; Nestler, A. 72; Newcombe, 
C. 216; Penhallow, D. P. 216; Porter, 
T. C. 448; Pound, 367; Pringle, C 
G. 287; Raciborski, 142; Rober 
H. F. 288; Schiffner, V. 142; Seymour, 
A. B. 448; Shear, C. A. 287; Smith, E. 


a8 
ebster, H. 287; Weittstein, R. von 142; 
Williams, Ry ae 448; Williams, T. oe 


143; 
fees xi eoralie 395; tenuifolius 
16 


Petri, L., 135 
aeny hack EF 399 
Phalacroseris Bolanderi Salenate 393 


INDEX TO VOLUME XXXI 


455 


5 gn pest ai! de 350 
Phoenix, geotropism of 419 

Phot deieed S, Goldfus on 440 
Phycomyces nitens, germination of spores 


si in 126, 241; of fungi 38; of 
reproduction in fungi, Klebs on 208 

Pierce, Newton B., 273; ‘‘Peach leaf curl” 
20 


sas aren 122, ptericlada 121 
us Sabiniana, geotropism of 416 

P: sea rave oak of ~~ 

Pitcainds Palmer 

Pittier’s ‘‘ Primitiae — Costaricensis ”’ 


131, 205, 

Plant W 

Plantago, Morris on 364 
Plasmodia of Fuligo septica 198 


Podochilus, Schlechter on 13 
Poison of Primula obconica, Nestler on 
20 


Polemonium Archibaldae 397; confertum 
392; filicinum 397 

Pollen, Correns on 361; germination of 
39 

Polanisia Sarpy es on 212 

Sart eh on 4 

ard, (. “86. ‘otk of 135 

Pollination, Vince on 444 

Pol i ulata 328 


gd pens eienaae Maxon on 212; 
vulgare 351 

Polystichum angulare 347 

Porphyra, Hus on 135 

Porter, Thom , death of 448 

Potamogeton penn ovule and embryo 


Potato see Brodie on 439 ° 
Co) oe, personal 367 
Preston, CY et2 
Pringle, Cyrus G., personal 287 
mula obconica, Nestler On poison of 
08 


i a hes of iano 34 
Pseudo omonas, campes Is oo 
Pg ee spans 2725 Stewar 272 
Pteris, aquilina 348; cretic 
Puccinia, Aniscanthii en Asteris 332; 
Baccharidis 331; Baccharidis-hirtellae 
334 multiflorae 331; Ber- 


Coulterophyti 335; Des- 

334; Mates 3333 espino- 

ferox 333; ii 330; 

334; heteruspora 330; 
3324 Iostep. 334;_Kuh- 

niae 330; iateripes 329; Marsdeniae 

330; Menthae 330; Oaxacana 331; 


456 


Philibertiae 330; Pitcairniae 3303 
se orsa 332; Ruelliae- cE oe One 
padre begin 330; it ace a 332; 
Wieck: € 333, 337; Xanthii 330 
Pucciniosira Brickelliae 336; pallidula 
336; shed 336 


Pyronema confluens, Harper on 
a ene an new genus, Ry fe: on 212 
R 


Raciborski, M., personal 142 


Ra hades s, Lewin o 
Raaia, Peoiarc ont ral iad epiphylla 


+ Indica...336; ati 336; 
spinulosa 336 Stan 
Rendle, A. B., work of 2 
Doren in “Opuntia 127; of Pyro- 


a, Har 139 
Respiration, ico and Russell on 443 
Bailey’s “ Botany” I 

ao ic 


orth America 
Dennert’s ‘e Plant 
sti ge Lata 
ngler’s ‘“ Pflanzenreich” 206; 
recs and Prantl’s “ Natiirlichen 
Pflanzenfamilien” 205; F 


bye S. 
“Labo oratory guide 


in elementa ary bacteriology 439; Goe 
bel’s “ Organogra der Pflanzen ” 


sary of botanic terms” 69; Jepson’s 


BOTANICAL GAZETTE 


[JUNE 


“Flora of a phair Se 
Marsha ie Pg OOk 
Max s and fern allie of 
pei Ort “ Para= 
of Ve rmont ” ee ; Peck’s 

dible oa ungi”’ 359, “Report of the 
state botanist of New York” 359; Per- 
erval’s. * te eyeing n 
Pie erce’s “Pea h le aft carl?’ 


S501 
Rhizoctonia, Duggar and Stewart on 
os 

Rhizome, structure of Eriocaulon 24 

Rhopalocn emis, soe n 138 

Ricinus, com , bacteriosis of 273 

Robinia Futuna, geotropism of 416 
3 


pe 

on, 441: 

ransmission ofa 
stimul 1 eae 

-ptbenps oe col ene 265 


Rubus 327 
Ruellia Bourguei 329 


13 
k of 443 
Rydberg, P. A. work 6f 212, 280 
Ss 


Salix glauca villosa 392 


work of of 2 

Sargent, Char 3S. f, Fei work of 280 
Saxifraga Draaleie austrina 394 
Scape of tare 25 

Schaffner, J. H 

Schiffner, Vict “he ae nal I 

— ea pusilla, Britton sass Taylor on 


Schlechter ore Meo of 136 

Sch erm “ Disease of black 
ioonet » 206; 

Schultz, O. E., work of 364 


1901 | INDEX TO VOLUME XXXI 457 


Schumann’s “ ia Kakteen” 132, 
20 Lauterbach’s ‘Flora 
beg deutschen peel in der Siid- 


Sciadophyllom lena 13 

Sci timus, Fernald 213; 
polyps prophium i os “wie 
cus, prophyllum of 2 

PRT a Co m, new gems, Schumann and 
Lauterbach o 

Scotland, Smith on Be cabee of 136 

Scott, D. H., work of 136, 279 


as 

on ; immature 443; 

Munson on 444 

revels, Fitting on ¢ meguspore of 213 
Senecio Crawfordii, Britton on 364; 


we 


Crostinien Nn 213 
Sequoia secpesenene Pierce on 442 
Seward, A. C., work of 283 
Seymeria virgata 330 
Seymour, A. B., — 448 
Serratula scariosa 406 
Shear nelius A., personal 287 
Sida Hol wayi 330 
Seria or Rydberg o 
“te i ringhiens and Russell on 443 
Sirrine, F. A., work of 44 
paar Niagarense, Robinson on 
213; officina ee Robinson on 213 
Slosson, E. E., 443 
ith, Erwi pines 216; John B 
work of 135; John Donnell 109; Jared 
G., personal 287; } ary ; Robert, 


443; W. G., 

Societies: International Botanical Asso- 
ciation 367; for Plant Morphology and 
Physiology 143, 216; for Promotion of 
iach er Science, 438; Western 
Naturalists 72 

Solanum sppentcuatom 328; Dulca- 

mara, Ter 


Solida, ago , Greene on 213 
Solution, oe ya nutrient 367 
sommier, S., work of 364 
€ 


uller on chemotaxis of 281 
pores, (ers of 38 
seals: in leaves of Lee evergreens in 
r, Miyake o 


y of Minn 


gion , geotro weaneage 
Stenotopsis, new pena Rydberg on 


Station, yovsanil botanical, of the Univer- 
wee 


Secutean: new genus, Rydberg on 212 


ppt bani pt nigra, ‘Beeniator of 


Ste LF G:, 

Sticke eae ou 

Stimuli, influence of on germinati of 
spores 52; Nemec on conden of 133 

Stomata, Brown and Escombe on diffu- 
sion thro ough 281 

Ston f 443 

ashlee Eduard, ern 287 

Sturgis, William C., work of 444 
ylidi s on 20 

Succulents, Brenner on 211 

Suckers, Peirce on 442 

Sudwor th, George B., work of 210 


Swertia e on 212 
2 se pce m, new Ee Sea Schumann 


d Lauterbach o 
Keniun. poe on ees aes of 284 
ff is 
Tapirira, Brandegee on 135 : 
Taraxacum, Greene on 213 y. 


Taxonom ve) P25. 212, 250; 364 
Taylor, ap pe oe of 363 
6 


Terras, Jame eg work ce £33. FAT 
Thalictrum, Greene on 
snp Rydberg on 212 


Toumey, James W., personal 216 
Tournefortia ion a: 327 

Townsend, C. O., 

Travel, notes of 352, 423 

oe in Nebraska 133; North American 


Me 
Tre lease, geges 131 
oe 
Tripsa a ribner and Merrill on 213 


ay ristachya, Scribner and Merrill on #13. 
Triumfet 6 


U 


Umbilicaria, Minks on 
Underw pee . M., 366; Seon. 216; 


work of 2 na 
Uredo, pehain 335; Lippiae 335; Polym- 
niae 32 


Uredospores, germination of 44 __ 
Urocystis anemones, germination of 
spores 44 


458 


Uromyces, Aegopoginis 328; Caladii cal 
ca Pye padiinus, germination of s 
siae 326; dolichosporus 3273 


S 


Ustilago, ROR of spores oe 


V 


bee gp igi folia 333; montanifolia 
233 aerocephala 338; trilobata 


of 134 
a, Im ool. and Bot. Society 216 
Vigna "asobiophos 328 
ntata 


ce 334; Palmeri 


3325 
ike 

Villaresin  antiicnuits IIO 

Viscum album 39 

dias Schrenk, see Schrenk 


W 


Wager, H., work of 140 
Nena bacteriosis 
Ward) I> My 202:: 
Son 
Webber, H. J., work of 444 
Mkbatxe, Hollis, personal 287 
Weinzirl, John, work 
Wellesley College, 
Department of 288 


272 
Lester F., work of 


of 279 
gift to Botanical 


BOTANICAL GAZETTE 


| JUNE 


Wettstein, Richard von, personal 142 
Wheeler, W. A., work of 361 
Whi of 282 


- 133, 135, 211, 289 

Wies ner’s “ Die Rohstoffe a Pflanzen- 

reiches’”’ 70 
bier ox, E. M., see Chesnut ; work of 361, 
Wildeman and Durand's * Contributions 

ala flore du Congo” 70; “ Illustrations 

de la flore du Congo” 205, 359 
Williams, R. S., personal 448; Thomas 

ae , personal 143 

Voody ogee radicans 349 
“lieing . C., work of 278 
Wrightson, | Phi lip G. 141 
Wyoming, Nelson on cryptogams of 443 


x 


Xanthium 330 
Xe paren, new fete Schumann and 


Lau 
Kslocte penpidhertealiee 109 


¥ 
e, Y., persona 
ee eae ad paler of 419 
vs 


Zizia aurea, Bissell on 135 
Zygadenus Colabaduetists 3043 elegans 
394 


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[resacaae J 


Cincinnati 
Asheville 


an 
Jacksonville 
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heft agen en AND DINING 
RS BY DA 

nen sLaPinG foe i Ste aah 

MEnT 

6 TRAINS DAILY , BE 

pile THE OHIO RIVER 


oe 
W.H.M“DOEL pass.act. CHASH ROCKWELL 
PRES.& GENL MOR. ee CHICAGO. TRAFF.MGR. 


NEW MODEL 


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Redmond DEAL IN 


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Kerr & Co e | Investment 


BANKERS 


41 WALL ST., N.Y. Securities 


List of current offerings sent on application 
Transact a general banking business 


Receive deposits subject to draft. Dividends and 
interest collected and remitted. Act as Fiscal Agents 
for and negotiate and issue loans of railroads, street 
railways, gas companies, etc. Securities bought and 
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Issue Travelers’ 
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THROUGH PULLMAN SERVICE 
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F YOU ARE CONTEMPLATING A TRIP, ANY POR- 
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Xx ALTON, IT WILL PAY YOU TO WRITE TO THE UNDER- 
SIGNED FOR RATES, MAPS, TIME-T. ABLES, ETC. 


Geo. J. CHARLTON, 
pe Se PASSENGER AGENT, 


"MADE FROM THE BEAN 


Waukesha Hygeia 
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FROM IT IS MADE THE FAMOUS 


BORO-LITHIA WATER GINGER ALE 
AND WILD CHERRY PHOSPHATE 
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THE WAUKESHA WATER CO. 
132 N. Jefferson Street, Chicago 


Telephones - - Monroe 1166 and 1168 


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Health Policies. 


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Manufacturers and Mechanics, Contractors, and Owners of 
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Paid-up Cash Capital, . . $1,000,000.00 
ASSETS, . . . .... 40,001,030.00 


Liabilities, . . (6 20,387,003.25 
EXCESS, 3% per cert basis, 4,543,126.81 
Life Insurance in force, . . . © $109,019,851.00 
Returned to Policy holders, .° 42,643,384.92 


J. G. BATTERSON, — 
S. C. (DUNHAM, Vice President. H, J. MESSENGER, Act 
JOHN E. MORRIS, Secretary. E. V. PRESTON, aerate t af Agencies, 


= 


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TRUE COIN EELS a SHIN 
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GEONPIERCE: (O'R BUTEALO- BOSTON: REO SORKD FiconeR EADS Ger Wy JOR LD 


TIME SAVING SYSTEMS ae L=<Ly Sy LS * | 
-see r Our Engraving ¢ iy 


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ng our s of c e 
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ACCOUNTANTS 2 a to people of taste and refine- 
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AUDITORS — Our Prices Prices 


There will be no guess work, no erratic theory in the 
work we do, emily ingen is s keener today, = it costs Are as low as hone ad our 
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more to do business than fore. Os standard can be nam 


must have perfect cimaitbarten. 
DEVISERS OF BUSINESS SYSTEMS Wesend you finely engraved 
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may be desired the true condition of affairs in detail. 


We stand on our record a Mo vik re and initial work 


NUFACTURERS = eprcalty 
of a Leaf Binding and Filing Devices and Supplie: High class correspondence 
We have experience to sell to those who want pe pec §, papers. 
willing £0 pay for results. SAdsons our Audit Depart- Xe 
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A $. D. Childs 5 Zo. 
x 140-142 Monroe St. 


Established 1837 CHICAGO. 


| BAKER- -VAWTER COMPANY 


GAS AS WS 


| ATcMISON, na HOLYOKE. mass. GAPE DE Des DE DS DS 


Cée HARTFORD RUBBER WORKS CO..HARTFORD, Conn. #f 


Big Four Route 


FROM 
CHICAGO 
TO 


Indianapolis, Cincinnati, Louisville, 


the 
South and Southeast. 
THE SCENIC LINE TO 
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via the Picturesque 
CHESAPEAKE & OHIO R’Y, 


he short line to 
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W. J. LYNCH, GP, & TA, W, P, DEPPE, Ass’t GP. & T.A. 
CINCINNATI, O. 
J. C. TUCKER, G.N.A. 234 Clark St., CHICAGO 


DAN-AMERICAN | 
EXPOSITION 


IS THE SHORTEST LINE 
22 BVFFALO & 
KANSAS CITY, ST. LOUIS, CHICAGO 
_AND INTERMEDIATE POINTS, 


F 


0. 8. CRANE, Gen’! 


call on neare: 


Ticket Agent, or address 
Pass’r and Tick gent, ST. LOUIS. 


Pit; FUR ESOUEL 
ROUTE 


TO 


N. E. A. MEETING 


Detroit, Michigan 
JULY 8:40 JULY: 12 
AND THE 
Pan-American Exposition 
Buffalo, New York 


MAY to NOVEMBER, 1901 


CITY TICKET OFFICE: 
249 Clark Street, cor. Jackson Boulevard 


NO HCE 


The EXCELLENT 
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SERVICE 


From CHICAGO 
: To the EAST 


Att Trains Darry 


6 8:30a.m. | Buffalo, Albany, and New York. 
ro 10330 a.m. | New York and Boston Special. 
be 
16 | 2:00 p.m, New England Express. 
22 | 5:30p.m, Lake Shore Limited 
| 
| 
26 8:30p.m. | Cleveland. 
28 10:35p.m. | New York and Boston Express, 
| j 
ski. Si. 90 ae RR ERE anise NeR DE 
32 3:00am, | Buffalo and East, 


} | 
City ticket office, 180 Clark St. 
F,.M.BYRON,G.W.A., | A.J.SMITH,G.P.A., 
Chicago, Til. | Cleveland, 


THE PHOTOGRAPHIC 
TIMES Fer may 


AS one of the most sg a persia ere he A ek isi liter sop “a er fee ee 
re contained, besides other features of v nete amples rork of t 
cently deceased Ht. 'P. Robinson, ped ee “the tien grea ie oh pet oats 
Many of ‘these are of full page size, prin ited on heavy plate paper, and the Sac. ce in 
photogravure is by the same artist. This nortan should be on the shelves of all who have 
any interest in i saree as an art. 
The June Number will contain many features of exceptional interest. 
A photogravure fri ontispiece, and many articles written by photographic experts which will 
be found to be of great value. 
To introduce the magazine to a still larger circle of readers, we will 
send the Photographic Times for May and for June as well as one other num- 
ber, all for twenty cents in stamps —Three for the price of one. 


The Photographic Times ts a monthly. 
Subscription price, $2.00 


The SCOVILL & ADAMS COMPANY of NEW YORK 


3 and 5 West Nineteenth Street, New York 


“IT’S ALL IN THE LENS”’ 


KORONACAMERAS 


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A 


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MICROSCOPES ¢ 


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ctor of Papers on MICRO-CHEMICAL ANALYSIS 
y PROF. E, M. CHAMOT, Cornell Univ., now running 


> 


Journal of 


Applied Microscopy 


Laboratory Method i 
a ethods a“ 
m 
Vol. IV January, 1901 No. s | © 
LEADING SUBJECTS mm 
Moses C. dees 
. H. GAGE, Cornell University, ..........---+. 1108 2 & LOMB ¢ 
Fire in the V fiesdins College at Cornell. 
SU ONGR se ov ee Ss A OA wei a etree a 1111 OPTICAL 
Lab: — Faetogreshy- ry . ce) 
tereo Photo-Micrography, . - 111 
The New York Botanical Garden. fe) CO MPANY 
D. ai ee = DOUGAL, ee ne Fe Kea 1115 
Pretiminary Study Uv 
CLARA “LANGENBECK, Wells College,» + es eee ee 1119 inl 
Micro-Chemicat Analysis, X. Pot 
EM. CHAMOT, Cornell University 1121 Me 
Easy Method of Mounting and Preservi me $9 
VEN: CAAA MMe ph ciyd tte te a5 als ain't 1129 vn 
Current ~ bye ma ‘ Ss] 
HAMBER etsity of Chicago, 31 
Cytology, eee a steal ates Microscopic: ei oa. 
AGNES M. CLAYPOLE, Cornell University, ......... 1133 m 
Normal and Pathological. Histology. m 
RICHARD M. PEAR D., Harvard College,...4-. 1136 
General Physiology 
“ RAYMOND PEARL, Universi ity of Michi 1138 
Current Bacteriol ture. 


‘ Tentrenetss i! ‘esleyan University, ... 1-6 ese 202s 1141 
Notes on Recent eralogic: sa tee, 


. 
( 
( 
( 
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=> . > © @@ @2686¢ 42844686 66636466869 


ALPRED J. MOSES, LEA Mcl. LUQU! yER, ~ +. 11446 
Modical Notes, ©... tee ee eee te eee enter sesens 1145 
News and ee ee ee ae eee ee a 1 
Publication ees: — mit saree kiaacca Soy Rochester, N. Y, 
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NEW YORK CHICAGO 
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A practical treaties compiled from Board of Health reports 
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treat them, with illustrations showing methods of house- 
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HENRY B. PLATT, 36 PLATT ST., NEW YORK. 


Dining Cars 
Service a la Carte 


OSES are always on the tables, 
i ' St. Louis to all the i rtant 
and delicate china, glass and ee cod ia cae 
5] . ’ yest such as 
silver add to the pleasur a dint — 
P .° ef ST. PAUL 
well cooked and daintily served IRMERPOLS 


KANSAS CITY 
DENVER 


u are going to any of 

° © ese places or to the 
Alt the Season’s Delicacies pene kindly let me send you 
at Very Moderate Prices time tables and other informa- 


tion about our train service. 


P. S. Eustis, 
Gen’! Pass. Agt., C. B. & Q. R. R., 
CHICAGO, ILL. 


IN URIC ACID DIA: 

UFFALO LITHIA WATER {8 dst uit 
MATI SM, ETC. THIS 

WATER DISSOLVES URIC ACID AND PHOSPHATIC SEDIMENTS, ETC., ETC. 
John V. Shoemaker, M.D., LL.D., Professor of Matera. Medica and sae pt in the 

Meilco Chivurgtal College of Philadelphia, etc., in the New York Medical Journal, June, 22 99s 

a s doubly efficient in Rh ti d a I 
The BUFFALO LITHIA WATER diecaie ee tiie ne and Posphatic ceiiandats: 

as well as other products difficult of elimination, while at the same time it exerts a moderately stimu- 

lant effect upon ne renal ce lls, and thereby pt ita f 


ool 
o 
“ 
[on al 
> 
Specie 
Fs) 
ZS 
=? 
or 
- 
tb) 
| 
° 
< 
2 
° 
= 
ms 
wi 
Cc 
1X} 
SF 
ra 
=| 
rs) 
ee. 
~ 
Me) 
po 
ies 
be 
i 
° 
3 


The intense sag produce ip 

avoided by prompt elimination. Beste 9 although the speedy removal of Uric Acid an 
other products of faulty tissue change is of. conspicuous benefit, yet to PREVEN their formation 
is a ee still more ea nt. when it corrects those 
This service is me the Fi WATER « digestive failures 
which are esporsible for the chau of deleterious materials.”’ 


The late Hunter McGuire, M.D., LL. D., Formerly President and Professor of Clinical 
Surgery, aa College of Medicine, ” Richmond, Va., and Ex-President of the American Medical 
Association, saps: 

ss as an alkaline seeker’ is invaluable. In Uric Acid 
spain ie indeed in diseases generally dependent 

upon a Uric Acid Diathesis, it is a remedy of e ordinary potency. | have prescribed it in 
cases of arta npesny A Gout which had rested the orittiedy eae aS with wonderfully good Pea 

I have used it’a in my own case, being a great sufferer from t malady, an ve 

derived more benctit from it than from any other remedy.”’ 


Dr. P. B. Barringer, Professor of Physiology and Surgery, University of Virginia: 

‘In more than twenty years of practice | have used Lithia as am ae -uric acid eee ae ine 
and have tried itina great variety of forms, both in the NATURA ATERSa ABLETS. 
As cs result of this experience, | have no hesitation in stating ms me baie result 1 have airs 
n ing to in preventing uric ac tg its in the 
compare with BUFFALO LITHIA WATER 10 ody. My experience with it as a solvent 
af old existing pia (calculi) has poe i gi & limited, and | hesitate to compare it ae wee fs 
other forms to their disadvantage; 
first class of conditions above set forth | feel that + BUFFALO LITHIA WATER & ALONE.” 
Dr. Thomas H. gga of Paris (Formerly of Baltimore), Suggestor cq Lithia as a Sol 

vent jor Uric Acid, say. 

** Nothing I could say wou ia z add 1 BUFFALO I have frequently 

rs 2 LITHIA WATER. scl ii wit, coo 


results in DIATHESIS, RHEUMATISM, and GOUT, and with. this object I ee 
ordered it to Europe. Lithia is in no form so valuable as where it exists in the carbona 3 
form in which nature’s mode of so vsti n and division in 


it is found in LITHIA WATER, water which has passed orca xk Lepidolite 
and Spondumne Mineral formations.”’ 


Dr. J. Rink Mallet, a. - Chemistry, University of Virginia.. Extract from report of analysis 
(teat under the action of BUEFALO 
ars under the acti te! WATER Spring No. 2. 
** It seems on the poor rable that the action of the Mon is PRIMARILY and MAINLY 
wipe upon UR ACID A D THE URATES, but when these constituents occur along 
with and as cementin age ra to S Pubepeanic or Oxa Oxalic Calculus materials, the latter may be 
so jaa g and broken (pili as to disintegrate the Calculus as a whole in these cases, also thus 
admitting of Urethral discharg 
James Sam Cabell, M. na) A.M., LL. D., Formerty Professor of Aiea and Surgery in t 
‘edical Department 6 the University of Virginia, and President of the N: nal Board of Heaith, dapat 
rT in. Uric Ac Acid og ars ig a well-known thera- 
WATER ea iapaes t should be recognized by the 


BUFFALO LITHIA 
profession as an article of Materia Medica.”’ 
_ BUFFALO LITHIA WATER is for sale by Grocers and Druggists generally. 


"TESTIMONIALS WHICH DEFY ALL IMPUTATION OR QUESTIONS SENT TO ANY ADDRESS 


IETOR. BUFFALO LITHIA SPRINGS . VIRGINIA 


BAKERS 
BREAKFAST 


< ie eit CHOCOLE. 
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‘Walter ee Mark 
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( Q 
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? 

> § 

5 

March 16, rgor. March 16, 1gor. r 

W EBEI R ae MPANY, WEBER COMPANY, 

" New y York. § 

5 onedd Th ) 

of the Ww eber pianos used ( 

by me in New York, San § 

ane ramets and el = here ) 

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and S have found them ( 

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4 Sincerely yours, 

keri Wkece “Epovarp DE RE 
\dieiletentbipcecaeiss 


The Weber Pianos used by the artists of the 
éeTa 


at the WEBER WAREROOMS, 108 Fifth : enue, 
corner Sixteenth Street, New York, at 


SPECIAL PRICES 


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Write promptly for ra and full particulars, 
mentioning this magazine.